Comprehensive Crop Science( 6049) Hand out

Compiled by: Shamuyarira.M [Bsc Agric Science,2020]

Mvurwi High School Publisher

1
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Contents
PLANT CLASSIFICATION ...................................................................................................................................... 5
CYTOLOGY(Cells) .............................................................................................................................................. 11
The cell cycle .................................................................................................................................................... 19
CELL DIVISION .................................................................................................................................................. 20
Stages of mitosis .............................................................................................................................................. 20
Meiosis ............................................................................................................................................................. 22
MINERAL COMPOSITION OF SOIL PARTICLES .................................................................................................. 28
Soil components............................................................................................................................................... 29
Soil water ......................................................................................................................................................... 30
PHYSICAL PROPERTIES OF THE SOIL ................................................................................................................. 32
Soil texture ....................................................................................................................................................... 32
Soil structure .................................................................................................................................................... 33
SOIL COLOR ...................................................................................................................................................... 35
Topography/relief ............................................................................................................................................ 37
SOIL AIR ............................................................................................................................................................ 38
Importance of Soil Aeration ............................................................................................................................. 39
SOIL BULK DENSITY, PARTICLE DENSITY AND POROSITY ................................................................................. 40
ORGANIC MATTER ........................................................................................................................................... 42
Factors affecting soil organic matter ............................................................................................................... 60
Composition of organic residues: .................................................................................................................... 60
Humus .............................................................................................................................................................. 64
Importance of manure ..................................................................................................................................... 64
Carbon Nitrogen Ratio ..................................................................................................................................... 65
PLANT NUTRIENTS ........................................................................................................................................... 66
BIOLOGICAL NITROGEN FIXATION ................................................................................................................... 69
PLANT GROWTH ............................................................................................................................................... 82
FLOWER AND FRUIT DEVELOPMENT ............................................................................................................... 84
VERNALISATION ............................................................................................................................................... 84
PHOTOPERIODISM ........................................................................................................................................... 87
PLANT GROWTH HORMONES ......................................................................................................................... 89
GAMETE FORMATION .................................................................................................................................... 100
POLLEN DEVELOPMENT ................................................................................................................................. 101
OVULE DEVELOPMENT ................................................................................................................................... 103
POLLINATION ................................................................................................................................................. 104
Pollination process ......................................................................................................................................... 105
DOUBLE FERTILISATION ................................................................................................................................. 106

2
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

SEEDS ............................................................................................................................................................. 107

TYPES OF GERMINATION ............................................................................................................................... 111
DORMANCY .................................................................................................................................................... 117
BREAKING SEED DORMANCY ......................................................................................................................... 118
PLANT – WATER RELATIONS .......................................................................................................................... 119
PHOTOSYNTHESIS .......................................................................................................................................... 128
C3 CARBON FIXATION .................................................................................................................................... 132
C4 carbon fixation ........................................................................................................................................... 132
C4 PATHWAY .................................................................................................................................................. 133
CRASSULACEAN ACID METABOLISM .............................................................................................................. 135
Comparison with C4 metabolism ................................................................................................................... 136
RESPIRATION .................................................................................................................................................. 138
ATP ................................................................................................................................................................. 142
PRINCIPLES OF CROP BREEDING AND BIOTECHNOLOGY ............................................................................... 143
CHROMOSOME .............................................................................................................................................. 144
THE STRUCTURE OF DNA ............................................................................................................................... 147
DNA REPLICATION .......................................................................................................................................... 148
PROTEIN SYNTHESIS ....................................................................................................................................... 151
GENE EXPRESSION ......................................................................................................................................... 152
EFFECTS OF ENVIRONMENT ON GENE EXPRESSION ...................................................................................... 154
MUTATIONS ................................................................................................................................................... 155
MENDELIAN LAWS OF INHERITANCE ............................................................................................................. 157
BREEDING....................................................................................................................................................... 160
Single hybrid cultivar (AB) .............................................................................................................................. 166
Double hybrid cross ....................................................................................................................................... 166
GENETIC ENGINEERING .................................................................................................................................. 167
WEEDS AND WEED CONTROL ........................................................................................................................ 170
CROP WEED COMPETITION(allelospoly): ....................................................................................................... 177
METHODS OF WEED CONTROL ...................................................................................................................... 181
CROP PESTS .................................................................................................................................................... 184
INTEGRATED PEST MANAGEMENT (IPM) ...................................................................................................... 193
MANAGEMENT TACTICS IN IPM..................................................................................................................... 194
CROP DISEASES .............................................................................................................................................. 196
Transmission of Plant Diseases ...................................................................................................................... 198
DISEASES CONTROL METHODS ...................................................................................................................... 202
Sprayer Calibration ........................................................................................................................................ 204
CROP PRODUCTION ....................................................................................................................................... 205
3
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

AGRONOMIC PRINCIPLES ............................................................................................................................... 205

AGRO-ECOLOGICAL ZONES OF ZIMBABWE .................................................................................................... 205
TILLAGE .......................................................................................................................................................... 209
PLANTING ....................................................................................................................................................... 213
ROTATIONS .................................................................................................................................................... 217
MAIZE ............................................................................................................................................................. 220
SOYABEAN PRODUCTION ............................................................................................................................... 224
CONSERVATION FARMING:PRINCIPLES AND PRACTICES ............................................................................... 228
PRINCIPAL CONSTRAINTS TO EXPANDED CROP PRODUCTION ...................................................................... 228
COLLOIDS AND CLAY MINERALS ...................................................................................................................... 42
Causes of negative charges in colloids ............................................................................................................. 43
Cation Exchange Capacity ................................................................................................................................ 43
Clay minerals .................................................................................................................................................... 45
Nutrient Uptake Mechanism ........................................................................................................................... 47
SOIL ACIDITY, SALINITY AND ALKALINITY ......................................................................................................... 48
Soil sampling .................................................................................................................................................... 55
Principles of soil sampling ................................................................................................................................ 56
Methods of soil sampling ................................................................................................................................. 56
CROP MANAGEMENT ..................................................................................................................................... 234
Maturity Indices and Harvest Indices............................................................................................................. 234

4
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 1

PLANT CLASSIFICATION

By the end of this topic, learners must be able to:

 classify plants
Defination
The arrangement of plants in taxonomic groups according to their observed similarities (including at
least kingdom and division in plants, and class, order, family, genus, and species).
Botany.
Botany, also called plant science(s), plant biology or phytology, is the science of plant life and a
branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this
field. The term "botany" comes from the Ancient Greek word βοτάνη (botanē) meaning "pasture",
"grass", or "fodder"; βοτάνη is in turn derived from βόσκειν (boskein), "to feed" or "to graze".
Brief History
Modern botany traces its roots back to Ancient Greece specifically to Theophrastus (c. 371–287 BC),
a student of Aristotle who invented and described many of its principles and is widely regarded in
the scientific community as the "Father of Botany". His major works, Enquiry into Plants and On the
Causes of Plants, constitute the most important contributions to botanical science until the Middle
Ages, almost seventeen centuries later.
During the 18th century, systems of plant identification were developed comparable to dichotomous
keys, where unidentified plants are placed into taxonomic groups (e.g. family, genus and species) by
making a series of choices between pairs of characters. The choice and sequence of the characters may
be artificial in keys designed purely for identification (diagnostic keys) or more closely related to the
natural or phyletic order of the taxa in synoptic keys.
Taxonomy
taxonomy (from Ancient Greek τάξις (taxis), meaning 'arrangement', and -νομία (-nomia), meaning
'method') is the science of defining and naming groups of biological organisms on the basis of shared
characteristics. Organisms are grouped together into taxa (singular: taxon) and these groups are given
a taxonomic rank; groups of a given rank can be aggregated to form a super-group of higher rank, thus
creating a taxonomic hierarchy. The principal ranks in modern use
are domain, kingdom, phylum (division is sometimes used in botany in place of
phylum), class, order, family, genus, and species. The Swedish botanist Carl Linnaeus is regarded as
the father of taxonomy, as he developed a system known as Linnaean taxonomy for categorizing
organisms and binomial nomenclature for naming organisms.
With the advent of such fields of study as phylogenetic, cladistics, and systematics, the Linnaean
system has progressed to a system of modern biological classification based on
the evolutionary relationships between organisms, both living and extinct.
Phylogenetic
In biology, phylogenetics /ˌfaɪloʊdʒəˈnɛtɪks, -lə-/ (Greek: φυλή, φῦλον – phylé, phylon = tribe, clan,
race + γενετικός – genetikós = origin, source, birth) is the study of the evolutionary history and
relationships among individuals or groups of organisms (e.g. species, or populations).
Cladistics (/kləˈdɪstɪks/, from Greek κλάδος, cládos, "branch")[1] is an approach to biological
classification in which organisms are categorized in groups ("clades") based on the most recent
common ancestor.
Biological systematics is the study of the diversification of living forms, both past and present, and
the relationships among living things through time.
5
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

How is the hierarchy formed?

 The hierarchy starts from the species (basic unit of classification).
 After the species, closely related species are grouped into a category (taxon) called Genus and
members of a genus have much in common morphologically. The number of plants in a genus
lends itself well to studies of biochemical, cytological, ecological and genetic relationships.
 Closely related genera form a Family-this grouping is frequently encountered in taxonomic studies
because it is small enough to allow relatively easy study of natural relationships among its
members. This is the most used category of the major taxa
 From Family there is Order, Class, Division and up to the Kingdom.
 Taxa from Kingdom to family are the major taxa while those below are minor taxa
VARIETY
 When one or more of the populations of a species is significantly different from the remaining
members it is given a varietal status. E.g. purple and yellow granadillas are varieties. Cabbage,
cauliflower and Brussels sprouts are varieties of the same species – they exist in nature as such.
 This is different from cultivar
Cultivar
 Short for cultivated variety.
This is a named group of plants within a cultivated species that is distinguishable by a character or group of
characters and that maintain its identity when propagated either sexually or asexually.
 This does not exist in nature but maintained by cultivation thus a result of cultivation, selection and breeding
and not by natural means
Scientific naming of plants
 Scientific names are Latin
 Genus and species are used to name plants – system is called binomial nomenclature and was first
used by a Swedish botanist Carl Linnaeus.
 The genus name should always be a singular noun and written with a capital initial letter.
 The specific scientific name is a combination of the generic name and the specific epithet
 Specific epithets themselves are not names therefore not used on their own to refer to plants. These
should always be written in small letters
 The full scientific name should be italised or underlined e.g. Citrussinensis or Citrus sinensis
 Names of taxa below species are named by combinations of the specific name and the appropriate
intraspecific epithet e.g. Brassica oleracea var. capitata for common cabbage
 Sometimes names upper with an abbreviation or a letter at the end-the abbreviation or letter stands
for the name of the author of that name e.g. Phaseolus vulgaris L.
IMPORTANCE OF CLASSIFICATION
Importance of classifying the Crop Plants:
1. To get acquainted with crops.
2. To understand the requirement of soil & water different crops.
3. To know adaptability of crops.
4. To know the growing habit of crops.
5. To understand climatic requirement of different crops.
6. To know the economic produce of the crop plant & its use.
7. To know the growing season of the crop
8. Overall to know the actual condition required to the cultivation of plant.

Crops plants are classified based on three criteria which are;

 Botanical classification – Based on generic and specific names

6
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Agricultural classification – Based on the uses or important of crops to man and animals
 Classification based on life cycle – Based on numbers of year or season from germination to
harvesting or death.
General Classification of Plants
1. By Growth habit:
Succulent plants – herbaceous or herbs (succulent seed plants possessing self-supporting stems)
Vine – a climbing or trailing herbaceous plant (Liana – a climbing or trailing woody plant)
Trees – having a single central axis
Shrub – having several more or less upright stems
2. By Leaf drops:
Deciduous – no living leaves during dormant (winter) season (apple)
Evergreen – retaining functional leaves throughout the year (spruce)
3. By Life span:
Annuals – plants that normally complete their life cycle during a single growing season (lettuce,
spinach, marigold)
Biennials – plant that normally completes its life cycle during a period of two growing seasons
(celery, carrot)
Ephemerals: They are crops that complete their life cycle once in three or four months and can
undergo two or three life cycles in a year e.g. tomato
Perennials – plants that grow year after year, often taking many years to mature.
These crops are classified in to two groups.
i) Herbaceous:
Herbaceous perennials are those with more or less soft succulent stems. In Temperate climates the tips
die off after season’s growth but root remains alive and produce new stem and tops on favorable
conditions. In other words their tips are annual while ground parts are perennials lie many years and
are classified as: a) Trees b) Shrubs c) Vines according to their habit of growth.
a) Trees: Trees are upright in habit and stems take the form of central axis e.g. Mango, guava,
Mandarins etc.
b) Shrubs: Shrubs have no main trunk but a number of erect or semi erect stems are seen but do not
forms the main frame work e.g. Hibiscus, Rose etc.
c) Vine: Both woody and herbaceous have stems which are flexible and not in position to keep their
branches and leaves erect. They either spread on the ground or require some support whether alive or
manmade e.g. Grape vines, Passion fruit etc.
4. By Temperature tolerance:
Tender plant – damaged or killed by low temperature
Hardy plant – withstands winter low temperatures
Wood hardy – a whole plant is winter hardy
Flower-bud hardiness – ability of flower buds to survive low winter temperatures (peach, ginkgo tree)
5. By Temperature requirements:
Cool-season crop – prefers cool temperatures (peas, lettuce)
Warm-season crop – prefers warm temperatures (tomato, pepper)
6. By Habitat or Site preference:
Xerophytes – prefers dry sites
Shade plants – prefers low light intensity
Acid loving – prefers low pH soils
Halophyte – prefers salty soils
7. Classification based on climate:
Based on temperature requirements and response to different climatic conditions, horticultural crops
have been classified in to three main groups and these are:
1. Temperate

7
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Temperate plants are commonly found in cold regions enjoying a mild and temperate climate. These
plants endure cold and go to rest or dormancy by shedding of all their leaves during winter e.g. Apple,
Plums cherry and almond etc.
2. Tropical
Tropical plants are those which do not tolerate severe cold but can tolerate warm temperatures of
about 37. Those plants need strong sunshine, warms, humidity and a very mild winter. They cannot
stand far against frost e.g. Papaya, Banana and Pineapple.
3. Sub -Tropical:
Sub -tropical plants like Orange, Litchi, Fig, Mango and cashew nut are intermediate in character.
They need warmth and humidity and can tolerate mild winters.
8. Use/Agronomic classification:
Grain crops: may be cereals as millets cereals are the cultivated grasses grown for their edible starchy
grains. The larger grain used as staple food is cereals. E.g. rice, wheat, maize, barley, and millets are
the small grained cereals which are of minor importance as food.
Pulse/legume crops: seeds of leguminous crops plant used as food. E.g. soybean, pea, cowpea etc.
Oil seeds crops: crop seeds are rich in fatty acids, are used to extract vegetable oil to meet various
requirements. E.g. Groundnut, Mustard, Sunflower, linseed etc.
Forage Crop: It refers to vegetative matter fresh as preserved utilized as food for
animals. Crop cultivated & used for fickler, hay, silage. Ex- sorghum, elephant grass, guinea grass, &
others etc.
Fibre crops: crown for fibre yield. Fibre may be obtained from seed. E.g. Cotton, sun hemp.
Roots crops: Roots are the economic produce in root crop. E.g. sweet, potato, sugar beet, carrot,
turnip etc.
Tuber crop: crop whose edible portion is not a root but a short thickened underground stem. E.g.
Potato, yam.
Sugar crops: the two important crops are sugarcane and sugar beet cultivated for production for
sugar.
Starch crops: grown for the production of starch. E.g. tapioca, potato, sweet potato.
Dreg crop: used for preparation for medicines. E.g. tobacco, mint, pyrethrum.
Spices & condiments/spices crops: crop plants as their products are used to flavour taste and
sometime colour the fresh preserved food. E.g. ginger, garlic, chilli, cumin onion, coriander,
cardamom, pepper, turmeric etc.
Vegetables crops: may be leafy as fruity vegetables. E.g. tomato.
Green manure crop: grown and incorporated into soil to increase fertility of soil. E.g. sun hemp.
Medicinal & aromatic crops: Medicinal plants includes cinchona, opium poppy, senna, belladonna,
iycorice and aromatic plants such as lemon grass, citronella grass, Japanese mint, peppermint, rose
geranicem, jasmine, henna etc.
9. Classification based on root system:
Tap root system: The main root goes deep into the soil. E.g. Grape, Cotton etc.
Adventitious/Fibre rooted: The crops whose roots are fibrous shallow & spreading into the soil. E.g.
Cereal crops, wheat, rice etc.
10. Classification based on economic importance:
Cash crop: Grown for earning money. E.g. Sugarcane, cotton.
Food crops: Grown for raising food grain for the population and & fodder for cattle. E.g. wheat, rice
etc.
11. Classification based on No. of cotyledons:
Monocots or monocotyledons: Having one cotyledon in the seed. E.g. all cereals & Millets.
Dicots or dicotyledonous: Crops having two cotyledons in the seed. E.g. all legumes & pulses.
12. Classification based on photosynthesis’ (Reduction of CO2/Dark reaction):

8
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

C3 Plants: Photo respiration is high in these plants C3 Plants have lower water use efficiency. The
initial product of C assimilation in the three ‘C’ compounds. The enzyme involved in the primary
carboxylation is ribulose-1,-Biophospate carboxylose. E.g. Rice, soybeans, wheat, barley cottons,
potato.
C4 plants: The primary product of C fixation is four carbon compounds which may be malice acid or
acerbic acid. The enzymes responsible for carboxylation are phosphoenol Pyruvic acid carboxylose
which has high affinity for CO2 and capable of assimilation CO2 event at lower concentration,
photorespiration is negligible. Photosynthetic rates are higher in C4 than C3 plants for the same
amount of stomatal opening. These are said to be drought resistant & they are able to grow better even
under moisture stress. C4 plants translate photosynthates rapidly. E.g. Sorghum, Maize, napter grass,
sesame etc.
Cam plants: (Cassulacean acid metabolism plants) the stomata open at night and large amount of
CO2 is fixed as a malice acid which is stored in vacuoles. During day stomata are closed. There is no
possibility of CO2 entry. CO2 which is stored as malice acid is broken down & released as CO2. In
these plants there is negligible transpiration. C4 & cam plant have high water use efficiency. These are
highly drought resistant. E.g. Pineapple, sisal & agave.
13. Classification based on length of photoperiod required for floral initiation:
Most plants are influenced by relative length of the day & night, especially for floral initiation, the
effect on plant is known as photoperiodism depending on the length of photoperiod required for floral
ignition, plants are classified as:
Short-day plants: Flower initiation takes place when days are short less than ten hours. E.g. rice, etc.
Long day’s plants: require long days are more than ten hours for floral ignition. E.g. Wheat, Barley
Day neutral plants: Photoperiod does not have much influence for phase change for these plants. E.g.
Cotton, sunflower. The rate of the flowering initiation depends on how short or long is photoperiod.
Shorter days, more rapid initiation of flowering in short days plants. Longer the days more rapid are
the initiation of flowering in long days plants.
14. Botanical classification
In scientific classification, plants are classified in a hierarchical format called taxonomic hierarchy.
In scientific classification, plants are classified in a hierarchical format called taxonomic hierarchy
shown below:
KINGDOM A kingdom is the highest rank of biological classification. All plants belong to the
kingdom Plantae. The total number of kingdoms varies from between five and eight.
DIVISION A division is the first major rank below the kingdom. The term "division" is not
interchangeable with "phylum." "Division" refers to ranking in botany while "phylum" is
for zoology." The kingdom Plantae has twelve recognized divisions. Divisions are
generally based on morphological differences.
CLASS Class follows the phylum and precedes the order. Classes are determined by professional
taxonomists. There is often no consensus in the scientific community on which plants
belong to which classes or what group constitutes a biological class.
ORDER An order follows super order and precedes suborder.
FAMILY A family follows the super family and precedes subfamily.
TRIBE The tribe follows the super tribe and precedes the sub tribe.
GENUS The genus follows the sub tribe and precedes the species name. The genus is the first part
of the binomial name.
SPECIES The species follows the super species and precedes the subspecies. The species name is
the second part of the binomial name.
CHECK YOURSELF
1. What is the definition of a species?
A. A group of individuals who are able to mate
9
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

B. A group of individuals who belong to the same phylum

C. A group of individuals who have similar characteristics
D. A group of individuals who have similar characteristics and can produce fertile offspring
2. Which order is correct for the order of taxonomy?
A. Kingdom Phylum Class Order Family Genus Species
B. Kingdom Animals Mammals Fish Bacteria
C. Kingdom Class Genus Species Family Order Phylum
D. Species Genus Family Order Class Phylum Kingdom
3. Who is Carl Linnaeus?
A. A scientist that created the 'binomial nomenclature'
B. A scientist that was from Sweden
C. A scientist that is famous for his work in taxonomy
D. All of the above
4. What does binomial nomenclature mean?
A. Two-naming system
B. Singular name
C. Genus Species
D. An organism's scientific name
5. What does classification mean?
A. Grouping similar organisms together
B. Observing living organisms under a microscope
C. Randomly grouping living organisms in order to investigate their characteristics
D. Creating a classroom display of the results of an investigation
6. The plant kingdom is divided into two important groups. What are they?
A. Grasses and trees
B. Green plants and non-green plants
C. Flowering and non-flowering plants
D. Coniferous and deciduous plants
7. Glycine max is the genus and species of what fruit?
A. Maize
B. Beans
C. Soyabeans
D. cowpeas
8. Which two taxonomic levels are used in the binomial system?
A. Genus and phylum
B. Family and species
C. Genus and species
D. Phylum and class
9. What is the scientific name for wheat
A. Triticum eastivum
B. Tagetes munuta
C. Triticum sativum
D. Hypogea arachis
10. Gymnosperms are
A. Plants that produces flowers
B. Plants that produces spores
C. Plants that bears fruits
D. Plants that lacks a true root
SECTION B
11. Explain how plants are classified.[15]

10
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 2

CYTOLOGY(Cells)

 draw a plant cell

 label parts of a plant cell
 explain functions of cell parts
 identify cell organelles
 outline functions of cell organelles
 Explain the relationship between organelles.

11
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

PLASMA MEMBRANE/ CELL MEMBRANE

CELL MEMBRANE

Membrane lipid bilayer is unit membrane that surrounds prokaryotic and eukaryotic cells. In
addition, all bio membranes form closed structures, separating lumen on inside from outside, and are
based on a similar bilayer structure. They control movement of molecules between inside and outside
of a cell and into and out of organelles of eukaryotic cells. In accord with importance of internal
membranes to cell function, total surface area of these membranes is roughly tenfold as great as that of
plasma membrane

Basic Structural Organization of Membrane Lipid Bilayer

Fluid mosaic model of S. J. Singer and Garth Nicolson 1972 best describes structure of membranes
bilayer.

Lipids form a bilayer in which nonpolar regions of lipid molecules in each layer face core of bilayer
and their polar head groups face outward, interacting with aqueous phase on either side.

Proteins are embedded in this bilayer sheet, held by hydrophobic interactions between membrane
lipids and hydrophobic domains in the proteins. Some proteins protrude from only one side of
membrane; others have domains exposed on both sides.

The orientation of proteins in bilayer is asymmetric, giving membrane “sidedness”: protein domains
exposed on one side of bilayer are different from those exposed on other side, reflecting functional
asymmetry.

The individual lipid and protein units in a membrane form a fluid mosaic with a pattern that, unlike a
mosaic of ceramic tile and mortar, is free to change constantly.

The membrane mosaic is fluid because most of the interactions among its components are
noncovalent, leaving individual lipid and protein molecules free to move laterally in the plane of the
membrane.

12
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2. CYTOPLASM

Structure - The jelly-like substance composed of mainly water and found between the cell membrane
and nucleus. The cytoplasm makes up most of the "body" of a cell and is constantly streaming.

Function - Organelles are found here and substances like salts may be dissolved in the cytoplasm.

3. NUCLEUS

Structure - The largest organelle in the cell. It is dark and round, and is surrounded by a double
membrane called the nuclear envelope/membrane. In spots the nuclear envelope fuses to form pores
which are selectively permeable
─ Spherical structure 10-20 micrometres in diameter
─ Surrounded by a double membrane called nuclear envelope
─ Contains nucleoplasm
─ Nuclear envelope compartmentalises chemical reactions taking place in the nucleus
─ Nucleoplasm contains chromosomes and nucleoli
─ Chromosomes contain DNA attached to proteins (histones)
─ Nucleoplasm also contains RNA (3 types of RNA)
─ Nucleus functions to control the synthesis of proteins
─ Controls the cell’s activities
─ Divide at the start of cell division, ensuring that daughter cells have exact copies of the cell’s
genetic material
─ To assemble ribosomes
THE FOLLOWING ORGANELLES ARE FOUND IN BOTH PLANT AND ANIMAL CELLS.

1. "ER" OR ENDOPLASMIC RETICULUM

The Endoplasmic Reticulum is a network of membranous canals filled with fluid. They carry
materials throughout the cell. The ER is the "transport system" of the cell.
There are two types of ER: rough ER and smooth ER.
a) Rough endoplasmic reticulum
─ Have ribosomes on them.
─ Their role is to manufacture proteins.
─ Rough endoplasmic reticulum is abundant in cell that either secrete proteins or that are growing
rapidly.
b) Smooth endoplasmic reticulum
─ Lacks ribosomes on its surface.
─ Abundant in cells that secrete steroids or lipid substances
Functions of endoplasmic reticulum
─ To provide area for biochemical reactions.
─ To act as a pathway for the transport and exchange of material.
─ To manufacture proteins e.g. enzymes
─ To manufacture lipids and steroids.
─ To collect and store any synthesized material.
─ To form a structural skeleton for maintaining cellular shape

13
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2. RIBOSOMES

Ribosomes are small particles which are found individually in the cytoplasm and also line the
membranes of the rough endoplasmic reticulum. Very small organelles consisting of a large subunit
and a small subunit.
─ They are made of roughly equal amounts of protein and RNA.
─ There are two types i.e. 70s and 80s ribosomes.
─ They are responsible for protein synthesis.
─ They are either bound to ER or lie free in the cytoplasm.
─ They form polysomes i.e. collection of ribosomes strung along messenger RNA

3. GOLGI BODY / APPARATUS

-Composed of a series of parallel membranes that are flattened fluid spaces.

─ The cristae are slightly curved the entire structure appear concave.
─ Functions of Golgi apparatus:
 Manufacture of glycoproteins which are required for secretions.
 Production of secretory enzymes.
 Production of carbohydrates e.g. those involved in the manufacture of new cell walls.
 Transport and storage of lipids.
 Formation of lysosomes.

4. LYSOSOMES

Lysosomes are small sac-like structures surrounded by a single membrane and containing strong
digestive enzymes which when released can break down worn out organelles or food. The lysosome
is also known as a suicide sac.
─ Functions of Lysosomes:
 To contain enzymes capable of digesting a wide variety of substances.
 To digest cytoplasmic organelles.
 To act as suicide bags which help to rapidly digest entire cells that are old.

5. MITOCHONDRIA

 0.5–1.0 μm, diameter / width ;

 double membrane ;
 inner membrane folded / cristae ;
 hold, stalked particles / ATP synthase / ATP synthetase ;
 site of ETC ;
 ref. H+ and intermembrane space ;
 ATP production ;
 oxidative phosphorylation / chemiosmosis ;
 matrix is site of, link reaction / Krebs cycle ;
 enzymes in matrix ;
 70S ribosomes ;
 (mitochondrial) DNA ;
 They are envelope bound and the inner membrane folds to form cristae.
 It consists of a matrix with few ribosomes, a circular DNA molecule and phosphate granules.

14
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 In aerobic respiration, cristae are the sites for oxidative phosphorylation and electron transport
chain.
 Matrix is the site for Krebs’s cycle of enzymes.

6. VACUOLES

Vacuoles are fluid filled organelles enclosed by a membrane. They can store materials such as food,
water, sugar, minerals and waste products.

7. CENTRIOLES
 Found as a pair near the nucleus.
 They are made up of bundles of tubules.
 They pull apart during cell division to produce a spindle made of microtubules which are involved
in chromosome movement.

ORGANELLES AND OTHER FEATURES FOUND ONLY IN PLANT CELLS:

1. CELL WALL

The cell wall is a rigid organelle composed of cellulose and lying just outside the cell membrane. The
cell wall gives the plant cell its box-like shape. It also protects the cell. The cell wall contains pores
which allow materials to pass to and from the cell membrane.

2. PLASTIDS

Plastids are double membrane bound organelles. It is in plastids that plants make and store food.
Plastids are found in the cytoplasm and there are two main types:

Leucoplasts - colourless organelles which store starch or other plant nutrients. (Example - starch
stored in a potato)

Chromoplasts - contain different coloured pigments. The most important type of Chromoplasts is
the chloroplast, which contains the green pigment chlorophyll. This is important in the process of
photosynthesis.

3. CENTRAL VACUOLE

The central vacuole is a large fluid-filled vacuole found in plants.

15
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Structure

Chloroplasts found in higher plants are generally biconvex or planoconvex shaped. In different plants
chloroplasts have different shapes, they vary from spheroid, filamentous saucer-shaped,
discoid or ovoid shaped.

They are vesicular and have a colourless centre. Some chloroplasts are in shape of club, they have a
thin middle zone and the ends are filled with chlorophyll. In algae a single huge chloroplast is seen
that appears as a network, a spiral band or a stellate plate.

The size of the chloroplast also varies from species to species and it is constant for a given cell type.
In higher plants, the average size of chloroplast is 4-6 µ in diameter and 1-3 µ in thickness.
Function
The chloroplasts are double membrane bound organelles and are the site of photosynthesis. The
chloroplasts have a system of three membranes: the outer membrane, the inner membrane and the
thylakoid system. The outer and the inner membrane of the chloroplast enclose a semi-gel-like fluid
known as the stroma. This stroma makes up much of the volume of the chloroplast, the thylakoids
system floats in the stroma.
Outer membrane - It is a semi-porous membrane and is permeable to small molecules and ions,
which diffuses easily. The outer membrane is not permeable to larger proteins.

Intermembrane Space - It is usually a thin intermembrane space about 10-20 nanometres and it is
present between the outer and the inner membrane of the chloroplast.

Inner membrane - The inner membrane of the chloroplast forms a border to the stroma. It regulates
passage of materials in and out of the chloroplast. In addition of regulation activity, the fatty acids,
lipids and carotenoids are synthesized in the inner chloroplast membrane.
Stroma
Stroma is an alkaline, aqueous fluid which is protein rich and is present within the inner membrane of
the chloroplast. The space outside the thylakoid space is called the stroma. The chloroplast DNA
chloroplast ribosome’s and the thylakoid system, starch granules and many proteins are found floating
around the stroma.
Thylakoid System
 The thylakoid system is suspended in the stroma. The thylakoid system is a collection of
membranous sacks called thylakoids. The chlorophyll is found in the thylakoids and is the sight for
the process of light reactions of photosynthesis to happen. The thylakoids are arranged in stacks
known as grana.
 Each granum contains around 10-20 thylakoids.
 Thylakoids are interconnected small sacks, the membranes of these thylakoids is the site for the
light reactions of the photosynthesis to take place. The word 'thylakoid' is derived from the Greek
word "thylakos" which means 'sack'.
 Important protein complexes which carry out light reaction of photosynthesis are embedded in the
membranes of the thylakoids. The Photosystem I and the Photosystem II are complexes that
harvest light with chlorophyll and carotenoids; they absorb the light energy and use it to energize
the electrons.
 Thylakoids are of two types - granal thylakoids and stromal thylakoids. Granal thylakoids are
arranged in the grana are pancake shaped circular discs, which are about 300-600 nanometres in
diameter. The stromal thylakoids are in contact with the stroma and are in the form of helicoid
sheets.
-The granal thylakoids contain only photosystem II protein complex, this allows them to stack
tightly and form many granal layers with granal membrane. This structure increases stability and
16
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

surface area for the capture of light.

-The photosystem I and ATP synthase protein complexes are present in the stroma. These protein
complexes act as spacers between the sheets of stromal thylakoids.
Functions of chloroplast:

 In plants all the cells participate in plant immune response as they lack specialized immune cells.
The chloroplasts with the nucleus and cell membrane and ER are the key organelles of pathogen
defence.
 The most important function of chloroplast is to make food by the process of photosynthesis. Food
is prepared in the form of sugars. During the process of photosynthesis sugar and oxygen are made
using light energy, water, and carbon dioxide.
 Light reactions take place on the membranes of the thylakoids.
 Chloroplasts, like the mitochondria use the potential energy of the H+ ions or the hydrogen ion
gradient to generate energy in the form of ATP.
 The dark reactions also known as the Calvin cycle take place in the stroma of chloroplast.
 Production of NADPH2 molecules and oxygen as a result of photolysis of water.
 BY the utilization of assimilatory powers the 6-carbon atom is broken into two molecules of
phosphoglyceric acid.

CHECK YOURSELF
1. Holds the genetic information (DNA) for the cell. It controls all cell activities.
A) nucleus
B) ribosome
C) mitochondria
D) lysosome
2. Food producer for the plant cell. It converts the energy of the sun into sugar.
A) cell membrane
B) chloroplast
C) nucleus
D) cytoplasm
3. Acts as the digestive system inside a cell. It helps to break down old or unneeded parts of the cell, and
substances that have been brought into the cell from the outside.
A) endoplasmic reticulum
B) lysosome
C) ribosome
D) mitochondria
4. Monitors and controls entry into and out of the cell.
A) vacuole
B) chloroplast
C) cell membrane
D) ribosome
5. Creates proteins. It can float within the cytoplasm or be attached to an organelle.
A) ribosome
B) mitochondria
C) nucleus
D) cytoplasm
6. Checks, makes necessary changes, packages and secretes proteins.
A) Golgi Bodies / Golgi Apparatus
B) mitochondria
C) cell wall
D) endoplasmic reticulum
7. Jelly-like fluid that fills the cells and suspends the organelles.

17
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

A) cytoplasm
B) lysosome
C) nucleus
D) chloroplast
8. Thick, rigid layer that surrounds the plant cell and provides support and structure.
A) mitochondria
B) nucleus
C) cell wall
D) chloroplast
9. Helps transport proteins.
A) endoplasmic reticulum
B) cell wall
C) lysosome
D) vacuole
10. In plant cells, a large, fluid-filled space inside the cell that helps the cell maintain its shape and may also be
used to store nutrients and waste products. In animal cells, small fluid spaces inside the cell that are used to
store nutrients and waste products.
A) cell wall
B) ribosome
C) chloroplast
D) vacuole
11. Produces energy to fuel the cell’s activities.
A) mitochondria
B) cytoplasm
C) cell wall
D) Golgi Bodies / Golgi Apparatus
[Link] of electron (e-) microscope are
A. magnify the image up to 1000 times
B. produce black and white image
C. artificially colorized
D. to produce monochramitc images
[Link] denser protoplasm is called as
A. Cytoplasm
B. Nucleoplasm
C. cell sap
D. Mitochondrion
SECTION B
14. State the functions of the following organelles: nucleus,RER,SER,mitochondria,lysosomes
Answer Key
1.A 2.B 3.B 4.C 5.A 6.A 7.A 8.C 9.A 10.D 11.A 12.A 13.B

18
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 3
CELL DIVISION
 describe the process of mitosis
 explain the significance of mitosis in crop production
 describe the process of meiosis
 explain the significance of meiosis in plant reproduction
 compare and contrast mitosis and meiosis

CELL CYCLE AND MITOSIS AND MEIOSIS

The cell cycle

Actively dividing eukaryote cells pass through a series of stages known collectively as the cell cycle:
two gap phases (G1 and G2); an S (for synthesis) phase, in which the genetic material is duplicated;
and an M phase, in which mitosis partitions the genetic material and the cell divides.
G1 phase. Metabolic changes prepare the cell for division. At a certain point - the restriction point -
the cell is committed to division and moves into the S phase.
S phase. DNA synthesis replicates the genetic material. Each chromosome now consists of two sister
chromatids.
G2 phase. Metabolic changes assemble the cytoplasmic materials necessary for mitosis and
cytokinesis. Intensive cellular synthesis mitochondrion, chloroplast divide. Energy store increases
mitotic spindle begins to form.
M phase. A nuclear division (mitosis) followed by a cell division (cytokinesis).
The period between mitotic divisions - that is, G1, S and G2 - is known as interphase.

19
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CELL DIVISION

Introduction
Every individual grows in terms of size and also has a certain capacity of reproduce offspring. This is
brought about by mitosis and meiosis respectively. Take it from this angle, we all started as a single
cell (when the ovum fused with the sperm cell) and this cell divided into several cell to produce a
multi-cellular organism. The growth of a young animal or young plant in to adult, the development of
sex cells, all involve cell multiplication.
Mitosis is a process of cell duplication, or reproduction, during which one cell gives rise to two
genetically identical daughter cells.

Stages of mitosis

Mitosis is a nuclear division which involves four stages which are prophase, metaphase, anaphase and
telophase. There is a resting phase called interphase between telophase and prophase. The stages of
mitosis are described below in detail:

Interphase cell

This is the stage between two mitotic divisions. At this stage the genetic material is visible as
chromatin. Sometimes this stage is referred to as resting stage. There is DNA synthesis and replication
of centrioles.

Prophase

The chromosome further shortens and thickens and each appears as a double strand consisting of two
identical sister chromatids attached to a centromere. The nucleolus and nuclear membrane gradually
disappear and the centrioles moves to opposite ends of poles of the nucleus.

Centriole replicates (just before prophase)

Prometaphase

The chromosomes, led by their centromeres, migrate to the equatorial plane in the midline of cell - at
right-angles to the axis formed by the centrosomes. This region of the mitotic spindle is known as the
metaphase plate. The spindle fibres bind to a structure associated with the centromere of each
chromosome called a kinetochore. Individual spindle fibres bind to a kinetochore structure on each
side of the centromere. The chromosomes continue to condense.

20
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Metaphase

During this stage the chromosomes line up around the equator of the spindle. They are attached by
their centromere to the spindle fibres.

Chromosome line up across the equator

Anaphase

The centromere divides to separate the two chromatids, which then move apart and migrate towards
the opposite poles of the cells. Once separated, the sister chromatids are referred to as daughter
chromosomes. The separated chromatids are pulled along behind the centromere.

Telophase

It is tamed the reorganization phase that result in the constitution of two daughter nuclei. As soon as
daughter chromosome reaches the poles of the cell, they lengthen and uncoil to form chromatin. The
spindle fibre disintegrates. The nucleoli reappear and nuclear membrane is reformed.

Division of the cytoplasm (cytokinesis) starts during the late telophase. The process separates the
cytoplasm and newly formed daughter nuclei into two daughter cells.

21
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Significance of mitosis

 Facilitate growth of an organism.

 Basis of asexual reproduction.
 Ensure genetic stability (No variation).
 Cell replacement and cell repair

Meiosis

Meiosis is a special type of cell division that occurs in sexually reproducing organisms. It reduces the
chromosome number by half, enabling sexual recombination to occur. Meiosis of diploid cells
produces haploid daughter cells, which may function as gametes. Gametes undergo fertilization,
restoring the diploid number of chromosomes in the zygote

Meiosis, on the other hand, is a division of a germ cell involving two fissions of the nucleus and
giving rise to four gametes, or sex cells, each possessing half the number of chromosomes of the
original cell.

The stages of meiosis can be broken down into two main stages which are meiosis 1 and meiosis II.
The stages are described in detail below.

Meiosis I

Interphase

This is the resting stage. DNA replication takes place during this stage.

Prophase I
Most of the significant processes of Meiosis occur during Prophase I. The chromosomes condense and
become visible while the centrioles form and move toward the poles. The nuclear membrane begins to
dissolve and the homologue pair up, forming a tetrad. Each tetrad is comprised of four chromatids (the
two homologues), each with its sister chromatid.
Prophase I is divided into five phases:
Leptotene: chromosomes start to condense.
Zygotene: homologous chromosomes become closely associated (synapsis) to form pairs of
chromosomes (bivalents) consisting of four chromatids (tetrads).
Pachytene: crossing over between pairs of homologous chromosomes to form chiasmata (sing.
chiasma).
Diplotene: homologous chromosomes start to separate but remain attached by chiasmata.
Diakinesis: homologous chromosomes continue to separate, and chiasmata move to the ends of the
chromosomes.
Prometaphase I
Spindle apparatus formed, and chromosomes attached to spindle fibres by kinetochores.
The diagram below show a nucleus during prophase I.

22
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

During crossing over genetic material from the homologous chromosomes is randomly swapped and
this creates four unique chromatids as shown in the diagram below. Since each chromatid is unique,
the overall genetic diversity of the gametes is greatly increased (as shown below)

Metaphase I

Microtubules grow from the centrioles and attach to the centromeres and the tetrads line up along the
cell equator.

Anaphase I

The centromeres break and homologous chromosomes separate (note that the sister chromatids are
still attached). Cytokinesis also begins.

Telophase I
23
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The chromosomes reaches the poles and may decondense (depends on species) while cytokinesis

reaches completion, creating two haploid daughter cells

Meiosis II

It is important to note that meiosis II is similar to mitosis.

Prophase II
-centrioles form, move toward the poles and the nuclear membrane dissolves.

Metaphase II

Microtubules grow from the centrioles and attach to the centromeres and the sister chromatids line up
along the cell equator

Anaphase II
The centromeres break and sister chromatids separate and cytokinesis begins.

24
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Telophase II

The chromosomes may decondense (depends on species) and cytokinesis reaches completion, creating
four haploid daughter cells

Roles of meiosis

o Meiosis reduces the chromosome number by half, enabling sexual recombination to occur.
o Meiosis and fertilization introduce genetic variation in three ways:
 Crossing over between homologous chromosomes at prophase I.
 Independent assortment of homologous pairs at metaphase I: the chromosomes in meiosis
undergo a recombination which shuffles the genes producing a different genetic combination
in each gamete, compared with the co-existence of each of the two separate pairs of each
chromosome (one received from each parent) in each cell which results from mitosis.
 Random chance fertilization between any one female gamete with any other male gamete.

Activity
 Compare and contrast mitosis and meiosis.
 What are the roles of these two nuclear divisions in agriculture?
Summary
In this unit we looked at cell division i.e. mitosis and meiosis and to summarise it all, I will give you a
quick comparison of those processes.

25
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Comparison chart
Meiosis Mitosis
Definition: A type of cellular reproduction in A process of asexual reproduction in
which the number of which the cell divides in two producing a
chromosomes are reduced by half replica, with an equal number of
through the separation of chromosomes in haploid cell
homologous chromosomes in a
diploid cell.
Function: sexual reproduction Cellular Reproduction & general growth
and repair of the body
Type of Reproduction: Sexual Asexual
Occurs in: Humans, animals, plants, fungi all organisms
Genetically: different Identical
Crossing Over: Yes, mixing of chromosomes can No, crossing over cannot occur.
occur.
Pairing of Homologues: Yes No
Number of Divisions: 2 1
Number of Haploid 4 2
Daughter Cells produced:
Chromosome Number: Reduced by half Remains the same
Steps: The steps of meiosis are The steps of mitosis are Interphase,
Interphase, Prophase I, Metaphase Prophase, Metaphase, Anaphase,
I, Anaphase I, Telophase I, Telophase and Cytokinesis
Prophase II, Metaphase II,
Anaphase II and Telophase II.
Karyokenesis: Occurs in Interphase I Occurs in Interphase
Cytokinesis: Occurs in Telophase I & Occurs in Telophase
Telohpase II
Centromeres Split: The centromeres do not separate The centromeres split during Anaphase
during anaphase I, but during
anaphase II
Creates: Sex cells only: Female egg cells Makes everything other than sex cells
or Male sperm cells
CHECK YOURSELF
____ 1. Animal cells do NOT have ____.
a. centrioles c. cell plates/walls
b. centromeres d. cytoplasm
____ 2. Most of the life of any cell is spent in a period of growth called ____.
a. telophase c. interphase
b. prophase d. anaphase
____ 3. Which choice is NOT a process of the cell cycle?
a. death c. development
b. division d. growth
____ 4. Which choice best describes the cell cycle?
a. Cells grow and develop during interphase. Cells reproduce during the mitotic phase.
b. Cells grow and develop during the mitotic phase. Cells reproduce during interphase.
c. The nucleus of a cell divides during interphase. The cytoplasm of a cell divides during the
mitotic phase.
d. The nucleus of a cell divides during the mitotic phase. The cytoplasm of a cell divides
interphase.
____ 5. How long does it take for most dividing human cells to complete a cell cycle?

26
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

a. a few minutes c. a year

b. a day d. about 7 years
____ 6. What is the first thing that happens when a new cell is produced?
a. It gets smaller. c. It divides again.
b. It gets larger. d. It rests.
____ 7. During which stage of interphase do cells perform their normal cell functions (such as growing
and making enzymes to digest your food)?
a. S stage c. G2 stage
b. G1 stage d. Mitosis
____ 8. During which stage of interphase do cells copy their DNA?
a. S stage c. G2 stage
b. G1 stage d. Cytokinesis
____ 9. During which stage of interphase do cells store energy in final preparation for use in the mitotic
phase?
a. G2 c. S
b. G3 d. S2
____ 10. During the S stage of interphase, pairs of identical chromosomes are held together by which
structure?
a. centromere c. connectroid
b. chromatid d. centromatid
____ 11. Identify two stages of the mitotic phase.
a. prophase and metaphase
b. cytokinesis and interphase
c. mitosis and interphase
d. interphase and metaphase
____ 12. If a cell has 22 duplicated chromosomes, how many chromatids does it have?
a. 22 c. 11
b. 44 d. 88
____ 13. What is the shortest phase of interphase?
a. G2 c. S
b. metaphase d. prophase
14. Which type of cell divides by the cell membrane pinching together until the two cells split apart?
a. plant c. cytoplastic
b. animal d. chromosomal
____ 15. Which best describes how a plant cell divides?
a. A new cell plate and wall forms in the middle of the cell and two new cells are formed.
b. The two cells twist apart.
c. The membrane pinches shut in the middle of the cell and the cells are split apart.
d. Plant cells do not divide.
____ 16. Which of the following is NOT a result of cell division?
a. reproduction c. growth
b. nutrition d. repair
____ 17. How long does it take a cell to complete the cell cycle?
a. 8 minutes
b. 1 year
c. 24 hours
d. The time it takes depends on the type of cell that is dividing.
1. C 2.C 3.A 4.A 5.B 6.B 7.B 8.A 9.A 10.A 11.A 12.B 13.A 14.B 15.A 16.B 17.A
SECTION B
[Link] in detail the process of mitosis and meiosis.[50]
19. State the importance of mitosis and meiosis in crop production.[10]

27
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 4
SOIL FERTILITY AND PLANT NUTRITION
 Identify the soil minerals.
 describe the four components of soil
 Discuss the significance of soil water to plant growth.
 explain the role of soil air in plant growth and microbial activity
 Describe the significance of soil horizons and catena effect.
 Discuss the significance of soil color, texture and structure in crop production.
 Describe the management practices of soil structure and texture in crop production.
 Discuss the importance of soil organic matter.
 Determine soil bulk and particle density and porosity.
 explain the roles of macro- nutrients and micro nutrients in plant growth and development
 identify constituents of soil organic matter
 describe the role of soil organisms in organic matter decomposition
 identify organisms involved in different stages of the decomposition process
 describe the carbon: nitrogen (C:N) ratio
 describe how the C:N ratio affects the rate of decomposition
 Determine fertilizer requirements in crops.
 Describe factors affecting soil organic matter levels.
 discuss the benefits of soil organic matter
 Describe the formation of clay and humus colloids.
 Describe the basic structure of clays.
 Explain the source of negative charges on clay and humus colloids.
 Explain the origins and significance of cation exchange and anion exchange capacity (CEC and
AEC).
 Determine cation exchange capacity (CEC) and base saturation percentage.
 Discuss the significance of base saturation and exchangeable sodium percentage (ESP).
 Discuss the causes of soil acidity and salinity.
 Discuss the effects of soil acidity and salinity on crop growth.
 Explain methods of correcting soil acidity and salinity.
 Carry out experiments on soil analysis.
 Determine soil pH and calculate liming requirements.

MINERAL COMPOSITION OF SOIL PARTICLES

28
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Primary silicate minerals=micas, hornblendes, feldspars

Other secondary minerals= oxides/hydroxides of Fe/Al
Secondary silicate minerals=kaolinite, illite, montmorillonite
PRIMARY MINERALS
Are those which have not been altered chemically since deposition and crystallization from molten
lava. Disintegration of rocks composed of primary minerals (by physical and chemical weathering)
releases the individual mineral particles. Many of these primary mineral particles become sand and silt
particles in parent materials and soils. Primary minerals weather chemically (decompose) and release
their elements to the soil solution. Some of the elements released in weathering react to form
secondary minerals.
SECONDARY MINERALS
Are formed from chemical weathering or transformations of primary minerals.A secondary mineral
results from the decomposition of a primary mineral or from the precipitation of the decomposition
products of minerals. Secondary minerals originate when a few atoms react and precipitate from
solution to form a very small crystal that increases in size over time. Because of the generally small
particle size of secondary minerals, they dominate the clay fraction of soils and are divided into two
broad groups, namely:
 secondary silicates
 Oxides/hydroxides of iron and aluminum.
Primary minerals are predominantly silicates, which have a crystalline structures based upon a simple
unit, called a silicon tetrahedron. These silicon tetrahedrons can be joined together in rows, sheets or
3-dimensional structures to form different types of minerals.

SOIL CHARACTERISTICS

Soil components

The soil consists of about half solids and half fluids (gas/liquid). The fluids are found in the soil pores
between the solid particles. Figure 1.1 below shows that there are four major components of soils.
This figure shows the approximate proportions by volume of these components in a loam soils.
NB The line dividing the air from the water is flexible. This means that the volume of air and water
vary in a soil depending on its condition and time of the season. For optimum plant growth, air and
water volumes should be roughly equal at about 25% of the total soil volume. The water in soil is
actually a solution and it contains many dissolved salts. This solution provides the nutrients to plants.
Oxygen is required for respiration of roots and other soil organisms. Mineral material has various
types of particles ranging from large ones that we can see (gravel, sands to those that are too small for
us to see e.g. clays. Organic fraction consists of fresh, decomposing and decomposed organic matter in
various stages of decay.

29
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Components of the soil on a volume basis

Soil water

Uses of soil water

 Transport nutrients in solution.

 Support chemical weathering reactions.
 For soil temperature regulation.
 For the maintenance of plant structure (cell turgidity).
 Imbibed by seeds and lead to germination (activate enzymes).
 For metabolism, thus supporting major cell constituents.
 Supporting of microbial activity.

Types of capillary pores

Micro-pores/capillary pores- they hold capillary water at field capacity.
Macro-pores- are air occupied (hold water in excess). Saturation point is reached when all pores are
occupied by water. It is also known as maximum retention capacity, water drains in 48 hrs to leave
soil at field capacity (drains by gravity).
Factors affecting capillary water amount
Soil texture-pore spaces increase as texture gets finer.
Relative proportion of micro and macro pores

Soil type relative proportion of pore spaces

Sandy soils micro< macro
Loamy soils micro= macro or are well drained
Clay soils micro> macro
Soil texture are granular structure which has (micro> macro) this hold more capillary water than other
structures.
Organic matter content- micro-pore capacity increase with an increase in organic matter and organic
matter turns into humus to form crumby structure.

Forces of water retention in soil

Hold water against drainage due to gravity
a) Adhesion forces- are the attraction of water molecules to soil surface, where water and soil
particles interface. The force attaches a thin film of water, and this is called hygroscopic water
or adhesion water.
b) Cohesion forces- are the attraction between water molecules and it enables thin film of water
around soil particles outside hygroscopic water. The energy is in soil water retention, the
greater the soil moisture tension the more tightly held is the water (plants fail to utilize it). Soil
moisture tension increases with a decrease in soil moisture.

Moisture and tension relationship

 At saturation point tension is almost zero and excess water around soil particles drains out, in
48hrs from saturation soil is at field capacity and moisture stabilizes.
30
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Field capacity
Is amount of water on mass to volume basis which a soil can hold or retain against drainage to gravity
or the amount of water left in a soil 2 days after saturation point and excess water has drained. Water
at field capacity differ with soil texture, structure and organic matter content and the tension holding
water to soil particles increase as crops extract moisture. Tension will reach a point when plants
cannot extract water anymore or smaller than needed. At the above point plants remain wilted even if
water is added. This is called permanent wilting point (wilting co-efficient).

Classification of soil water

Physical classification- this is classification according to tension or force holding the water.

 Gravitational water or drainage water-this is excess water above field capacity that drains. This
water is held at very low tension (0.3 atmospheres). The water can cause leaching of minerals.
 Hygroscopic water- is extremely thin film of water held tightly to the soil particles at a high
tension of about 31 atmospheres. This is adsorbed water not in capillary and is not available to
plants. The water is present even in dry soils.
 Capillary water- this water which is held between hygroscopic water and field capacity or is the
film of water surrounding soil particles outside hygroscopic water and is held in micro pores. This
is the most available water to plants.

Biological classification
It is water classification according to availability to plants for uptake.

 Superfluous water- it is undesirable water or gravitational water and it causes leaching, water
logging and reduce aeration.
 Available water- is water held between field capacity and permanent wilting point (PWP).
 Undesirable water- is water held at 15 atmospheres and above in tension. This water is not
available to plants.
Available water capacity (AWC) of soil is the amount of water a soil can hold between field capacity
and permanent wilting point and this water on soil texture and structure.

Water movement in soil

 Saturation flow is moisture at which tension is almost zero, water is actively being drained by
gravity down the profile. It is downward water movement through saturation or near saturation soil
through gravity (percolation). Amount of percolation depends on amount of rainfall, infiltration
capacity of a soil, rate of penetration to lower horizons. In general it depends on soil organic
matter, texture and structure of horizons. Sandy soils have high percolation rate while heavy soils
have low percolation rate.
 Unsaturated flow is the water moving along moisture tension gradient, water moves from region of
low tension to region of high tension.
 Vapour flow water vapour movement along vapour pressure gradient by diffusion. Vapour moves
from high to low pressure or from soil to atmosphere. Evaporation from soil decreases with an
increase in relative humidity and evaporation increases with an increase in temperature and will
increase with the wind.

Summary

31
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

This unit has covered soil minerals, how the physical properties of the soil affect water absorption and
air circulation in the soil. It also constitutes the properties of soil which promote maximum production
of crops. This section also indicates the type of water which is available for plant uptake in the soil

CHAPTER 5

PHYSICAL PROPERTIES OF THE SOIL

Objectives
At the end of this unit students should be able to:

 Describe the significance of soil horizons and catena effect.

 Discuss the significance of soil colour, texture and structure in crop production.
 Describe the management practices of soil structure and texture in crop production.
 Discuss the importance of soil organic matter.
 Determine soil bulk and particle density and porosity.

Soil texture

It is the relative proportion of sand, silt and clay in any given soil; it is also referred to as the
coarseness/finess of the soil. Texture is a permanent property that does not readily change from year to
year. Once we know the amount of sand, silt and clay in a soil we can therefore place the soil into a
textural class using a textural triangle. Examples of textural classes are sandy loam and loamy sand
soils.

32
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Particle size analysis estimates the percentage sand, silt and clay contents of a given soil and is often
reported as percentage by weight. This can be done qualitatively (using a Finger Assessment
Technique).Finger assessment technique provides the broad textural category of soil. The more sticky
and mouldable a wet soil is, the greater the clay content.
Or quantitatively using sieving and sedimentation. The sieving and sedimentation techniques are two
complimentary methods.
Sieving
It is a simple procedure which separates soil particles > 50 micrometres in diameter. A sample is
emptied into an assemblage of sieves of known diameters, with the largest sieve on top and smallest at
the bottom. After a period of shaking, the sand grains will settle into the different sieves. These
fractions can then be weighed and their size distribution calculated.
Sedimentation
Sedimentation on the other hand is used to separate particles which are < 50 micrometres (silt and
clay) in diameter. The two major sedimentation techniques are the hydrometer and the pipette
method.
The hydrometer method.
The hydrometer is an instrument which measures the density (g per litre) of a soil-water suspension at
a given time.
The first stage is the dispersion of soil into individual particles. This is achieved through mechanical
shaking in calgon (sodium hexametaphosphate) solution. Individual soil particles are often bound into
aggregates hence the need for dispersion. In clay soils with high organic matter content, there may be
need to oxidize cementing organic compounds using hydrogen peroxide. In highly calcareous soils, it
is important to dissolve carbonates using acids.
The hydrometer method is based on Stokes Law which states that:
The settling velocity of a particle is the net difference between its downward forces (gravity) against
the buoyancy (resistance to fall) by surface friction and movement of the water.
- When soil particles are suspended in water they tend to sink
- The settling velocity of a particle is a function of liquid temperature, viscosity and specific gravity of
the settling particle.
- The particles are assumed to be spherical (weakness of Stokes Law).
According to Stokes Law the settling velocity of a particle is proportional to the square of the radius of
the particle.
V = 2r2 (Ps – Pw) g
9n
v = terminal settling velocity, r = particle radius, Ps = particle density, Pw = density of water, g =
acceleration due to gravity, n = viscosity of water.
High temperatures result in reduced viscosity (why?)
The sand grains settle first because they are larger followed by silt and the clay remains as a colloidal
suspension.
Significance of soil texture
To deduce useful information about the soil chemical and physical behaviour such as its likely nutrient
content, water holding capacity, aeration status, easy of tillage and potential problem. The other reason
is to know the soil capillarity (upward movement of water against force of gravity.

Soil structure

This is the arrangement/combination of primary soil particles into secondary particles, aggregates
which are separated from one another by planes of weaknesses or an association or aggregation of
primary soil particles sand, silt and clay into clusters or particles called peds. The particles are bound
33
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

together by materials such as organic matter, silica and iron oxides. Between the peds are pores or air
spaces and soil structure is determined when soil is dry.
Type of soil structures
There are four types of structures. These are:

 Spheroidal structure which can be granular and crump structures

 Blocky which can be angular or sub angular blocky structures
 Prismatic which can be prism like or columnar
 Plate structure

Importance of crumby structures to a farmer

 They provide adequate micro-spores for good aeration, they provide oxygen for root respiration
and germination, and microbial activities like decomposition and chemical weathering reactions
(oxidation) are activated.
 It improves the drainage of the soil. The peds are separated by micro spores and this enable the
soil absorb water easily (reduce runoff).
 Shape and size of peds- allows good soil seed contact for imbibing water and germination.
 Easy root penetration, there is increased soil volume for nutrient and moisture uptake by plants.
 The soil is easily workable and requires fewer droughts.
Methods of forming and keeping crumb structures
Tillage destroys soil structure, practice conservation farming. Timing of cultivation, do not work on
soil when too wet or too dry. When the soil is dry it is pulverized into dust and clay form clods
(structure is destroyed). Incorporation of farm manure, green manure, compost and plant residues
improve soil structure (crumb) and decaying organic matter bind soil particles into crumbs. Addition
of lime results in flocculation or binding together of clay particles to form crumbs.
Soil aeration
Pores are spaces between soil particles or non-solid part of soil through which water circulates, air
circulates and plant roots penetrate. Pore spaces constitute 50% by volume of soil, half is occupied by
water and percentage volume occupied by air and water is called porosity. Soil air has more carbon
dioxide and water vapour, excess water fills micro and macro pores and gases diffuse out of soil and
low oxygen for root respiration and oxidation reactions results.

Effects of poor soil aeration

Most plant roots fail to respire and die without oxygen for long except for rice in water logged areas.
Oxygen deficiency impairs root ability to absorb nutrients. Tomatoes in saturated soils where oxygen
is deficiency fails to absorb water and wilts (wet wilting). Anaerobic conditions reduce microbes’
population (nitrifying bacteria) and denitrifying bacteria increases and this will reduces available
nitrogen in the soil. Anaerobic bacteria carryout reduction reaction producing toxic substances like
hydrogen sulphate and acetic acid (toxic to plant roots).

Factors affecting soil aeration

 Soil texture – the finer the texture the greater the porosity of a soil and it is dominated by micro
pores holding moisture.

34
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Soil structure – more crumby structures had more porosity, crumby peds do not join and have
more air spaces.
 Soil depth – the deeper you go the less the porosity, this is due to compaction of the soil.
 Application of organic matter – improves soil structure to more crumby and greater pore capacity.

Methods of increasing soil aeration

Soil loosening improves aeration by encouraging air circulation (deep and thorough ploughing,
hoeing). Addition of organic matter leads to crumb structure.

SOIL COLOR

The most considered colour is that of “B horizon” on the upper slope. A horizon colour is mashed by
humus and it is brown or black coloured. The colour of C horizon varies and is beyond reach of many
crops. The colour is determined on moist soil using Munsell colour chart. The system has a colour
chart book with 3 variables.
i) The Hue represents dominant spectral colour e.g. red, yellow and yellow red and this scores 0 -
10.
ii) Value represent amount of light reflected and gives lightness or darkness of colour and score
from 0 – 10(0 – black and 10 – white).
iii) Chroma represents purity of colour, colours which are greyer.
Significance of soil colour
A soil colour change along a slope is due to differences in drainage. Upper slope soils are well
drained, middle slope soils are imperfectly drained while lower and lower most slopes are poorly
drained.

Interpretation of different soil colours

Red colour
Red to reddish brown soils is found on the upper slopes. It indicates well drained and well aerated
soils, the soils are dominated by hydrated iron III oxide (red in colour) and the manganese dioxide
(product of oxidation) adds to the red colour.

Brown soil colour

Brown colour is found in middle slopes. It indicates imperfect drainage and aeration this is because of
lateral moisture movement from upper slopes. The soil remains moist for long period and the brown
colour results due to partly reduced and hydrated iron oxides.

Yellow brown to yellowish grey soils

This colour is found on the lower slopes and the colour is given by reduced and hydrated iron
compounds. This colour indicates further depressed drainage and aeration and the soils are wet over a
long period than middle slopes soils.

Grey or gley soils

Part or whole soil profile is water logged where these soils are found. It indicates very poor drainage
and aeration. Grey or bluish or greenish or gley colour is exhibited. Gley soils results from completed

35
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

reduction of iron III oxide to iron II oxide/complete iron reduction. Most of iron combines with
organic matter as iron II oxide and iron III hydroxide that give grey colour.
Black
Soils are found on lower and lower most slopes. The colour is associated with 2:1 clay minerals,
montimorrillonite is dominant. In high rainfall areas, colour varies along slope and montimorrillonite
gives black colour. Soils are derived from marfic and ultra marfic rocks. The areas are generally low
lying and poorly drained and aerated, the soil should not be confused for vertisols (black soils) derived
from basalt in low rainfall.

Other soil characteristics on the slope

Mottling is the red or yellow spots or blotches interspaced with dominant colour of soil. These
indicates seasonal water table fluctuations, colours are caused by repeated reduction and oxidation of
iron and manganese due to increase and decrease in water table. Oxidation occurs when water table
falls and reduction when soils are water logged or when water table increases. Mottles are only found
in part of soil where water table varies, amount of mottles indicates degree of water table fluctuations.
Concretions are grains or nodules of varying size and hardness as well as shape and colour. They are
made up of chemical compounds in soil, oxides of iron and manganese are hard black nodules when
there is a water table fluctuation, the concretion form hard compact layers impermeable to roots and
this limits effective soil depth.

Soil profile and Horizons

Profile it is the vertical cross-section through a soil extending from surface to parent rock. If a deep
hole is dug in a soil, distinct horizontal layers are noticed.
Horizons it consist of individual layers different from each other. N.B. each soil has its own distinct
profile characteristics.

Reasons for studying the soil profile

It tells us more about the formation properties of soil; it will help in deciding how best one can use his
soil. As weathering progresses further clay minerals and sesquioxides are leached 15 – 50cm and
deposited. The removal of the minerals and sesquioxides is called elluviation and the deposition of
these sesquioxides and minerals to form part of B horizon is called illuviation.

O- Horizon (organic matter)

A-Horizon (top soil)

E-Horizon (elluviation)

B-Horizon (subsoil illuviation zone)

C-Horizon (mineral zone/weathering rock)

R-Horizon (unweathered bed rock)

36
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Figure: Soil horizon.

Horizons and their characteristics

O- Horizon
This is an organic rich layer often found at the surface in humid climate. It may contain
decomposed/undecomposed organic matter and this do not commonly occur in Zimbabwe.
A-Horizon
This layer support most plants and life and is usually is darker than the subsoil horizon.
B-Horizon
This horizon is characterized by accumulation of clay, iron, aluminum and humus. It is often lighter in
colour than the A horizon and is more structured.
C –horizon
This horizon contains weathered rock in transition between the weathered soil and the unweathered
parent materials.
R- Horizon
This horizon is made up of continuous hard bed rock which may or may not be the parent material of
the soil above it.

Activities

 Discuss hydrolysis as a chemical weathering which affect soil formation.

 Discuss five factors which affect soil formation.
 Describe how human activities affect soil formation.
 Draw a well labelled soil catena and describe it
Summary
From this unit we have noticed that weathering produces a soil which consist of a mixture of minerals
at various stages of decomposition and this depends on the original mineral composition, their
resistance to weathering, the amount of water present, temperature and oxidation or reduction reaction.
Leaching also removes soluble constituent from soil when excess water drains down a soil, soil turns
acidic and the bases are reduced e.g. calcium, magnesium, potassium and sodium. Quartz, kaolinite,
iron III oxide and aluminium oxide remains. With time and interaction of many other factors soils
with different horizons are formed as the end product of weathering.

References
Bear, F. E. (1964). Ed. Chemistry of the Soil. Oxford and IBH Publishing, India.
Foth, H. D. (1984). Fundamentals of Soil Science. John Wiley and Sons, USA.

Topography/relief

This modifies soil profile in three ways:

 By influencing rainfall absorbed and therefore affecting the amount of moisture in the soil.
37
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 By influencing the rate of soil erosion.

 By influencing the amount of subsurface water movement which will transport soil material from
one place to the other.
The main influence is on the movement and distribution of water within and over soils. The steeper
the slope the greater the runoff and erosion, less water enters the soil to be available for chemical and
biological weathering. Therefore soils on the steep slopes are usually shallow and poorly formed and
this influences vegetation growth and this can be explained by the soil catena concept.

Soil catena
This is a sequence of soils derived from the same parent material but differ in properties due to the
different topographic positions they occupy. The differences in soil properties arise because of the
differences in soil moisture regime that exists between soils in the sequence.
Crest position soils lower slope valley bottom: vlei, accumulation
Often shallow with subject to of bases and clay, may be permanently
Rock outcrops water logging or gleyed
Red Runoff carries during the
Bases and rainy season River
Brown Soil (mottles)
Water table and water movement Yellow Black/gley soils

Figure 2.1: Soil Catena

In the upland position, just below the crest, sufficient rainfall penetrates the soil to allow for
weathering and development of a deep profile. This site is well drained (above water table) and water
moves internally and carries bases down slope and the soil is mostly composed of stable iron oxide
and is red in colour. Lower down slope, the water table is now closed to the surface and influences the
soil morphology. The soil experiences seasonal water logging which induces formation of mottles in
the subsoil and the soil is brown due to the iron oxide being reduced. Further down slope the soils
become permanently water logged and all iron is reduced to the ferrous state. There is little stream
flow and the poor drainage encourages accumulation of bases, organic matter and presence of active
swelling clays. The clays combine with organic matter to produce black soil above the waterline. Soils
below the waterline which are permanently wet exhibit gley colour.

SOIL AIR

Are the gases found in the air space between soil components or gases occupying the free pore space
in soil. The primary natural soil gases include nitrogen, carbon dioxide and oxygen. The oxygen is
critical because it allows for respiration of both plant roots and soil organisms. Other natural soil gases
are atmospheric methane and radon. Some environmental contaminants below ground produce gas
which diffuses through the soil such as from landfill wastes, mining activities, and contamination by
petroleum hydrocarbons which produce volatile organic compounds.

38
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

In nutrient management, soil aeration influences the availability of many nutrients. Particularly, soil
air is needed by many of the microorganisms that release plant nutrients to the soil. An appropriate
balance between soil air and soil water must be maintained since soil air is displaced by soil water.

Air can fill soil pores as water drains or is removed from a soil pore by evaporation or root absorption.
The network of pores within the soil aerates, or ventilates, the soil. This aeration network becomes
blocked when water enters soil pores. Not only are both soil air and soil water very dynamic parts of
soil, but both are often inversely related:

 An increase in soil water content often causes a reduction in soil aeration.

 Likewise, reducing soil water content may mean an increase in soil aeration.
 Since plant roots require water and oxygen (from the air in pore spaces), maintaining the balance
between root and aeration and soil water availability is a critical aspect of managing crop plants.

Soil air is very different than the above-ground atmosphere. A significant difference is between the
levels of carbon dioxide. Since the soil contains high amounts of carbon dioxide, oxygen levels may
become limited. Since plants must have oxygen to live, it is important to allow proper aeration in the
soil. See Tables 3 and 4 for references to soil air composition.

Factors Affecting the Composition of Soil Air:

1. Nature and condition of soil: The quantity of oxygen in soil air is less than that in atmospheric air.
The amount of oxygen also depends upon the soil depth. The oxygen content of the air in lower layer
is usually less than that of the surface soil. This is possibly due to more readily diffusion of the oxygen
from the atmosphere into the surface soil than in the subsoil. Light texture soil or sandy soil contains
much higher percentage than heavy soil. The concentration of CO2 is usually greater in subsoil
probably due to more sluggish aeration in lower layer than in the surface soil.

2. Type of crop: Plant roots require oxygen, which they take from the soil air and deplete the
concentration of oxygen in the soil air. Soils on which crops are grown contain more CO2 than fallow
lands. The amount of CO2 is usually much greater near the roots of plants than further away. It may
be due to respiration by roots.

3. Microbial activity: The microorganisms in soil require oxygen for respiration and they take it from
the soil air and thus deplete its concentration in the soil air. Decomposition of organic matter produces
CO2 because of increased microbial activity. Hence, soils rich in organic matter contain higher
percentage of CO2.

4. Seasonal variation: The quantity of oxygen is usually higher in dry season than during the
monsoon. Because soils are normally drier during the summer months, opportunity for gaseous
exchange is greater during this period. This results in relatively high O2 and low CO2 levels.
Temperature also influences the CO2 content in the soil air. High temperature during summer season
encourages microorganism activity which results in higher production of CO2.

Importance of Soil Aeration

1. Plant and root growth: Soil aeration is an important factor in the normal growth of plants. The
supply of oxygen to roots in adequate quantities and the removal of CO2 from the soil atmosphere are
very essential for healthy plant growth.

39
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

When the supply of oxygen is inadequate, the plant growth either retards or ceases completely as the
accumulated CO2 hampers the growth of plant roots. The abnormal effect of insufficient aeration on
root development is most noticeable on the root crops. Abnormally shaped roots of these plants are
common on the compact and poorly aerated soils. The penetration and development of root are poor.
Such undeveloped root system cannot absorb sufficient moisture and nutrients from the soil

2. Microorganism population and activity: The microorganisms living in the soil also require oxygen
for respiration and metabolism. Some of the important microbial activities such as the decomposition
of organic matter, nitrification, Sulphur oxidation etc. depend upon oxygen present in the soil air. The
deficiency of air (oxygen) in soil slows down the rate of microbial activity.

For example, the decomposition of organic matter is retarded and nitrification arrested. The
microorganism population is also drastically affected by poor aeration.

3. Formation of toxic material: Poor aeration results in the development of toxin and other injurious
substances such as ferrous oxide, H2S gas, CO2 gas etc. in the soil.

4. Water and nutrient absorption: A deficiency of oxygen has been found to check the nutrient and
water absorption by plants. The energy of respiration is utilized in absorption of water and nutrients.
Under poor aeration condition (this condition may arise when soil is water logged), plants exhibit
water and nutrient deficiency

5. Development of plant diseases: Insufficient aeration of the soil also leads to the development of
diseases. For example, wilt of gram and dieback of citrus and peach.

SOIL BULK DENSITY, PARTICLE DENSITY AND POROSITY

Soil water and air occupy voids in the soil, called pore spaces. The pore system in soil provides the
conduits for air and water exchange and houses roots and microbes. Soil porosity is the amount of
pore volume (%age of pore space). A medium textured, well-aggregated soil contains about 50% pore
space and is in good condition for plant growth when the pores hold an equal distribution of air and
water. Pore size affects pore activity. Big pores, macropores, facilitate free-water drainage, aeration,
evaporation, and gas exchange. Mesopores, medium-size pores, are essential to capillary water
distribution, and micropores provide water storage sites. Macropores are most prevalent in sandy soils
and well-aggregated soils, but can be converted to micropores by compaction. Medium-textured soils
have an abundance of mesopores. Clays promote aggregation but can also be readily compacted. Clays
also increase water storage by providing an abundance of micropores. Thus, texture and structure, plus
the level of induced compaction, are the main properties governing amount and type of pore space in
the soil. Organic matter affects porosity through its enhancement of soil aggregation.
Porosity can be calculated if bulk density and particle density are known. Bulk density is soil mass
divided by unit volume. In its natural state, a soil's volume includes solids and pores, therefore, a
sample must be taken without compaction or crumbling to correctly determine bulk density.

Bulk density = Oven dry soil weight / volume of soil solids and pores
Bulk density of mineral soils commonly ranges from 1.1 to 1.5 g/cm3 in surface horizons. It increases
with depth and tends to be high in sands and compacted pan horizons, and tends to be low in soils with
abundant organic matter. Tillage operations loosen soils and temporarily lower bulk density, while
compaction processes raise bulk density. High bulk densities correspond to low porosity. Natural soil-

40
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

forming processes that increase aggregation reduce bulk density, but excessive tillage and raindrop
impact on bare soil destroy aggregation and increase bulk density.
Methods of determining bulk density (Clod, Core and Sand replacement method)
1. Core method
A sharpened, open-ended cylindrical metal container, or ring, is carefully pushed into a soil until the
level of the ring is flush with the soil surface. The core is then carefully extracted, excess soil is
trimmed away until flush with the ends of the ring. In the laboratory, the core plus soil are oven-dried
and the soil is weighed.
The bulk density is then calculated by dividing the oven dry mass of soil by the total volume of the
core.
The volume of the core is given by:
Volume = (π) (r2) (h)
2. Sand replacement method
This method is mainly used by engineers. A hole is made in the surface of the soil, about 100 mm
diameter by 150 mm deep and the extracted soil is weighed after oven drying to obtain (Ms). The
volume of the hole is determined by filling it with a measured quantity of dry sand of known density
(D = M/V). By dividing the mass of excavated soil with the volume, the bulk density is determined.
Example: Calculate the mass (kg) of soil in 1 hectare of top soil 0-0.25 m in depth, if the bulk density
is 1300 kg m-3.
Factors affecting bulk density

1. Pore space: Since bulk density relates to the combined volume of the solids and pore spaces, soils
with high proportion of pore space to solids have lower bulk densities than those that are more
compact and have less pore space. Consequently, any factor that influences soil pore space will affect
bulk density.
2. Texture: Fine textured surface soils such as silt loams, clays and clay loams generally have lower
bulk densities than sandy soils. This is because the fine textured soils tend to organize in porous grains
especially because of adequate organic matter content. This results in high pore space and low bulk
density. However, in sandy soils, organic matter content is generally low, the solid particles lie close
together and the bulk density is commonly higher than in fine textured soils.

3. Organic matter content: More the organic matter content in soil results in high pore space there by
shows lower bulk density of soil and vice-versa.

PARTICLE DENSITY
Particle density is the volumetric mass of the solid soil. It differs from bulk density because the
volume used does not include pore spaces.

Particle density = oven-dry soil weight / volume of soil solids

Particle density represents the average density of all the minerals composing the soil. For most soils,
this value is very near 2.65 g/cm3 because quartz has a density of 2.65 g/cm3 and quartz is usually the
dominant mineral. Particle density varies little between minerals and has little practical significance
except in the calculation of pore space.
Porosity is that portion of the soil volume occupied by pore spaces. This property does not have to be
measured directly since it can be calculated using values determined for bulk density and particle
density. Finding the ratio of bulk density to particle density and multiplying by 100 calculates the
percent solid space, so subtracting it from 100 gives the % of soil volume that is pore space.

41
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

% solid space = (bulk density / particle density) x 100

% porosity = 100 - (% solid space)

Sample Calculation of Porosity:
A 260 cm3 cylindrical container was used to collect an undisturbed soil sample. The container and soil
weighed 413 g when dried. When empty the container weighed 75 g. What is the bulk density and
porosity of the soil?
Types and functions of pores
Macropores > 50 micrometres (diameter)
Mesopores 0.5 – 50 micrometres
Micropores < 0.5 micrometres
Pores have three main functions and these are: transmission pores, which are vital for easy
movement of water, nutrients and air, storage pores which are vital for retention of water for use by
crops and Residual pores which are water tightly bound and plants cannot utilize it
A good agricultural soil has a combination of the three pore types.

A. To determine bulk density:

Sample Volume = 260 cm3; Sample Weight = 413 - 75 = 338 g; Bulk density = 338 g/260 cm3= 1.3 g
/cm3

B. To determine porosity:
Bulk density = 1.3 g /cm3; Particle density = 2.65 g /cm3; Porosity = 100 - (1.3/2.65 x 100) = 51%

COLLOIDS AND CLAY MINERALS

Introduction

We have covered the physical properties of soil in the previous unit. This and the next unit cover the
chemical properties. During the course of this unit try to link the chemical and physical properties of
the soil and both influences soil properties.

Objectives
By the end of this section you should be able to:
 Define the terms; colloid, flocculation, cat ion exchange capacity, base saturation and give
their significance in agriculture.
 Describe the structures of the different groups of clays and relate them to their properties.
 Describe the nutrient uptake mechanism in plants

The chemical properties of the soil are largely determined by the clay content of the respective soil
Clay particles are small (<0.002mm in diameter) and together with humus particles which are equally
as small they form what are known as colloids. Clays are inorganic colloids which are either residues
of the original minerals or they may be secondary or even tertiary products of the parent material.
Humus falls in the organic colloids class are made up decomposed plant, animal and microbial matter.
These two colloidal forms may at time be combined with each other (Bear, 1964). As shall be

42
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

discussed later, clays are the last products in the decomposition of rocks whilst humus is the last
product of the decomposition of organic materials.

Colloids have a large surface area to volume and as such they tend to have very high charges. Colloids
have two common characteristics which arise as result of the high charges that surround them;
flocculation and cat ion exchange capacity (CEC). The charges are mainly negative and they are
caused by several factors some of which are discussed below.

Causes of negative charges in colloids

i. Isomorphous substitution is the replacement of an ion (cat ion) by another one of more or
less the same size but of a different charge. Negative charges are going to arise when a cat
ion of a lower replaces one with a higher charge on the colloidal structures. Cat ions are
found within the layers of the colloidal structure s as shall be explained when the structures
of clays get described in this section. Al3+ can replace Si4+, Mg2+ and Fe2+ can replace Al3+
in the colloidal structures. This replacement is going to create extra unsatisfied negative
charges.
Dissociation of groups like the siloxane groups (-Si-OH) found at the edges of clay
crystals. These charges are pH dependent, as the pH increases, in other words as the
concentration of hydroxyl (-OH) ions increases, these charges also increases as shown by
the reaction shown below
SiOH + OH……………SiO- + H2O

ii. Presence of carboxylic, phenolic, enolic and imide chemical groups these groups
normally have the hydrogen ion on their –OH groups removed, living behind –O- groups
which are negatively charged. The dissociation is also dependent on the pH of the soil.
iii. Unsatisfied bonds at the edges of the clay crystals

Flocculation
It is the coming together of soil particles to form crumbs. Colloids are naturally negatively charged,
the causes of the negative charges have been discussed above. The negative charges cause colloidal
repulsion since like charges repel. However in the natural soil there are reserves of positively charged
ions, the cat ions, these are attracted to the negative charges that are on the surfaces of colloids. As a
result of the attraction, the repulsive forces operating between colloidal particles are going to be
cancelled as the cat ions are going to wedge themselves between the repelling anions. As a result
these colloidal structures are going to come together to form soil particle groupings that have a
definite structure and individual structures are also called peds. The agricultural significance of
flocculation has already been discussed under soil structure when you were introduced to the
significance of the crumb structure to agriculture.

Cation Exchange Capacity

Since clay crystals are negatively charged, they attract cat ions, the positively charged ions towards
themselves. Some of the cat ions that can be attracted include; K+, Ca2+, Al3+, H+, Na+, Mg2+, Mn2+,
Zn2+, Cu2+, NH4+, Fe2+. These cat ions are held tightly to the colloids and therefore they are not prone
to being leached. Although these ions are held in the colloids, they are differentially held depending
on the concentration of the different groups and the concentration of the charge. They can actually
displace each other from the attraction sites and the displacement depends on the concentration of the

43
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

charge around the cat ion and the number of ions of a given substance in solution. The strength of the
charge depends on the valency of the ion and the size of the ion. Generally smaller ions have a more
concentrated charge and also those ions with a higher valency like Al3+ and Fe3+ have stronger
attractions than those with lower valencies like mono-valencies (+) and di-valencies (2+). Those ions
with a higher charge are going to replace those cat ions with a lower charge from the attractions sites
into the soil solution and those ions with a higher number in solution will displace those ions which
are lower in number from the active sites into solution. For the most common cations, the
replaceability series is usually Al > Ca > Mg > K > Na. This ability to interchange on the soil surfaces
is called cat ion exchange (CE). Cat ion exchange can be defined as the interchange between a cat ion
in solution and another cat ion on the surface of any surface active material (Foth, 1984)

The sum total of all exchangeable cat ions that are adsorbed onto the soil expressed in
milliequivalencies per 100g of oven dry soil is referred to as Cat ion Exchange Capacity. A
milliequivalency is equivalent to one gram of hydrogen or it is the amount of any ion that will
combine or displace one milligram of hydrogen (Buckingham and Brady, 2003). CEC can also be
referred to as an expression of the negative charge per unit mass of soil (White, 1997).

There are two types of cat ions that can be exchanged in the soil; the basic and the acidic groups.
These ions have opposing effects. The acidic ions consists of H+, Al3+, Mn2+ and to some extend
NH4+. The other group is com[posed of such ions as Ca2+, Na+, Mg2+, NH4+ and K+. these bases are
very important in the general nutrition of plants and they are also important soil ameliorants
preventing excessive acidity whose effects are going to be discussed in this section, they act as liming
materials with this respect. The content of these ions can be expressed as percentage base saturation
which is given by the following formula;

Percentage base saturation = Ca2+ + Mg2+ + Na+ + K+ * 100

CEC

In other words the exchangeable bases are expressed as a fraction of all the cat ions that are in the soil,
the higher the figure the better. Percentage base saturation can also be referred to as Base Exchange
Capacity (Wild, 1996).

Importance of CEC in agriculture

 Soils with a higher CEC tend to hold nutrients against leaching, soils with higher CECs
therefore have a natural reserve of nutrients.
 In soils with a higher CEC, fertilizers can all be applied preplanting without any fear of the
nutrients being leached as in sands, this is going to save a lot on labour requirements.
 There is however a negative impact of high CEC, any chemical that is applied to that soil is
likely to have its residual effects felt easily. This can be a disadvantage if herbicides are
applied and they persist in the soil. This is going to affect the rotation that a farmer is going to
follow especially if the crops that will be following each other are from different families.
Persistence of chemicals can only be an advantage in the case of fertilizers, fungicides and
herbicides if the persistence is going to span the length of the life of the commercial plant. If
the CEC gets very high, applied chemicals may be very strongly bound to the soil particles and
they may not be available for plant uptake, if they are herbicides the weeds are not going to
take up the chemical, fertilizers will not be availed to the plants of interest, in short the efficacy
of applied chemicals is going to be reduced. It will follow that the farmer will have to increase
the rates applied. In general, higher rates of herbicides are used in clay soils. The case is

44
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

slightly different with fertilizers because the higher concentration of other cat ions will ensure
that the cat ions of importance are going to be released into the soil due to cat ion exchange.
 Linked to the above discussion, the response to a slight increase in an added chemical is felt,
more in lighter soils of lower CECs than in soils with higher CEC which tend to trap a lot of
the added ions.

Clay minerals

Mineralogical organization of clays

All clays are made up of one or all of the two basic structures. The two basic structures are the silicon
tetrahedral units and the aluminum octahedral units.
The silicon tetrahedral units are made up a silicon ion which is surrounded by four closely packed
oxygen ions which are equally spaced in all dimensions, i.e. they are in four dimension arrangement.

The octahedral layer is made up of an aluminium ion which is surrounded by six hydroxyl ions at the
centre. It is the arrangements of these two basic units in different formats that will give the different
types of clays that are there as shall be discussed later. All clays are formed out of the chemical and
physical alterations of feldspars, amphiboles’ and pyroxenes followed by their decompositions and
subsequent recrystallisation.

Clay mineral types

There are two main groups of clays, the silicate clays and the hydrous oxide clays with the former
being of the most important significance. The silicate group can be divided into three realizable
groups; kaolinite group, montmorillonite group, the mica group.

Kaolinite Group
As the name implies, kaolinite represents the group but the group also includes other clays like
anauxite, halloysite and dickite. The general structure of these clays is that they are made up of one
layer of aluminium octahedral layer which is joined to one layer of the silicone tetrahedrons, hence
these clays are referred to as the 1:1 clays. The layers are tightly fixed by the covalent bondings that
are attained through mutual oxygen atoms shared between the layers. Adjacent 1:1 units are held
together by hydrogen bonds which are sometimes referred to as the van de Waals forces. These are
just weak forces but they still manage to hold the adjacent units together. These clays are non-
expanding as the other groups as you shall notice in the ensuing discussion. This is because cat ions
and water cannot enter the area in between the layers; as such the effective surface area of these clays
is restricted to the area outside the layers. This results in these clays having a low, actually the lowest
CEC of all the three groups of the silicate clays which ranges between 3 to 15 meq/100g of soil. The
hydrogen bonds will allow these clays to form very large particles when compared with the other
types of clays in the silicate group. Since cat ions and water cannot enter the region between layers,
these soils have very low; plasticity, cohesion, shrinkage, and swelling properties.

Montmorillonite Group
This group is sometimes referred to as the smectite group. Montmorillonite is the best representative
of the group which also includes nontronite and saponite. The basic structure is made up of two
tetrahedral silicate layers that sandwhich an octahedral aluminium layer in between them. As such
these soils are sometimes referred to as 2:1 clays the silicon tetrahedral layers and the aluminium
octahedral layer are held together by covalent bonds in mutually shared oxygen atoms. Adjacent 2:1
units are held together by share oxygen linkages as well. Unlike the kaolinite group, these clays have
hydrogen bonding and as such they cannot form large particles. In between the structural units, there
are negative charges which act as adsorptive systems. These are formed as a result of isomorphous

45
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

substitution of Al3+ by Mg2+. This results in these soils attaining a very high CEC since the negative
sites can attract a lot of cat ions. The group has the highest CEC in the silicate clays which ranges
from80-150meq/100g of soil. Cat ions and water are able to move in the region between the layers
within the basic units and the areas between the basic units and such there are both internal and
external adsorptive areas. The entrance of water between the layers allows the soils to expand when
they absorb water when it rains or when the soils are irrigated. When they loose this water, these soils
are going to shrink. When they shrink they tend to crack, this happens usually in the dry season and
this may help in the aeration of the soils. The water in between the layers will also result in the
lubrication of the interlayer region and the different layers can easily slide over each other hence
another characteristic of these soils is that of slipperiness when they are wet. The presence of water
together with cations makes these soils plastic and hence sticky when wet. These soils are very
difficult to work when they are wet as they result in the skidding of farm implements and the sticking
effect can jam implements as well. These soils are common in low lying waterlogged areas and in
areas where the soils originate from basalt rocks as the soils in Chisumbanje. Montmorillonite is
formed under mild weathering conditions (slightly acidic to alkaline) coupled with relative abundance
of magnesium and absence of excess leaching. It may also be formed when illite looses the potassium
that it contains.

The Mica Group

This is also known as the illite group. Like the montmorillonite group, this group is also a 2:1 clay
whose basic unit is made up of two silicon tetrahedral layer that are sandwiching an aluminium
octahedral layer. However in the interlayer region, there is a substitution of aluminium ions largely by
potassium ions. The potassium plays a stabilizing role to the layers’ interface region of the structural
units. As a result of this stabilizing effect, illite clays are less expansive when compared with
montmorillonite but they are better than those of the kaolinite group. Infact all the properties of this
group are intermediate those of the montmorillonite and the kaolinite groups. The CEC of this group
ranges between 20 and 40meq/100g of soil. When the potassium levels of the soil falls, the illite soils
can release the potassium ions that are in their interlayer regions, doing so until the an equilibrium
develops between the reserves and the K+ in the soil. Usually this point is never reached and there is
continued release of the ions for as long as the soil is moist and there is uptake of the potassium ions.
Sometimes the release rate of the ions may not be sufficient to meet the demand for the potassium ions
and this situation is worsened by the fact that most plants can luxuriantly take up potassium; that is to
say they can take up the nutrient beyond their nutritional needs. In this case there is a need to have an
external addition of potassium in fertilizers to meet this demand. Otherwise in Zimbabwe, generally,
the amounts of potassium in the soil would have been sufficient to meet the need of plants. In short
these clays have an inherent richness of potassium ions. It is important to make a point here that the
potassium ions in the interlayer region are deemed as non-exchangeable unlike those that are found on
the surface of the clays which have a ready access to the outside environment and are therefore readily
exchanged (Wild, 1996).

Illite is formed directly from mica or from potash feldspars by recrystalisation or it may be formed
when montmorillonite is invaded by potassium ions.

Hydrous clays
These clays do not posses neither the tetrahedral not the octahedral layers, rather they are made up of
hydrated oxides of different elements. In other words the oxides of the respective elements are found
in association with water molecules. Representatives of oxide clays include gibbisite (Al(OH)3) and
hematite (Fe2O3). These soils are in the advanced stages of weathering are usually found in humid
tropics. They have very stable aggregates, they have low degrees of plasticity and shrinkage. Their
stability is even more when they occur together with kaolite since kaolinite has a partial negative

46
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

charge and the oxide clays have a partial positive charge at times. This attraction is so stable that the
aggregates may behave like sands even though they are 100% clay. Infiltration of water into these
soils is very rapid and because of their stability they resist erosion. These soils are also easily tilled
(Foth, 1984).

Nutrient Uptake Mechanism

Nutrients are taken up through an active mechanism; the process is regarded as active because there is
a requirement for energy. This is unlike osmosis which is passive with water simply moving down a
water potential gradient without a need for energy. The energy is required because nutrients will be
pushed against a concentration gradient since normally nutrients are more concentrated within the
plant roots rather than outside.

Nutrients in soil solution are moved from the soil towards the roots passively; the movement being
driven by the uptake of water which is chiefly driven by the transpiratory pull of water. This
movement is there diffusion in essence and the nutrients will be moving by mass flow. The passive
movement will change once the nutrients reach the root surface. At the root surface, there are receptor
sites that recognize specific nutrients or nutrient compounds. Nutrient ions are going to be attached to
these receptor sites. Linked to the receptor sites there are carrier molecules which can traverse the
cell wall and the cell membrane of the plant carrying the nutrient. The nutrient molecule alone would
not be able to pass through the membrane without the aid of the carrier molecule. It is believed that the
attachment of the nutrient molecule to the carrier molecule and its subsequent movement requires
energy. The energy is obtained from energy molecules especially adenosine tri-phosphate (ATP)
which is produced during photosynthesis and respiration. This is the reason why anything that stops
respiration and photosynthesis will also stop the uptake of nutrients. Waterlogging drives oxygen out
of the soil, the oxygen is needed for respiration, plants that are growing in waterlogged environments
initially manifest deficiency of phosphorous since this nutrient has very limited movement in the soil
hence its uptake is quickly manifested. When the carrier molecule reaches inside the cells (vacuoles)
of the plant roots, it releases the nutrient it would be carrying and it is recycled back to the root surface
to be attached to another nutrient molecule.

Summary

The active uptake process of nutrients; requires energy, moves nutrients against a concentration
gradient (vacuolar sap is many times more concentrated than the soil solution), makes use of carrier
molecule which move specific molecules. So specific are these carriers that they are able to
distinguish between even closely related molecules like HPO42- and H2PO4- (Foth, 1984) any factors
that affect metabolism will also affect nutrient uptake and these include; supply of substrate from
photosynthesis, temperature and soil oxygen which may be affected by soil compaction. Soil moisture
will affect the movement of nutrients towards the root surface and this rate can also be affected by the
rate of transpiration. I am sure from you crop physiology lectures you now know the parameters which
affect the transpiration rate in plants. The concentration of roots is usually directly related to the
uptake rate of nutrients and this parameter is very important in the uptake of phosphorous.

Activities
 What are the importances of CEC in agriculture?
 Identify the sources of negative charges of a colloid

References

47
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Bear, F. E. (1964). Ed. Chemistry of the Soil. Oxford and IBH Publishing, India.
Foth, H. D. (1984). Fundamentals of Soil Science. John Wiley and Sons, USA.
Wild, A. (1996). Soil and the Environment. Cambridge University Press, UK.
White, R.E. (1997). Principles and Practice of Soil Science. Blackwell Publishers, USA

SOIL ACIDITY, SALINITY AND ALKALINITY

Introduction

Soil acidity plays an important role in crop production. The acidity of a soil is determined by the
dominant nutrients in the soil. So it is important to go through the previous be studying this unit.

By the end of this section one should be able to;

 explain the causes or the sources of acidity
 explain the effects of acidity on plant growth
 explain the correction of the acidity through the chemistry of liming
 describe liming with respect to; types of lime, rates of application, times and methods of
application, measurement of lime requirement and residual values of lime
 give the causes of soil salinity and alkalinity, effects of these conditions on plant growth and
describe and explain the reclamation of these soils
 give the need for soil sampling giving the principles and methods of sampling
 describe soil analysis and the interpretation of results

Soil acidity
Soil acidity is the measure of hydrogen ion (H+) concentration in a given substance. Acidity is
measured on a pH scale which has a scale of 0-14. pH is actually a measure of acidity or alkalinity of a
substance with the later being the measure of the concentration of hydroxyl ions (OH-). pH
measurement is based on the dissociation of water into the hydrogen and hydroxyl ions as shown
below;

H2O……………H+ + OH-

Trace amounts of these ions are found in all aqueous systems and they are found in inverse
proportions. The only exception is when they are found in equal amounts at the neutral point which is
at pH 7. Only pure distilled has a pH of 7. At pHs below 7, there will be more of the hydrogen ions
than the hydroxyl ions and such a substance is termed as being acidic. Some of the substances of
agricultural importance that are found in this category include; ammonium nitrate, ammonium
sulphate, urea, humic acids that are found in humus, carbonic acid which is found in acid rain which is
important in weathering of rocks. If the pH is above 7, such a substance is termed as being basic or
alkaline. Some of the substances of agricultural importance which are found in this category include;
all liming material like calcium oxides, magnesium oxides and the elemental forms of calcium,
magnesium and sodium. Soil pH plays a very significant role in the health of a soil as shall be
discussed later on. Most plants do well in a pH range of 5.5 to 6.5. There are however some extreme
and rare cases that need either higher or lower pHs.

Causes of soil acidity

48
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Leaching of bases like calcium, magnesium and sodium which have a neutralizing effect on
excess levels of hydrogen ions. Once they are leached there is going to be a lot of hydrogen
and aluminium ions in the profile they are moving away from resulting in increased acidity.
 Poor management of soils, his especially applies to the addition of lime materials to crop lands.
In most cases especially in Zimbabwe, farmers rarely add liming material which is important
in increasing pH.
 Acidity may be caused by the parent material of the soil, generally some parental material like
granite are acidic. This is the reason why most sand soils which are common in Zimbabwe
tend to be acidic since they originate from granite. Soils that are formed from limestone rock
on the other hand tend to be alkaline in nature.
 Soils that are very rich in aluminium and manganese ions tend to be hydrolysed by water. The
hydrolysis is accompanied by the release of hydrogen ions as shown in the following
equations;

Al3+ + H2O……………..Al (OH)2+ + H+

Al (OH)2+ + H2O………Al (OH)2+ + H+
Al (OH)2+ + H2O………Al (OH)3 + H+
Al (OH)3 + H2O………..Al (OH)4- + H+
(Ross, 1996)
As can be seen from the above reaction there is going to be a release of a single hydrogen ion
for every aluminium ion which is hydrolised and the hydrogen ions which are released may be
as many as four per ion of aluminium which is completely hydrolysed. The hydrolysis is
however not complete sometimes. The hydrogen ions so produced are going to contribute to
the acidity of the respective soil.
 pH can be contributed by the phenomenon of acid rain. Acid rain is formed when the oxide of
nitrogen, sulphur and carbon react with rain forming their respective acids as illustrated by the
following reactions;

CO2 + H2O…………………………..H2CO3 In this reaction water reacts with carbon dioxide

and a weak carbonic is formed.

SO2 + H2O…………………..H2SO3………………….H2SO4 In this reaction sulphur dioxide

reacts with rain and a weakily acidic precussor of sulphric acid before finally the sulphric acid
itself is formed.

NO2 + H2O……………………..HNO3 Here nitrogen dioxide reacts with rain forming nitric
acid.
All these reactions result in the addition of hydrogen ions which are formed when these acids
are dissolved in water as shown in the example below using carbonic acid;

H2CO3 ……………………….2H+ + CO32- (Wild, 1996)

 The overapplication of acidic fertilizers especially nitrogen and sulphate fertilizers. These
fertilizers can be oxidized in the presence of oxygen to form either nitric acid or sulphate acid or
both.
Some of the notable reactions with this respect include the following;
When urea is dissolved in water, there is going to be formation of carbonic acid and nitric acid
as shown in the following reaction

49
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CO(NH2)2 + 2H2O…………..(NH4)2CO3………………2HNO3 + H2CO3 + 2H2O It is

important to note that this reaction is what takes place when urea is applied in the soil, there is
initially formation of the ammonium ion (NH4+) and this is later converted into the nitrate ion
(NO3-) all these forms of nitrogen can be used by the plant but if they are present in excessive
amounts then acidity is going to set in as nitric acid is already an acid and ammonium
oxidation can yield hydrogen ions. It is important to note at this point that the above reaction
also explains why there is a time lag between the application of urea and the response of the
plant to the applied fertilizer, the above steps have to occur first before the nitrogen in urea is
availed to the plant. The reaction can stop in the absence of moisture and if the urea is not
covered it simply volatilizes into the atmosphere. It also follows that if there is a lot of rain
during the application of urea, there may be no need to cover the urea since the above reaction
will occur.

If the urea is applied in excess amounts, the ammonium ions that are formed in the initial
stages of the reaction may also be oxidized by oxygen and there is going to be the release of
hydrogen ions; NH4+ + 3O…………….NO-2 + H+ + H2O
Another important acidic fertilizer is ammonium sulphate (NH4)2SO4, if present in excessive
amounts, the fertilizer can be oxidized to form nitric and sulphric acid as shown in the
following equation;

(NH4)2SO4 + 8O……………………2HNO3 + H2SO4 + 2H2O

Sometimes you would hear some farmers especially the small scale communal farmers saying
the use of fertilizer is detrimental to their fields, “It kills the field” they normally say, to an
extend this may be true but only if there is misuse and poor management of the lands. It is
highly recommended that farmers take their soils for analysis periodically so that the nutrient
status and the pH can be corrected as shall be discussed later in this section.
 Microbial activity on organic matter can help increase the acidity since the decomposition process
can also release organic acids. Although these acids are very weak, their effect can help in the
increase in hydrogen concentration. An example is when simple sugars like glucose are oxidized,
they are going to form an acid like pyruvic acid as shown below;
C6H12O6 + 2O…………………….2CH3COCOOH + 2H2O
An increase in soil organic matter content will increase the amount of carboxylic and phenolic
groups which tend to release hydrogen ions when in solution thereby increasing acidity.
 An interesting cause of acidity occurs in swampy areas where elemental sulphur can be found.
Sulphur reducing bacteria found in swampy areas are going to reduce the sulphur through the
addition of hydrogen to form hydrogen sulphide (H2S). This is a gas which smells like rotten eggs
if one gets to a swampy or marshy area. This reaction happens in times when the area is inundated
with water. When the area gets drained or the area dries out in the dry season, these waterlogged
areas become aerated and there is oxidization of the hydrogen sulphide to form sulphric acid as
shown in the reaction below;
H2S + 4O…………………..H2SO4
Sometimes the sulphur is found bound to iron in compounds known as iron pyrites (FeS2).
These iron pyrites can still be oxidized like the hydrogen sulphide gas to form sulphric acid as
shown below;
FeS2 + 3O + H2O………………..H2SO4 + Fe (Foth, 1984)

It will be too much to expect you to memorise all the equations given above but their inclusion will
help you to understand the processes that take place and your answer will be more credible if you can
attempt to put some of them though it will be acceptable if you can express your understanding in
words.

50
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

So you have learnt about the cause of acidity in the soil, but of what significance is soil acidity to the
farmer. In the following section we discuss the significance of acidity to crop production.

Effects of acidity on crop growth

Acidity affects the availability of nutrients in a big way, some nutrients become less available and
some become available in excessive amounts that may be toxic to plants. There are other plants which
can thrive on acidic soils, some of them include; tea, rice, cranberry, blueberry, rhododendron and
azalea (Millar and Turk, 1998).
 Calcium and magnesium become more soluble in acidic soils and they are easily leached away
beyond the reach of plant roots.
 Nitrogen, sulphur and phosphorus become immobilized especially if the soil is rich in organic
matter, when immobilized they become tightly held and plants cannot access them for their
use.
 Nitrogen gets taken up in the ammonium form (NH4+). The plant can take this form up but it
still has to be changed to the nitrate form (NO3-) which is the metabolisable form. If the
ammonium ion is present in high amounts it can be toxic to plants and can lead to the death of
especially young seedlings like the seedlings of tobacco where ammonium nitrate is not
recommended for use in seedbeds because the ammonium ions that are released will result in
high seedling mortality.
 Phosphorous ions change in their form depending on the prevailing pH. Some of the forms are
not absorbed by plants. Refer to phosphorous under nutrients.
 At low pH, the concentration of iron, manganese, chrome, copper, nickel and zinc are going to
increase. These elements are referred to as trace elements and they are required by plants at
very low concentrations roughly around 50 parts per million. If they go higher than this
concentration they become toxic to plants which may end up dying or they may have impaired
growth which will reduce yield. The most dangerous ion with this respect is aluminium whose
concentration rises under acidic environments, it can easily displace other important cations
from the active uptake sites preventing them from being taken up by plants. Together with
nickel, the ions are known to impair root growth, affected roots become club shaped which
tends to reduce nutrient and water uptake.
 At low pH molybedenum and phosphorus form insoluble compounds with iron and calcium
and the so formed compound cannot be taken up plants meaning that these nutrients will not be
available to the plant.

Correction of low pH
If the pH is found to be too low, then means have to be found of increasing it. This is almost
always done through the application of lime. Lime can be defined as any material which is basic in
nature and can therefore increase the pH thereby countering or neutralizing the low pH. The
materials used are almost always calcium or magnesium based and some of the materials that can
be used include the following; Calcium oxide (CaO), magnesium oxide (MgO), calcium hydroxide
(Ca(OH)2, magnesium hydroxide Mg(OH)2, calcium carbonate also known as calcite (CaCO3),
magnesium carbonate (MgCO3) and dolomite (CaMg(CO3)2.

Carbonates are derived from naturally occurring limestone, shellfish deposits, and precipitated
carbonates but by far the most common and commercially viable source is limestone rock. In
Zimbabwe the rock is found in abundance in many areas but the most common one is the mine in
the Concession area. Even when finely ground the carbonates both calcium, magnesium, and
dolomitic limes dissociate slowly in solution and they take a long time to effect the necessary
change and this may be up to a year.

51
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The oxides are produced by heating the carbonates, a reaction which produce the respective oxide
and carbon dioxide as show in the reaction below;
CaCO3 + heat………………..CaO + CO2.
CaMg(CO3)2 + heat……………….MgO + CaO + 2CO2.
This form of lime is quick acting when compared with carbonate form. It is sometimes referred to
as quick lime or burnt lime since it is heated in large kilns. Its so caustic that its effects may be
felt in the soil within one or two weeks of application.

The hydroxides are formed through the addition of water to the respective oxides as exemplified
by the addition of water to calcium oxide to form calcium hydroxide;
CaO + H2O……………..Ca (OH)2
These limes are sometimes referred to as the hydrated limes because of the addition of water. Of
the three groups these are the quickest in effecting change in the pH of a soil.

The chemistry of liming

Your syllabus requires that you be able to describe chemistry of liming, in other words how does
liming increase the pH of a soil. This part has to be memorized as it is examinable. An example is
illustrated here using calcium carbonate. The steps may be slightly different for the different forms
of lime but the final product formed is calcium bicarbonate (Ca (HCO3)2) or magnesium
bicarbonate (Mg (HCO3)2). If calcium carbonate is used the following reactions are what will be
occurring in the soil.

Calcium carbonate dissociates in the soil

CaCO3……………..Ca2+ + CO32-

The carbonate ion will react with water forming a hydroxyl ion and a hydrogen carbonate ion;
CO32- + H2O…………………..HCO3- + OH- and,

There is a further reaction where water reacts with the hydrogen carbonate to form carbonic acid
and another hydroxyl ion;
HCO3- + H2O………………….H2CO3 + OH-

The carbonic acid produced is very unstable and it breaks down into its constituent components,
carbon dioxide and water;
H2CO3 ………………………. H2O + CO2
Liming
Commonly soils become acidic with continuous cultivation, to raise the pH, lime, oxides, OH, CO3 of
Ca and Mg is added
Calcium oxide CaO
Is also called unslaked lime, burnt lime or quick lime. It is a white powder manufactured by roasting
calcitic limestone.
The purity of the burned lime depends on the purity of the raw material. It reacts almost immediately
when added to the soil, it should be mixed completely with soil as it rapidly cakes and becomes
ineffective.
Ca(OH)2
Referred to as slaked lime, hydrated or builders lime. It is a white powdery substance, difficult and
unpleasant to handle. It is prepared by hydrating CaO.
CaCO3 and dolomitic lime MgCO3
Is made from mined deposits. Quality depends on impurities

52
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Slags
Are bi-products of industrial reactions. Eg blast furnace slag is a bi-product of the manufacture of pig
iron.
Liming material is selected on its neutralizing value, its degree of fines and its reactivity. Neutralizing
values of all liming materials are determined by comparing them to the neutralizing value of value of
pure CaCO3 (100).

The hydroxyl ions (OH-) that are produced are going to react with the excess hydrogen ions (H+)
that would be causing the low pH thereby neutralizing the soil. As the hydroxyl ions are used up,
more CaCO3 is broken down until either the CaCO3 or the hydrogen ions are finished. Ca2+ or
Mg2+, from the dissociation of the liming material in the initial stages may be used in the
displacement of hydrogen ions on surface areas resulting in either Ca2+ or Mg2+ being dominant on
the surface of exchange sites thereby creating a reserve of bases which are highly needed by the
plant. The presence of hydrogen ions tends to exclude the uptake of these bases and other
important nutrients.

The reactions that would have happened up to this point can be summarized by the following
reaction;
CaCO3 + 2H+……………………….Ca2+ + H2O + CO2

As long as the carbon dioxide (CO2) is present in large amounts, it is going to push the balance of
the reaction towards the production of carbonic acid which may increase the acidity which you
will be trying to lower and therefore the carbon dioxide has to be removed. The carbon dioxide can
react with any of the liming materials in the presence of water to form the corresponding salts;
CaCO3 + H2O + CO2…………………………Ca (HCO3)2

The salt molecules formed are harmless; they can simply dissolve to form the respective ions in
solution. The formation of the salt therefore suppresses carbonic acid activity.

The effect of liming can therefore be summarized by the following reactions depending on the
material that is used;
CaCO3 + H2O + CO2………………………………Ca (HCO3)2
Ca (OH)2 + H2O + 2CO2………………………. Ca (HCO3)2
CaO + H2O + 2CO2………………………. Ca (HCO3)2
MgCO3 + H2O + CO2………………………. Mg (HCO3)2
Mg (OH)2 + H2O + 2CO2………………………. Mg (HCO3)2
MgO + H2O + 2CO2………………………. Mg (HCO3)2 (Ross, 1996)

Measurement of lime requirement

The amount of lime that is required to bring about a desired pH change is dependent on a number
of parameters which may include the following;
 The change in pH required, the more it should be increased the more the lime that should be used.
Generally for a 0.1 change in pH one would require 100kg of liming material.
 The chemical composition of the liming material, as already alluded to some liming materials are
quick acting when compared with the others, those that are quickly used may be applied in smaller
amounts since they quickly change the chemical nature of the soil.
 The fineness and distribution of the liming material, finely ground lime has a larger surface area
and therefore it quickly reacts with hydrogen ions and the effect is increased when the lime is well
distributed in the soil profile whose pH is to be increased. This is the reason why lime has to be
incorporated by ploughing or by discing it into the soil.
53
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 The buffer capacity of the soil, a buffer is a material which maintains the pH of a substance around
a certain region, in other words a well buffered substance has very low variations of pH. A buffer
can act as either an acid or it can act as an acid depending on the acidity of the environment in
which it finds itself. Normally buffers are weak acids. If a soil is well buffered it will require less
lime.

The lime requirement can be measured through one of several ways and some of them are discussed
below.

I. Buffer solution method- the soil is shaken a buffer solution and the pH decrease is measured.
The decrease is related to the amount of acidity which will determine the amount of lime to be
added.
II. Incubation method- the soil is incubated with varying amounts of lime in a moist
environment. After the incubation the pH of the different samples is going to be measured and
the amount which would have adjusted the pH to the required level is the one that can be used.
The information can also be plotted on a graph with lime amount on the x-axis and the
resultant pH on the y-axis and this can be used in the future.
III. The Hutchison-Mclenan method-this method was developed by two scientists whom it was
named after. It is one of the titration methods which can be used to determine lime
requirement. The soil is shaken with a known amount of calcium carbonate. The excess is
going to be titrated with a standard acid, one whose concentration is known so that the amount
of acid used in the netraisation can be used to determine the amount of lime that is required to
bring a certain change in the pH can be determined. This method is largely determined in the
laboratory, there may be a need to support these findings with field trials since field conditions
may vary a little with lab findings.
IV. Dye method- is used to determine the pH so that reference can be made to past calibrations so
that the amount of lime can be determined usually from pre-prepared graphs.
V. Electrometric methods- these make use of pH meters again to measure the pH so that the
amount of lime to be added can be determined from prepared graphs.

Methods of lime application

Whatever method that is used to apply the lime, it should be evenly distributed and it should be
thoroughly mixed within the soil profile where there is a need of pH adjustiment and normally this up
to the ploughing depth. Distribution can be achieved through the use of lime boxes or lime spreaders
which spread the lime evenly on the soil surface. This should be followed by discing or ploughing in
of the lime using harrows or ploughs. This is the point where conservation agriculture has some
hinderance since there will be no cultivation to incorporate the lime. Lime can be applied any time of
the year though this depends on the type of rotation, system of farming and the form of lime that is
applied.

Methods to lower pH

Residual value of lime

The residual effects of lime in a soil can be felt for about 5-8years though this it dependent on a
number of factors. If lime used is in the carbonate form which is a slow release form then the residual
effects can be felt for a long period of time, if however, the form used is the hydrated form then it is
quickly used up because it is quickly released and hence the residual effects are going to be felt for a
few years maybe 2-5years. The amount of residual lime will also depend on the amount of nitrogenous
and other acidic fertilizers that are added to the soil. If this amount is high it means that the need for
the control of acidity will be high and this will result in quick depletion of the lime content in the soil.

54
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The climate of the land also plays an important part in determining the level of residues in the soil. If
the rainfall amount is high then the leaching rate of bases will also be high and therefore the residual
effects of lime will not be felt for long.
Saline and Sodic Soils
Soils may become naturally alkaline though weathering, lack of leaching and poor drainage.
Exchangeable Sodium Percentage. – used to distinguish between sodic and non-sodic soils
ESP ==== Exch Na * 100
CEC
Soils with an ESP of greater than 9 are classified as sodic in Zimbabwe, they have a very bad
structure, poor drainage and hard to work.
Sodium Adoption Ratio (SAR) is used to separate out sodic soils.
SAR === [Na+] cmolesckg-1
√0.5([Ca2+ ] + [Mg2+])
This shows the relative concentration of Na to Ca, and Mg
Electrical conductivity
Electrical conductivity (EC) is measured in deciSiemens/m. The greater the quantity of salts the more
the solution conducts the electrical current
2.8.5 Saline Soils
These soils contain mainly Ca and Mg salts. There are enough salts in the soil to seriously impair plant
growth. The soil solution has a high osmotic potential and cause plasmolysis of plant cells. Salts are
brought up to the surface by evaporating water, and they are called white alkali soils. Salinity results
in soil flocculation. Saline-sodic soils have appreciable quantities of Ca/Mg and enough Na to affect
plants. These soils are more difficult to work.
2.8.6 Sodic soils
Have very high Na concentrations. The soil pH is greater than 8.5 due to the hydrolysis of water by Na
Clay-Na + H2O ----------- Clay- H+ + Na+ + OH
Conditions results in toxicity of nutrients ie too much availability of other nutrients
The increase in pH cause fixation of many plant nutrients. The sodium destabilizes clay, causing its
dispersion. Infiltration of rainfall is therefore reduced due to the dispersed clay which clogs pores.
Sodic soils develop from granite or karoo formations in drier areas as these are rich in Na minerals.
Sodium has the opposite effects of salinity on soils, that is, the forces that bind clay particles together
get disrupted when too many large sodium ions come between them.

Soil sampling

Sampling is the selection of a small fraction of the whole so that the characteristics of the whole can
be inferred from the fraction. Sampling is necessary because it reduces costs, space and time required
to evaluate a certain characteristic about the whole population. In soil sampling it is impossible to take
soil from the whole field so that the characteristics of that soil may be ascertained and as such only a
fraction of the soil must be taken so that it can be taken to labs that do the analysis and these may be
several hundreds of kilometers away. As such soil sampling should be governed by certain principles
which shall be discussed here and these help to ensure that the results that you are going to get are
going to reflect the properties of the whole field. However; before we look at these principles let us
look at the need for soil sampling which may sound more or less the same like the introductory
discussion.
Need for soil sampling
 It breaks bulkiness of the soil so that smaller amounts of soil can be used to determine
characteristics about the whole field from which the soil was taken as such there is saving on space
for storage, transportation and also time and monetary resources during the handling.

55
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 The sampled soil is used to determine the nutrient content of the soil so that the proper fertilizers
can be applied at the right rates.
 The sample is also used to determine the soil pH so that the right material can be used to adjust the
pH, a farmer may use lime if the pH is too low or they may use acidic fertilizers if the pH is too
high and it has to be brought down.
 The sample may also be used to determine the physical characteristics of the field from which it
has been taken and these may include the following; soil structure, texture, bulk density, particle
density (these may help in the determination of water holding capacity, drainage and aeration) and
colour among many other parameters.

Principles of soil sampling

 A soil sample should be representative of the whole field from which it was taken. As such the
sample should be a composite one which is made up of several subsamples that would have been
taken around the whole field not just from one corner.
 A composite sample must be taken from each field for which recommendations are required.
Significant changes in a field must be sampled separately. By significant, it means the changes
affect a large piece of the land and some of the changes include; changes in soil colour, texture,
drainage, slope and vegetation among some of the parameters. If there is a large variation for areas
in the same field then the samples will have to be taken separately. Minor variations in the field
will have to be ignored since their impact is very negligible. Uncharacteristic abnormalities have
to be avoided and examples of such areas include; anthills, animal holes, ridges, compost sites,
areas where fertilizer or manure was heaped and drain sites among many other areas which are not
typical of large chunks of the field.
 Each composite sample must be made up of at least ten subsamples no matter how small the field
for the determination is. The number can only be increased not reduced, increase is more
appropriate in larger pieces of land and in land which has just been ploughed. The subsamples
must be taken randomly across the field without any bias as this may give false results. There are
some methods that may be used to avoid this and these include completely random sampling
across the field and zig-zagging across connecting to corners moving diagonally and this must be
done for all the corners.
 The sample must represent the full depth of the soil. All corrections that may be required are going
to be done in the rooting zone and as such the soil has to be sampled at least to a depth of about
30cm which is the depth of the top soil.
 When samples are being taken for nitrogen determination, it will be best if the samples are going
to be taken after the rains have stopped. If determinations are made during the rains, there may be
further loss of the nitrogen during the season and whatever corrections were made may be lower
than the actual requirement.
 If samples are required for pH determination, then the samples can be taken anytime since the
hydrogen ions which cause or determine acidity are not easily leached since they are tightly held to
soil particles.

Methods of soil sampling

Spade method
Dig a V-shaped hole to plough depth which is about 30cm. Do not brush away litter or loose soil on
the surface. Cut a uniform slice of soil along one arm of the V shape taking proper care not loose any

56
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

of the soil especially the surface soil. This is going to be done at each sub-sampling point. This
method only works where the ground is firm, where it is loose like in recently ploughed lands the
method is not really suitable since a lot of the soil from the surface is going to fall back into the hole
and the results may not be reflective of the true properties of the soil in question.

The trowel method

A normal garden trowel is used to gather subsamples in recently ploughed lands. The setback with a
trowel is that the representativeness of the subsample is reduced since its effectiveness is reduced with
increasing depth. To try and counter this you are supposed to increase the number of subsamples that
are taken per piece of land over the recommended 10. The method therefore becomes rather tedious
and even so the results will not be really representative, it will require someone good at the technique.

Auger method
This is the most rapid method and it is more efficient in taking samples that are more representative of
the soil depth. The method can be used by almost anyone who is able bodied since room for mistakes
is reduced. The equipment has a screwing effect and as it is screwed into the soil it will be augering
soil into a collection bucket packing it tightly as it does so. When the bucket is full the soil is
discarded in a collection container as a subsample. This method cannot be used in very wet lands since
the soil can get sticky therby preventing the proper augering of the soil. In soils which are very loose,
the soil will not remain in the bucket, it will be more suitable to use the trowel method. In very dry
lands it will be almost impossible to screw the auger into the soil, it will be more suitable to use the
spade method.

The augers come in different sizes which vary with the diameter of the bucket.

Preparing the samples for analysis

When all the subsamples have been collected in one container, there is going to be some thorough
mixing of all the subsamples to ensure the representativeness of the composite sample which is made
up of all the subsamples. The composite sample is allowed to dry if it has some moisture in it and the
drying should be natural, artificial drying is known to destroy some components which are of
significance. When the soil is ready, about a kilogram of soil is drawn from the composite sample and
packed in a proper package which can have the following information labeled on it; name of farmer,
name of field, date sampled and purpose of sampling. This prepared pack is send for analysis at proper
labs which may be private or public. The private laboratories include those at fertilizer companies like
ZFC and Windmill and also at the Zimbabwe Sugar Association Experimentation Station in Chiredzi.
There is also the quasi-government Tobacco Research Board Kutsaga Station and the soil science and
chemistry sections of institutions of higher learning like universities and polytechnic colleges.
Normally there is a fee to be paid for the service rendered. Fertilizer companies normally do pH
determinations for free but they will charge you for nutrient content determinations and the
recommendations. It is important to note here that fertilizer companies normally make
recommendations using their product which to me becomes easier for the farmer rather than being
given numbers for nutrient requirements which may be difficult for the farmer to formulate on farm.

Precautions
 All implements and all containers that are going to be used for the exercise must be clean and
free from chemicals that may alter the actual chemical contents of the soil e.g. used fertilizer
sacks should not be used.
 Avoid the peripheries of the field as these normally may have too much or too little fertilizer
applied to them.

57
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Summary
For maximum crop production, soil pH should be neutral. Acidic and saline soils are not good for crop
production. Such soil should be rehabilitated if crop production is to be done. Soil sampling and
analysis should be done correctly so as to get correct results.

Activities

 How do you think crop production can be affected by salination?

 What are the possible causes of saline soils?

References
Buckman, H.O. and Brady, N.C. (2003). The Nature and Properties of Soils. Macmillan Publishers,
London.
Ross, A. (1996). The Nature of Soils. Lakeland University Press, USA.
White, R.E. (1997). Principles and Practice of Soil Science. Blackwell Publishers, USA.
Wild, A. (1996). Soil and the Environment. Cambridge University Press, UK.

CHAPTER 6

ORGANIC MATTER

Introduction
The use of organic matter as fertilizer is popular to resource poor small holder farmers in Zimbabwe.
Farmers in most communal areas cannot afford artificial fertilizers. Organic fertilizer does not only
provide nutrients to the soil but also improve soil structure, microbial activities and soil temperature.
Therefore, organic inputs are proving to be vital in boosting agricultural production in Zimbabwe. As
future farmers, it is crucial to understand the role and management of organic matter to the best of our
advantage. We should develop agronomic practices that aid in maintaining organic matter at
reasonable levels in the soil.

Objectives
By the end of this unit students should be able to:

 Define and state the importance of organic matter.

 identify constituents of soil organic matter
 describe the role of soil organisms in organic matter decomposition
 identify organisms involved in different stages of the decomposition process
 Describe factors affecting soil organic matter levels.
 Discuss the benefits of soil organic matter.
 describe the carbon: nitrogen (C:N) ratio
 describe how the C:N ratio affects the rate of decomposition
 Describe the carbon cycle and the C: N ratio.
 Explain the importance of C: N ratio.

Definition of organic matter

58
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Organic matter is material that has come from a once living organism, is capable of decaying, or
product of decaying, or is composed of organic compounds. Organic matter originates from two
sources: primary and secondary sources. The primary sources are plant residues (roots ,stems and
leaves ).Generally, the moisture content of organic matter from plants is about 75% leaving a dry
matter of 25%.The dry matter is composed of carbon, oxygen, hydrogen, and other elements like
nitrogen, phosphorous, sulphur, potassium and magnesium. Decomposition of organic matter results
in the formation of humus. Humus is the last substance of decomposition. Another source of organic
matter is animal bodies, products and manure. The decomposition of animal remains provides a
secondary source of organic matter. They are referred to as secondary because animals consume plant
tissues .The secondary sources include bodies of soil organisms such as bacterial, fungi, beetles, ants
etc. and scavenging and manure from animals .Animal manure is widely used as organic fertilizer by
many smallholder farmers. It varies in composition depending on the type of animal, age, health and
feed given to the animal.

Soil organic matter

Soil organic matter (SOM) is the organic matter component of soil, consisting of plant and animal
residues at various stages of decomposition, cells and tissues of soil organisms, and substances
synthesized by soil organisms. SOM exerts numerous positive effects on soil physical and chemical
properties, as well as the soil’s capacity to provide regulatory ecosystem services. Particularly, the
presence of SOM is regarded as being critical for soil function and soil quality.
The positive impacts of SOM result from a number of complex, interactive edaphic factors; a non-
exhaustive list of SOM's effects on soil functioning includes improvements related to soil structure,
aggregation, water retention, soil biodiversity, absorption and retention of pollutants, buffering
capacity, and the cycling and storage of plant nutrients. SOM increases soil fertility by providing
cation exchange sites and acting as reserve of plant nutrients, especially nitrogen (N), phosphorus (P),
and sulfur (S), along with micronutrients, which are slowly released upon SOM mineralization. As
such, there is a significant correlation between SOM content and soil fertility.
SOM also acts as a major sink and source of soil carbon (C). Although the C content of SOM is
known to vary considerably, SOM is typically estimated to contain 58% C, and the terms 'soil organic
carbon' (SOC) and SOM are often used interchangeably, with measured SOC content often serving as
a proxy for SOM. Soil represents one of the largest C sinks on the planet and plays a major role in the
global carbon cycle. Therefore, SOM/SOC dynamics and the capacity of soils to provide the
ecosystem service of carbon sequestration through SOM management have received considerable
attention in recent years.
The concentration of SOM in soils generally ranges from 1% to 6% of the total topsoil mass for most
upland soils. Soils whose upper horizons consist of less than 1% organic matter are mostly limited to
desert areas, while the SOM content of soils in low-lying, wet areas can be as high as 90%. Soils
containing 12-18% SOC are generally classified as organic soils.
It can be divided into three general pools: living biomass of microorganisms, fresh and partially
decomposed residues, and humus: the well-decomposed organic material. Surface plant litter is
generally not included as part of soil organic matter.
Sources of soil organic matter
The primary source of organic matter contained in soil is vegetal. In forest or prairies, as well as
agricultural fields, dead plants are transformed by different kinds of living organisms. This process
involves several steps, the first being mostly mechanical, and becoming more chemical as it

59
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

progresses. The small living beings that work on that decomposition chain are themselves part of the
soil organic matter, and form a food web of organisms that prey upon each other and are preyed upon.
There are also other animals that consume living vegetal material, whose residues are passed to the
soil. The products from the living organism’s metabolism are the secondary sources of soil organic
matter that also includes the dead corpses of these organisms. Some animals, like earthworms, ants
and centipedes contribute to both vertical and horizontal translocation of organic material.
Additional sources of soil organic matter include plant root exudates and charcoal.

Factors affecting soil organic matter

1. Climate
2. Natural vegetation
3. Texture
4. Drainage
5. Cropping and Tillage
6. Crop rotations, residues and plant nutrients.

1. Climate: Temperature and rainfall exert a dominant influence on the amounts of N and organic
matter found in soils.
a) Temperature: The organic matter and N content of comparable soils tend to increase if one moves
from warmer to cooler areas. The decomposition of organic matter is accelerated in warm climates as
compared to cooler climates. For each
matter and N increases by two to three times.
b) Rainfall: There is an increase in organic matter with an increase in rainfall.
Under comparable conditions, the N and organic matter increase as the effective moisture becomes
greater.
2. Natural Vegetation: The total organic matter is higher in soils developed under grasslands than
those under forests.
3. Texture: Fine textured soils are generally higher in organic matter than coarse textured soils.
4. Drainage: Poorly drained soils because of their high moisture content and relatively poor aeration
are much higher in organic matter and N than well drained soils.
5. Cropping and Tillage: The cropped lands have much low N and organic matter than comparable
virgin soils. Modern conservation tillage practices helps to maintain high OM levels as compared to
conventional tillage.
6. Rotations, residues and plant nutrients: Crop rotations of cereals with legumes results in higher
soil organic matter. Higher organic matter levels, preferably where a crop rotation is followed.

Composition of organic residues:

Plant residues contain 75% moisture and 25% dry matter.

This 25% is made up of Carbon (10-12%), Oxygen (9-10%), Hydrogen (1.5-2.5%), N (1-2%) and
mineral matter (1-3%).
Composition of plant tissues:
 Carbohydrates
 Celluloses 20-50%
 Hemicellulose 10-30%
 Starch, Sugar 1-5%
 Proteins 1-15%

60
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Fats, waxes, tannins 1-10%

 Lignin 10-30%
Inorganic residues (mineral matter)
1. Water insoluble
Proteins, Peptides, Nitrogenous Peptones and S containing materials
2. Water soluble
(No3, NH4 compounds)
Soil organic residues
 Non Nitrogenous Carbohydrates (celluloses Hemicellulose, Starch, Sugar etc.)
 Ether solubles (Fats, oils, waxes, resins etc.) Lignins

The organic matter is also classified on the basis of their rate of decomposition
1. Rapidly decomposed: Sugars, starches, proteins etc.
2. Less rapidly decomposed: Hemicelluloses, celluloses etc.
3. Very slowly decomposed: Fats, waxes, resins, Lignins etc.
Decomposition of soil organic matter:
Different organic residues contain different organic compounds. There is great variation in the rate of
decomposition of organic residues. Sugars, starches and simple proteins are very rapidly decomposed.
On the other hand Fats, waxes and Lignins are very slowly decomposed.
Hemicellulose, celluloses and protein are intermediate. Even though the composition may vary the end
products are more or less the same.
The general reactions taking place during decomposition are:
1. Enzymatic oxidation of the bulk with the release of Co2, water, energy and heat
2. Essential elements are released (N, P, S etc.) and immobilized by a series of reactions.
3. Formation of compounds which are resistant to microbial action.

Molecules very resistant to microbial action is formed either through modification of compounds or by
microbial synthesis
Under aerobic conditions the products formed are
Co2, NH4, NO3 , H2PO4, SO4, H2O and essential plant nutrients like Ca, Mg, Fe, Cu, Zn etc.
Under anaerobic conditions
CH4, organic acids like lactic, propionic, butyric, NH4, various amine residues (R-NH2) H2S,
ethylene (CH2=CH2) and humic substances.
A. Decomposition of soluble substances:
When glucose is decomposed under aerobic conditions the reaction is as under:

Under partially oxidized conditions,

hydroxy acids (Citric, lactic etc.) or
Alcohols (ethyl alcohol etc.)
Ammonification – organic N - Polypeptides – Peptides – amonoacids – NH3 or NH4
i) Ammonification: The transformation of organic nitrogenous compounds (amino acids, amides,
ammonium compounds, nitrates etc.) into ammonia is called ammonification. This process occurs as a
result of hydrolytic and oxidative enzymatic reaction under aerobic conditions by heterotrophic
microbes.
ii) Nitrification: The process of conversion of ammonia to nitrites (NO2) and then to nitrate (NO3 -)
is known as nitrification. It is an aerobic process by autotrophic bacteria.
Ammonia Nitrite Nitrate
NH4 + O2 NO2 + 2H+ + H2O + energy
NO2 + O2 NO3 - + energy

61
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

iii) Denitrification: The process, which involves conversion of soil nitrate into gaseous nitrogen or
nitrous oxide, is called Denitrification. Water logging and high pH will increase N loss by
Denitrification.
2. Under anaerobic conditions:
C6H12 O6 (Glucose) - Lactic acid, butyric acid Ethyl alcohol are formed
Protein and other N compounds are converted into elemental N.
B. Decomposition of Insoluble Substances
i) Breakdown of Protein: During the course of decomposition of plant materials, the proteins are first
hydrolysed to a number of intermediate products.
Aminization: The process of conversion of proteins to amino acids.
Ammonification: The process of conversion of amino acids and amides to ammonia.
ii) Breakdown of cellulose: The decomposition of the most abundant carbohydrates. Hydrolysis
oxidation

This reaction proceeds more slowly in acid soils than in neutral and alkaline soils. It is quite rapid in
well aerated soils and comparatively slow in poorly aerated soils.
iii) Breakdown of Hemicellulose: Decompose faster than cellulose and are first hydrolysed to their
components sugars and uronic acids. Sugars are attacked by microbes and are converted to organic
acids, alcohols, carbon dioxide and water.
The uronic acids are broken down to pentose and CO2. The newly synthesized hemicelluloses thus
form a part of the humus.
iv) Breakdown of Starch: It is chemically a glucose polymer and is first hydrolysed to maltose by the
action of amylases. Maltose is next converted to glucose by maltase.
C. Decomposition of ether soluble substances:
Fats =glycerol + fatty acids
Glycerol CO2 + water
D. Decomposition of lignin: Lignin decomposes slowly, much slower than cellulose. Complete
oxidation gives rise to CO2 and H2O.
Sulphur containing organic compounds:
Converted to SO4 -2 + H+ + energy by sulphur oxidizing bacteria.
P containing organic compounds:
Various microorganisms mineralize phospholipids and other organic P compounds in the presence of
phosphates enzymes H2PO4 and HPO4-2 depending on soil PH.
Mineralisation: The biological conversion of organic forms of C, N, P and S to inorganic or mineral
forms is called mineralization.
Immobilization: The conversion of inorganic forms of C, N, P and S by the soil organism into
organic forms is called Immobilization.
Factors affecting decomposition
1. Temperature: Cold periods retard plant growth and organic matter decomposition. Warm summers
may permit plant growth and humus accumulation.
2. Soil moisture: Extremes of both arid and anaerobic conditions reduce plant growth and microbial
decomposition. Near or slightly wetter than field capacity moisture conditions are most favourable for
both processes.
3. Nutrients: Lack of nutrients particularly N slows decomposition.
4. Soil pH: Most of the microbes grow best at pH 6 to 8, but are severely inhibited below pH 4.5 and
above pH 8.5.
5. Soil Texture: Soils higher in clays tend to retain larger amounts of humus.
6. Other Factors: Toxic levels of elements (Al, Mn, B, Se, Cl), excessive soluble salts, shade and
organic phytotoxins in plant materials.
Role of organic matter

62
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

1. Organic matte creates a granular condition of soil which maintains favourable condition of aeration
and permeability.
2. Water holding capacity of soil is increased and surface runoff, erosion etc., are reduced as there is
good infiltration due to the addition of organic matter.
3. Surface mulching with coarse organic matter lowers wind erosion and lowers soil temperatures in
the summer and keeps the soil warmer in winter.
4. Organic matter serves as a source of energy for the microbes and as a reservoir of nutrients that are
essential for plant growth and also hormones, antibiotics.
5. Fresh Organic matter supplies food for earthworms, ants and rodents and makes soil P readily
available in acid soils.
6. Organic acids released from decomposing organic matter help to reduce alkalinity in soils; organic
acids along with released CO2 dissolve minerals and make them more available.
7. Humus (a highly decomposed organic matter) provides a storehouse for the exchangeable and
available cations.
8. It acts as a buffering agent which checks rapid chemical changes in pH and soil reaction.

ESSENTIAL FUNCTIONS PERFORMED BY DIFFERENT MEMBERS OF SOIL

ORGANISMS (BIOTA)

Functions Organisms involved

Maintenance of soil structure Bioturbating invertebrates and plant roots, mycorrhizae and some other
micro-organisms
Regulation of soil Most Bioturbating invertebrates and plant roots
hydrological processes
Gas exchange and carbon Mostly micro-organisms and plant roots, some C protected in large compact
sequestration (accumulation biogenic invertebrate aggregates
in soil)
Soil detoxification Mostly micro-organisms
Nutrient cycling Mostly micro-organisms and plant roots, some soil- and litter-feeding
invertebrates
Decomposition of organic Various saprophytic and litter-feeding invertebrates (detritivores), fungi,
matter bacteria, actinomycetes and other micro-organisms

Suppression of pests, Plants, mycorrhizae and other fungi, nematodes, bacteria and various other
parasites and diseases micro-organisms, collembola, earthworms, various predators

Sources of food and Plant roots, various insects (crickets, beetle larvae, ants, termites),
medicines earthworms, vertebrates, micro-organisms and their by-products

Symbiotic and asymbiotic Rhizobia, mycorrhizae, actinomycetes, diazotrophic bacteria and various
relationships with plants and other rhizosphere micro-organisms, ants
their roots

Plant growth control (positive Direct effects: plant roots, rhizobia, mycorrhizae, actinomycetes, pathogens,
and negative) phytoparasitic nematodes, rhizophagous insects, plant-growth promoting
rhizosphere micro-organisms, biocontrol agents Indirect effects: most soil
biota

63
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Humus

As vegetal material undergoes decomposition, some microbial resistant compounds are formed. These
include modified lignins, oils, fats and waxes. Secondly, some new compounds are synthesized, like
polysaccharides and polyuronids. These materials form the basis for humus. New reactions take place
between these compounds and some proteins and other nitrogen containing products, incorporating
thus nitrogen and avoiding its mineralization. Other nutrients are also protected in this way from
mineralization.

Humic substances classification

There is a classification into three groups, based on solubility in acids and alkalis, and also related to
stability.
 Fulvic acid is the group which contains the materials that have the lowest molecular weight, and
are soluble in acids and alkali, and susceptible to microbial attack.
 Humic acid group contains the intermediate materials, with medium molecular weight, soluble in
alkali, but insoluble in acid, and intermediate resistance to microbial attack.
 Humin is the generic name for the materials with highest molecular weight, that are darkest in
colour, insoluble in acid and alkali, and with the most resistance to microbial attack.

Functions of humus:

 improved fertilizer efficiency;

 long life N - for example, urea performs 60-80 days longer;
 improved nutrient uptake, particularly of P and Ca;
 stimulation of beneficial soil life;
 provides magnified nutrition for reduced disease, insect and frost impact;
 salinity management - humates “buffer” plants from excess sodium;
 Organic humates are a catalyst for increasing soil C levels.

Importance of manure

Organic matter is a sponge structure with mineral elements fixed in it. The minerals include: nitrogen,
phosphorous, carbon, sulphur just to mention a few. During mineralization the elements are released
and become available to plant roots. Moreover, humus has a high cation exchange capacity (CEC).
CEC is the sum total of the exchangeable cations that a soil can adsorb. This is because of many
humic colloids present in humus. Refer back to chemical properties of soil to get more about CEC
.The presents of high CEC implies that humus is rich in plant materials and is fertile.
Organic matter helps in the aggregation of soil particles thereby improving soil structure. Soil
structure is the aggregation of individual soil particles to form larger units called aggregates. A well-
structured soil has a good aeration, drainage and roots can penetrate easily through it.
Many clay soils are subjected to erosion. Application of organic matter loosens the soil thus increasing
infiltration while reducing runoff. An increase in organic matter from 1-3% can reduce erosion up to
20-23% because of the stable soil aggregates formation caused by organic matter. Therefore organic
matter is crucial in maintaining soil fertility through soil erosion reduction.

64
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Light textured soils drain quickly. This results in the need for frequent irrigation and leaching of
important plant nutrients. Organic matter behaves differently and is somewhat like a spongy structure
with the ability to absorb and hold up to 90% of its weight in water. Moreover it also acts as mulch
and this reduces the irrigation frequency in the case of a cropping land, it then reduces the cost of
production.
Microorganisms thrive well in soil rich in organic matter content. There are plenty of nutrients for
them and a conclusive temperature for their multiplication. In agriculture, microorganisms are
important as they are involved in organic matter mineralization and nitrogen fixation.
Considering the above points it can be noted that organic matter is very useful in agriculture. More
attention should be focused on maintaining good levels of organic matter in the soil.

Carbon Cycle
Carbon constitutes the major components of every living organism. This can be explained by
considering the carbon cycle. Carbon cycle is a complete series of processes through which all carbon
atoms in existence rotates. The carbon cycle is the greatest natural recycler of carbon atoms.
Plants, animals and soil interacts to make up a basic cycles of nature. Plants absorb carbon dioxide
from the atmosphere and use it, combine it with water they get from the soil to make the substance
they need for growth. Photosynthesis incorporates carbon atoms from carbon dioxide into sugars.
Animals then consume these plants to build their body. Carbon atoms in the sugars will then be fixed
in animal bodies. Carbon atom is lost from these animals through breathing and decomposition back
into the atmosphere. Ultimately, the same carbon atoms can move through many organisms and even
end in the same place where it begun.

Carbon Nitrogen Ratio

The balance of the nitrogen and carbon is important in the soil processes like mineralization and
immobilization. Carbon to nitrogen ratio is the ratio of the mass of carbon to the mass of nitrogen in a
substance. All substances tend to have more carbon and less nitrogen. The ratio is an indicator of
nitrogen limitation of plants and many other organisms. The carbon to nitrogen ratio of the organic
matter added to the soil influences the rate of decomposition of organic matter resulting in either
mineralization or immobilization of soil nitrogen. If the added organic matter contains more nitrogen
in proportion to carbon then nitrogen is released into the soil from the decomposing organic matter.
On the other hand if organic matter has a less amount of nitrogen in relation to carbon then the
microorganisms will utilize the soil nitrogen and the soil nitrogen will be immobilized and will not be
available to plants.

Summary
Organic matter is important as it provides nutrients to the soil and improves some physical properties
of soil like soil structure. The carbon cycle is important as it is involved in the recycling of carbon. C:
N ratio should be kept minimum for good plant growth.

Activities

 Describe the importance of organic matter in agriculture.

 What is the significance of C: N ratio in crop production?

References

65
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Foth, H. D. (1984). Fundamentals of Soil Science. John Wiley and Sons, USA.
Wild, A. (1996). Soil and the Environment. Cambridge University Press, UK.
White, R.E. (1997). Principles and Practice of Soil Science. Blackwell Publishers, USA.

CHAPTER 7

PLANT NUTRIENTS

Introduction
During the previous unit, we noted that organic matter is an important source of nutrients. In this unit
we are going to discuss the importance of specific nutrients in crop production. In general most plants
grow by absorbing nutrients from the soil. The mineral nutrients which come from the soil are
dissolved in water and absorbed through plant roots. They are not always enough of these nutrients in
the soil for plants to grow healthy. That is why farmers use fertilizers to add the nutrients to the soil.
The mineral nutrients are divided into two groups: macro and micro nutrients. Macro nutrients are
those nutrients required by plants in large quantities while micro nutrients are required in small
quantities.

Objectives
By the end of this unit students should be able to:

 Explain the roles of macro- nutrients and micro nutrients in plant growth and development.
 Determine fertilizer requirements in crops...

Macro Nutrients
Nitrogen
Soil nitrogen is found in amine group (-NH2), ammonia (NH3), nitrite (NO2-) and nitrates (NO3-)
forms. However plants can only access it when it is in ammonium or nitrate form. Nitrogen is very
mobile in the soil and therefore is prone to leaching. In most cases nitrogen is applied when required
by plants. It is therefore of paramount importance for farmers to know the growth stages of their crops
that requires more nitrogen. Nitrogen is an essential component of many plant compounds that are
required for different metabolic processes. The compounds include proteins, enzymes, nucleic acids
and growth regulators. It is also part of energy transferring molecules like chlorophyll, ADP and ATP.
Deficiency of soil nitrogen results in stunted growth as they would be limited production of protein
and other materials essential for the production of new cells .The shortage of chlorophyll will give rise
to a pale green colour. The paleness starts in older leaves as nitrogen will be translocated to other
plants. Nitrogen is lost to the atmosphere and deep underground. It is lost through erosion, nutrient
mining (removal of plant residues), leaching and denitrification. Farmers are encouraged to minimize
nitrogen loss through improving soil structure and avoiding prolonged water logging which causes
denitrification.

Phosphorous
It is very immobile in the soil and, depending on soil pH it is usually found in unavailable forms.
Under low PH conditions it is fixed by iron and aluminum while under high PH condition it is fixed
by calcium. It is readily available under neutral conditions. Phosphorus is important in the germination
and growth of seeds, the production of flowers in fruits and the growth of roots. Like nitrogen,
phosphorus is an essential for the process of photosynthesis as it helps with the transformation of solar
energy into chemical energy. It is also involved in the formation of all oils. Phosphorus is limited in
66
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

most soils because it is released very slowly from insoluble phosphate. The phosphorus deficiency in
plants is characterized by an intense green coloration in leaves. If the plant experiences high
phosphorus deficiency the leaves turn purple, become denatured and show signs of necrosis. Older
leaves will show signs of deficiency because phosphorus is very mobile in plants. Usually excess
phosphorus causes potassium deficiency.
Potassium
Potassium is absorbed by plants in larger quantities (luxury consumption) than any other mineral
element except nitrogen and in some cases calcium. The nutrient is found in tree forms: trapped
between clay layers (relatively unavailable), absorbed on the surface of clay colloids (exchangeable)
and in the soil solution (available). Available potassium supply maximum crop production depends on
the type of clay minerals in the soil parent mineral and its resistance to weathering. Potassium
enhances disease resistance in plants by strengthening stalks and stems, contributes to a thicker cuticle
which guards against diseases, controls the turgor pressure within plants to prevent wilting and
enhance fruit size, flower, texture and development. Potassium is also involved in the activation of
enzymes. More than 80 enzymes in plants are activated by potassium. It is also involved in the
translocation of assimilates especially carbohydrates. Crops like potatoes and sugarcanes are heavy
feeders of potassium. It is mobile in plants and will move from lower to upper leaves. If soils have
inadequate potassium, the margin of lower leaves turn brown. This development of a dead tissue is
accompanied by stripped appearance in the remainder of the leaves. The stripping assimilated with
potassium deficiency can be easily confused with deficiency symptoms for sulphur, magnesium and
zinc.

Secondary macro nutrients

Calcium
Calcium is responsible for construction of cell and promoting proper function of growing tissue.
Fortunately calcium occurs naturally in organic soil and there is usually no need for any calcium
supplements to be added in the field. A calcium deficiency in the soil occurs usually in acid soils.
Deficiency symptoms of sulphur usually occurs in young growth since .If calcium is deficient the
terminal part of the plant may be malformed and disintegrated may take place. Marginal colorations of
leaves may occur and it moves inwards firstly in young leaves.

Magnesium
It also occurs naturally in organic soils which usually makes adding any magnesium supplement to
garden soil rather superfluous. Magnesium is part of chlorophyll and thus plays a part in
photosynthesis. Magnesium deficiency symptoms are similar to that of maize: yellowing of older
leaves.

Sulphur
Most chemicals and organic fertilizers contain sulphur. Sulphur makes its deficiency symptoms very
rare. The element is responsible for the synthesis of protein and plant enzymes. Sulphur deficiency
symptoms are the same as nitrogen as sulphur form part of plant protein and play a role in the
formation of chlorophyll.
Micro nutrients
They are also known as trace elements. The micro nutrients of plants are made up of: Iron (Fe), Zinc
(Zn), Manganese (Mn), Boron (B), Molybdenum (Mo) and Copper (Cu). These are elements required

67
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

by plants in biological nitrogen fixation. Molybdenum deficiency produces whip tail in some
vegetation such as cauliflower, broccoli and other brassicas spp. The molybdenum deficiency will
reduce the activity of the symbiotic and non-symbiotic nitrogen fixing microorganism.

Boron is involved in cell division and growth of meristems. It also functions to facilitate normal seed
setting, fruit development, and successful pollination and encourages pollen grains germination and
development. Boron deficiency can manifest itself in various forms that are developed on the type
plants and their age of the plant. In some fruits like apple it causes die back, yellowing and resetting of
Lucerne crops.
Iron is an important element and is greatly concerned with photosynthesis .It is very insoluble and is
rendered still more so high PH conditions. Symptoms of shortages are yellowing of the whole leaf,
although extreme cases the leaf can turn white, the older leaves usually being more affected than
younger leaves.
Manganese is much concerned with photosynthesis activities. That is why any shortage produces a
leaf molting effect. Toxicity of manganese is much more common problem. A blue black coloration
develops on leaf tips in addition to a general dropping.
Summary
Plants require nutrients for growth and development. The nutrients are macro (required in larger
quantities) and micro nutrients (required in smaller quantities). If the micronutrients are supplied in
large quantities, they become toxic to plants.

Activities

 Describe the functions of nitrogen, phosphorous and potassium.

 List the deficiency symptoms of four micro elements.

DETERMING NUTRIENT STATUS AND NEEDS

 Common diagnostic tools to determine nutrient needs of plants include deficiency symptoms,
chemical soil tests and plant tissue tests
 What do you think are some of the disadvantages of using deficiency symptoms as a diagnostic
tool?
Soil analysis
 Begins with soil sampling
 If soils are wrongly sampled then a wrong recommendation will be given even though the
analysis of the wrong sample will be correct representative sample is needed
and nutrient content changes through the soil profile hence soil samples should be taken
from a consistent depth
-51cm, ploughed fields to plough
depth and fields planted or to be planted in rows but not ploughed there is need to core to the depth
where 75% of plant roots will be found
Plant /foliar analysis (Leaf analysis)

While soil analysis reveals the levels of essential soil nutrients, leaf analysis shows the grower
exactly what the plant has successfully absorbed
t analysis can be helpful in detecting nutrient deficiencies before they affect pant health and
yield.
68
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

fertilization program as it shows exactly what the plant has succeeded in taking up.

they affect plant health or yield

s could
make a difference in plant growth as well as times when accumulation is at its maximum or steady
state -leaf samples should be taken when most vegetative growth has subsided
o perform a leaf analysis
for each block once every three years to ensure maintenance of adequate nutrient levels.
 If nutritional problems are suspected in a given planting it’s good to take both leaf and soil tests

BIOLOGICAL NITROGEN FIXATION

All plants, including forage crops, need relatively large amounts of nitrogen (N) for proper growth and
development. Biological nitrogen fixation (BNF) is the term used for a process in which nitrogen gas
(N2) from the atmosphere is incorporated into the tissue of certain plants. Biological nitrogen fixation
(BNF) occurs when atmospheric nitrogen is converted to ammonia by an enzyme called
a [Link] overall reaction for BNF is:

N2 + 8 H+ + 8 e− → 2 NH3 + H2

The process is coupled to the hydrolysis of 16 equivalents of ATP and is accompanied by the co-
formation of one molecule of H2. The conversion of N2 into ammonia occurs at
a cluster called FeMoco, an abbreviation for the iron-molybdenum cofactor. The mechanism proceeds
via a series of protonation and reduction steps wherein the FeMoco active site hydrogenates the
N2 substrate. FeMoco is the primary cofactor of nitrogenase. Nitrogenase is the enzyme that catalyses
the conversion of atmospheric N2 into ammonia (NH3), through the process known as nitrogen
fixation. Containing iron and molybdenum, the cofactor is called FeMoco. Its stoichiometry is
Fe7MoS9C.

In free-living diazotrophs, the nitrogenase-generated ammonium is assimilated into glutamate through

the glutamine synthetase/glutamate synthase pathway.

The microbial genes required for nitrogen fixation are widely distributed in diverse environments.

Enzymes responsible for nitrogenase action are very susceptible to destruction by oxygen. For this
reason, many bacteria cease production of the enzyme in the presence of oxygen. Many nitrogen-
fixing organisms exist only in anaerobic conditions, respiring to draw down oxygen levels, or binding
the oxygen with a protein such as leghemoglobin.

Only a select group of plants is able to obtain N this way, with the help of soil microorganisms.
Among forage plants, the group of plants known as legumes (plants in the botanical family Fabaceae)
are well known for being able to obtain N from air N2.

In forage production, this process can be very important because it means that the much needed N can
be obtained from three sources: the atmosphere via BNF, the soil, and from fertilizers. Forage
producers who find ways to maximize the amount of N obtained from the atmosphere via BNF will be
able to reduce their fertilizer costs while maintaining soil fertility, high levels of forage protein, and
high yields.

69
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The process by which some forage crops can incorporate N2 from the air into their tissues involves a
host plant (also known as the macrosymbiont). For example: alfalfa and a microorganism (also known
as the microsymbiont) that is associated with the host plant function in what is called a symbiotic
relationship or symbiosis. A symbiotic relationship is one in which two organisms form a mutually
beneficial relationship. With most forage crops the second organism is a bacteria that occurs naturally
in the soil. The bacteria that is most often involved with forage crops is popularly known as rhizobia,
because it is classified as part of the bacterial genus known as Rhizobium.

Microorganisms that fix nitrogen

Diazotrophs includes cyanobacteria (e.g. the highly significant Trichodesmium and Cyanothece), green sulfur
bacteria, and Azotobacteraceae, rhizobia and Frankia.

Cyanobacteria play key roles in the carbon and nitrogen cycle of the biosphere. In
general, cyanobacteria can use various inorganic and organic sources of combined nitrogen,
like nitrate, nitrite, ammonium, urea, or some amino acids.

Root nodule symbioses

Legume family

Plants that contribute to nitrogen fixation include those of the legume family – Fabaceae – with taxa
such as kudzu, clovers, soybeans, alfalfa, lupines, peanuts, and rooibos. They
contain symbiotic bacteria called rhizobia within nodules in their root systems, producing nitrogen
compounds that help the plant to grow and compete with other plants. When the plant dies, the fixed
nitrogen is released, making it available to other plants; this helps to fertilize the soil. The great
majority of legumes have this association, but a few genera (e.g., Styphnolobium) do not. In many
traditional and organic farming practices, fields are rotated through various types of crops, which
usually include one consisting mainly or entirely of clover or buckwheat (non-legume
family Polygonaceae), often referred to as "green manure".

The efficiency of nitrogen fixation in soil is dependent on many factors, including the legume as well
as air and soil conditions. For example, nitrogen fixation by red clover can range from 50 - 200 lb/acre
depending on these variables.

Although the process involves a number of complex biochemical reactions, it may be summarized in a
relatively simple way by the following equation:

N2 + 8H2+ 16ATP ------> 2 NH3 + 2H2+ 16ADP + 16 Pi

The equation above indicates that one molecule of nitrogen gas (N2) combines with eight hydrogen
ions (also known as protons) (8H+) to form two molecules of ammonia (2NH3) and two molecules of
hydrogen gas (2H2). This reaction is conducted by an enzyme known as nitrogenase. The 16
molecules of ATP (ATP = Adenosine Triphosphate, an energy storing compound) represent the
energy required for the BNF reaction to take place. In biochemical terms 16 ATP represents a
relatively large amount of plant energy. Thus, the process of BNF is 'expensive' to the plant in terms
of energy usage.

What is the ultimate source of this energy needed for BNF? The sun, via the process of
photosynthesis. As ammonia (NH3) is formed it is converted to an amino acid such as glutamine. The

70
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Nitrogen in amino acids can be used by the plant to synthesize proteins for its growth and
development.

IMPORTANCE OF BNF

Use of inorganic fertilizers led to worldwide, ecological problems, such as the formation of coastal
dead zones. Biological nitrogen fixation, on the other hand, offers a natural means of providing
nitrogen for plants. It is a critical component of many aquatic, as well as terrestrial ecosystems across
our biosphere.

1. Greater use of BNF will reduce society's current dependence on synthetic N fertilizers. The
production of widely used synthetic N fertilizers such as anhydrous ammonia requires the use of
relatively large amounts of energy from non-renewable energy sources such as natural gas.
Distribution and application of these fertilizers also requires relatively large amounts of non-
renewable energy sources such as diesel fuel.

2. Greater use of BNF can help enhance environmental quality by reducing problems with air and
water pollution. The over-application of synthetic N fertilizers has been linked to excessive nitrate
(NO3-) levels in groundwater in a number of locations around the United States. Excessive nitrate
concentrations may have detrimental effects on human health. Both the manufacture and application
of synthetic N fertilizers involves burning non-renewable fossil fuels such as natural gas, diesel fuel,
and gasoline, which have been shown to contribute to air pollution.

3. Greater use of BNF can help lower production costs and thus increase profit margins for producers.
The use of crops that fix nitrogen in crop rotations can significantly reduce N fertilizer needs for crops
in the rotation. For example, following alfalfa with corn in a rotation will allow profitable yields of
corn with less expense for fertilizer purchase and application.

4. Greater use of BNF can help enhance sustainable food production by improving soil fertility and
tilth. Some producers have found that the use of so called green manure crops is a more sustainable
fertilizer alternative than purchased, synthetic fertilizer. Green manure crops are crops grown
specifically to be incorporated into the soil rather than for harvest. Using green manure crops that fix
atmospheric N2 may potentially increase soil N levels and organic matter content over time.
Additional organic matter in soils generally improves the tilth of a soil, where tilth refers to desirable
physical properties of soil such as proper drainage, water holding capacity, aeration, and structure.

TEST YOURSELF

1. Hue denotes
(A) Dominant spectrum (B) Lightness or brightness
(C) Purity (D) Intensity
Ans. A
2. Phosphorus uptake in alkali soil in the form of _____
(A) H2PO4– (B) HPO42–
3–
(C) PO4 (D) H3PO4
Ans. C
3. Which fertilizer produce acidity in soil
(A) Ammonium sulfate (B) Sodium nitrate
(C) Calcium ammonium nitrate (D) Calcium nitrate
Ans. A
4. In munsell colour chart hue 10 represents as ______

71
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

(A) Red (B) Black

(C) Grey (D) Yellow
Ans. D
5. Available water held between
(A) Saturation to wilting point (B) Field capacity to hygroscopic coefficient
(C) Only field capacity (D) Field capacity to wilting point
Ans. D
6. Soil colloidal particle shows the phenomena
(A) Plasticity (B) Adhesion and cohesion
(C) Flocculation (D) All of the above
Ans. D
7. The central cation on silica tetrahedron is _____
(A) Al3+ (B) Si4+
2+
(C) Fe (D) None of the above
Ans. B
8. Mica is a type of clay mineral
(A) 1:1 (B) 2:1
(C) 2:1:1 (D) None of the above
Ans. B
9. Hydrogen bond found in which clay mineral
(A) Kaolinite (B) Montmorillonite
(C) Vermiculite (D) Beidelite
Ans. A
10. Base saturation in lateritic soil
(A) < 40 % (B) > 40%
(C) < 20% (D) > 20%
Ans. B
11. Immobilization of sulphur occur in soil when C:S ratio is _____
(A) < 300:1 (B) > 300:1
(C) > 400:1 (D) > 200:1
Ans. C
12. Denitrification is a process of ______
(A) Oxidation (B) Reduction
(C) Hydration (D) Carbonation
Ans. B
13. Microorganism involve in conversion from nitrite to nitrate
(A) Nitrosomonas (B) Nitrobactor
(C) Pseudomonas (D) Bacillus
Ans. B
14. Most of the soil organism is _____
(A) Psychrophiles (B) Mesophiles
(C) Thermophiles (D) All of the above
Ans. B
15. Optimum growth of bacteria in soil
(A) Acidic (B) Alkali
(C) Neutral to slightly alkaline (D) Slightly acidic to neutral
Ans. C
16. Which genera of bacteria found maximum in soil?
(A) Azotobacter (B) Pseudomonas
(C) Bacillus (D) All of the above
Ans. C
17. Which nutrient influence the nodule formation and nitrogen fixation in soil?
(A) Ph (B) N
(C)P (D) All of the above
Ans. D
72
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

18. Nitrification process ceases at pH

(A) < 5 (B) < 6
(C) < 4 (D) < 7
Ans. A
19. Denitrification mostly takes place in _____
(A) Aerobic (B) Anaerobic
(C) Both (D) None of the above
Ans. B
20. Nitrification rapid in _____ season
(A) Winter (B) Rainy
(C) Summer (D) All of the above
Ans. C
21. The peak microbial population was observed in soil generally after addition of residue
(A) 3-4 week (B) 2-3 week
(C) 3-4 days (D) 1-2 week
Ans. D
22. Which element suppressed Downey mildew of pear millets?
(A) Chloride (B) Fluoride
(C) Bromide (D) Lithium
Ans. A
23. Improper development of wheat inflorescence is due to deficiency of _____
(A) Molybdenum (B) Boron
(C) Calcium (D) Magnesium
Ans. B
24. Feldspars is primary mineral that occurs pre-dominantly in ________
(A) Igneous rock (B) Sedimentary rock
(C) Metamorphic rocks (D) All of the above
Ans. A
25. Bedrock is absent in which soil?
(A) Black soil (B) Red soil
(C) Alluvial soil (D) Forest soil
Ans. C
26. Which soil occur mainly a soil crust problem?
(A) Sandy soil (B) Silty clay loam
(C) Loamy soil (D) Clayey soil
Ans. B
27. Red colour of soil is due to _____
(A) Hematite (B) Geothite
(C) Glauconite (D) Maghemite
Ans. A
28. Black colour of soil is due to _____
(A) Titaniferous magnetite (B) Geothite
(C) Glauconite (D) Organic matter
Ans. A
29. Micronutrient flow is mainly a _____ process in soil
(A) Mass flow (B) Diffusion
(C) Interception (D) None of the above
Ans. B
30. When organic matter added into the soil effect on bulk density
(A) Increased (B) No effect
(C) Decreased (D) Both A and C
Ans. C
31. Which clay mineral is responsible for cracking in black soil?
(A) Kaolinite (B) Illite
(C) Vermiculite (D) Montmorillonite
73
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Ans. D
32. pH meter measures
(A) H ions activity (B) H ions concentration
(C) Al ions (D) All of the above
Ans. A
SECTION B

1. Explain primary soil minerals and 2 give examples [4]

2. Define a soil catena and explain its significance.[4]
3. Outline the significance of soil air.[4]
[Link] the functions of: boron,nitrogen,copper,magnesium.[8]
[Link] the importance of Biological Nitrogen Fixation in crop production.[4]
[Link] the aid of diagrams describe the significance of soil horizons [8]
[Link] the importance of soil organic matter in relation to CEC, soil pH, plant nutrients and moisture
retention. (8)
[Link] and explain constituencies of soil organic matter. (8)
b)Identify organisms involved in different stages of the decomposition process of organic matter. (6)
c) Describe how CN ratio affects the rate of decomposition. (4)

CHAPTER 9

PLANT GROWTH
-describe the types of plant meristems
MERISTEMS
Key Terms
meristem: the plant tissue composed of totipotent cells that allows plant growth

74
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

undifferentiated: describes tissues where the individual cells have not yet developed mature or
distinguishing features, or describes embryonic organisms where the organs cannot be identified
apical: situated at the growing tip of the plant or its roots, in comparison with intercalary growth
situated between zones of permanent tissue
Plant growth occurs in areas called meristems, which are the site of repeated cell division of
unspecialised cells. These cells differentiate, and become specialised in relation to the function they
will perform.
Meristems are actively dividing, tissues responsible for both primary and secondary growth.
Meristems
Meristematic cells are also responsible for keeping the plant growing. The plant body that develops
after germination depends on the activities of meristematic tissues. Meristematic tissues are lumps of
small cells with dense cytoplasm and proportionately large nuclei that act like stem cells in animals.
That is, one cell divides to give rise to two cells. One remains meristematic, while the other is free to
differentiate and contribute to the plant body. In this way, the population of meristem cells is
continually renewed. Elongation of both root and shoot takes place as a result of repeated cell
divisions and subsequent elongation of the cells produced by the apical meristems. In some vascular
plants, including shrubs and most trees, lateral meristems produce an increase in root and shoot
diameter (girth).
Apical Meristems
Apical meristems are found at the tips of stems and at the tips of roots just behind the root cap. The
plant tissues that result from primary growth are called primary tissues.
Unspecialised cells, which undergo the following sequence to become a functional part of the plant
New unspecialised cells become available at the meristems, the site of mitosis
These cells become elongated and undergo vacuolation
They become specialised (differentiated) to perform a particular function
They then form part of a permanent tissue which performs a particular role within the plant
The cycle continues for growth and regeneration purposes

During periods of growth, the cells of apical meristems divide and continually add more cells to the
tips of a seedling’s body. Thus, the seedling lengthens. The elongation of the root and stem forms
what is known as the primary plant body, which is made up of primary tissues. The primary plant
body comprises the young, soft shoots and roots of a tree or shrub, or the entire plant body in some
herbaceous plants. Both root and shoot apical meristems are composed of delicate cells that need
protection.
The Shoot Apical Meristem (SAM) gives rise to organs like the leaves and flowers, while the Root
Apical Meristem (RAM) provides the meristematic cells for the future root growth. The cells of the
shoot and root apical meristems divide rapidly and are considered to be indeterminate, which means
that they do not possess any defined end fate. The root apical meristem is protected from the time it
emerges by the root cap. Root cap cells are produced by the root meristem and are sloughed off and
replaced as the root moves through the soil. A variety of adaptive mechanisms protect shoot apical
meristem during germination. The epicotyls or hypocotyl (“stem like” tissue above or below the
cotyledons) may bend as the seedling emerges to minimize the force on the shoot tip. In the monocots
(a late evolving group of angiosperms) there is often a coleoptile (sheath of tissue) that forms a
protective tube around the emerging shoot. Later in development, the leaf primordia cover the shoot
apical meristem which is particularly susceptible to desiccation.
Apical Meristem Function
The apical meristem is found at the ends of roots (root apical meristem) or the tops of shoots (shoot
apical meristem) of a plant, and is responsible for the plant’s growth in length or height. This type of
growth is known as primary growth. The presence of an apical bud (or terminal bud) exerts apical
dominance over the axillary buds, ultimately promoting vertical growth and hindering lateral growth.

75
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

When the apical bud is removed, signals blocking growth from the axillary buds disappear to activate
lateral growth.
Shoot Apical Meristem
The shoot apical meristem, found above ground, is composed of undifferentiated cells that have one of
three destinies. They can develop to become one of three primary meristems: the protoderm, ground
meristem, or procambium. The protoderm will go on to form the epidermal tissues of the plant; the
ground meristem will form the cortext and pith of the plant; and the procambium will become xylem
and phloem, the vascular tissues of the plant.
The shoot of a plant also includes its leaves, which grow from the sides of the apical meristem. The
beginning growth of the leaf at the node results in a bump, or an axillary bud, at the node. If the
terminal bud is in close proximity to the axillary bud, the axillary buds will remain dormant. However,
if the terminal bud is removed or if the distance between the terminal bud and axillary bud increases—
as happens when the plant grows—then the inhibiting factors exerting apical dominance diminish or
disappear, allowing for the growth of leaves at the lateral buds of the apical meristem. When flowering
plants are ready to bloom, the shoot apical meristem of the plant becomes an inflorescence meristem
where petals, sepals, stamens, and other flower parts emerge.
Root Apical Meristem
The root apical meristem, found below ground, is responsible for the growth and development of a
plant’s roots. The root meristem produces cells in a bilateral direction, meaning that it yields two types
of tissues at the same time. One tissue comprises the main roots of the plant that supply proliferative,
undifferentiated cells for continued growth, and the other forms a root cap that protects the apical
meristem and the source of new cells. Because the roots are growing and the root cap is continuously
being ground down into the soil, cells of the root cap are constantly being shed and replaced by new
cells, as provided by the main root. This is typical of a tap root. Lateral root meristems account for the
lateral growth of roots from the main root, into vastly branched root systems. Lateral root growth help
to increase the plant’s efficiency in water and nutrient absorption, nutrient storage, and stability for
aerial growth.
The apical meristem gives rise to three types of embryonic tissue systems called primary meristems.
Cell division continues in these partly differentiated tissues as they develop into the primary tissues of
the plant body. The three primary meristems are the protoderm, which forms the epidermis; the
procambium, which produces primary vascular tissues (primary xylem and primary phloem); and the
ground meristem, which differentiates further into ground tissue, which is composed of parenchyma
cells. In some plants, such as horsetails and corn, intercalary meristems arise in stem internodes,
adding to the internode lengths.
Lateral Meristems
Lateral meristems are responsible for secondary thickening, which is required by perennial plants that
grow year after year, and need the structural support.
Most trees, shrubs, and some herbs have active lateral meristems, which are cylinders of meristematic
tissue within the stems and roots. Although secondary growth increases girth in many nonwoody
plants, its effects are most dramatic in woody plants which have two lateral meristems.

Within the bark of a woody stem is the cork cambium, a lateral meristem that produces the cork cells
of the outer bark. Just beneath the bark is the vascular cambium, a lateral meristem that produces
secondary vascular tissue. The vascular cambium forms between the xylem and phloem in vascular
bundles, adding secondary vascular tissue on opposite sides of the vascular cambium. Secondary
xylem is the main component of wood. Secondary phloem is very close to the outer surface of a woody
stem. Removing the bark of a tree damages the phloem and may eventually kill the tree. Tissues
formed from lateral meristems, which comprise most of the trunk, branches, and older roots of trees
and shrubs, are known as secondary tissues and are collectively called the secondary plant body.
Intercalary meristem

76
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

In angiosperms, intercalary meristems occur only in monocot (in particular, grass) stems at the base of
nodes and leaf blades. Horsetails also exhibit intercalary growth. Intercalary meristems are capable of
cell division, and they allow for rapid growth and regrowth of many monocots. Intercalary meristems
at the nodes of bamboo allow for rapid stem elongation, while those at the base of most grass leaf
blades allow damaged leaves to rapidly regrow. This leaf regrowth in grasses evolved in response to
damage by grazing herbivores.
Floral meristem
When plants begin developmental process known as flowering, the shoot apical meristem is
transformed into an inflorescence meristem, which goes on to produce the floral meristem, which
produces the sepals, petals, stamens, and carpels of the flower.
In contrast to vegetative apical meristems and some efflorescence meristems, floral meristems cannot
continue to grow indefinitely. Their future growth is limited to the flower with a particular size and
form. The transition from shoot meristem to floral meristem requires floral meristem identity genes,
that both specify the floral organs and cause the termination of the production of stem
cells. AGAMOUS (AG) is a floral homeotic gene required for floral meristem termination and
necessary for proper development of the stamens and carpels. AG is necessary to prevent the
conversion of floral meristems to inflorescence shoot meristems, but is identity gene LEAFY (LFY)
and WUS and is restricted to the centre of the floral meristem or the inner two whorls. This way floral
identity and region specificity is achieved. WUS activates AG by binding to a consensus sequence in
the AG’s second intron and LFY binds to adjacent recognition sites. Once AG is activated it represses
expression of WUS leading to the termination of the meristem.
Through the years, scientists have manipulated floral meristems for economic reasons. An example is
the mutant tobacco plant "Maryland Mammoth." In 1936, the department of agriculture of Switzerland
performed several scientific tests with this plant. "Maryland Mammoth" is peculiar in that it grows
much faster than other tobacco plants.
Apical dominance
Apical dominance is the phenomenon where one meristem prevents or inhibits the growth of other
meristems. As a result, the plant will have one clearly defined main trunk. For example, in trees, the
tip of the main trunk bears the dominant shoot meristem. Therefore, the tip of the trunk grows rapidly
and is not shadowed by branches. If the dominant meristem is cut off, one or more branch tips will
assume dominance. The branch will start growing faster and the new growth will be vertical. Over the
years, the branch may begin to look more and more like an extension of the main trunk. Often several
branches will exhibit this behavior after the removal of apical meristem, leading to a bushy growth.
The mechanism of apical dominance is based on auxins, types of plant growth regulators. These are
produced in the apical meristem and transported towards the roots in the cambium. If apical
dominance is complete, they prevent any branches from forming as long as the apical meristem is
active. If the dominance is incomplete, side branches will develop.
Recent investigations into apical dominance and the control of branching have revealed a new plant
hormone family termed strigolactones.
ON THE BASIS OF ORIGIN:
Meristems are 3 types:
PROMERISTEM: Also referred as embryonic meristem. Present at the apices of the shoot and root
tip. It divides to form primary meristem.
PRIMARY MERISTEM: Found below the pro-meristem in the shoot and root tip & intercalary
meristem. They divide to produce primary permanent tissue that forms the primary plant body.
Primary meristems
Apical meristems may differentiate into three kinds of primary meristem:
Protoderm: lies around the outside of the stem and develops into the epidermis.
Procambium: lies just inside of the protoderm and develops into primary xylem and primary phloem.
It also produces the vascular cambium, and cork cambium, secondary meristems. The cork cambium

77
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

further differentiates into the phelloderm (to the inside) and the phellem, or cork (to the outside). All
three of these layers (cork cambium, phellem, and phelloderm) constitute the periderm. In roots, the
procambium can also give rise to the pericycle, which produces lateral roots in eudicots.
Ground meristem: develops into the cortex and the pith. Composed
of parenchyma, collenchyma and sclerenchyma cells.
These meristems are responsible for primary growth, or an increase in length or height, which were
discovered by scientist Joseph D. Carr of North Carolina in 1943.
SECONDARY MERISTEM: It is formed later in the plant body. It develops from primary
permanent tissue due to dedifferentiation. They divide to produce secondary permanent tissue that
forms the secondary plant body.
There are two types of secondary meristems, these are also called the lateral meristems because they
surround the established stem of a plant and cause it to grow laterally (i.e., larger in diameter).
Vascular cambium, which produces secondary xylem and secondary phloem. This is a process that
may continue throughout the life of the plant. This is what gives rise to wood in plants. Such plants are
called arborescent. This does not occur in plants that do not go through secondary growth (known
as herbaceous plants).
Cork cambium, which gives rise to the periderm, which replaces the epidermis.

ON THE BASIS OF POSITION:

Meristems are 3 types:
APICAL MERISTEM: It is present at the root and the shoot tip. Consists of pro-meristem & primary
meristem. Divide to produce primary permanent tissue that increases the length of the plant. It gives
rise to organs like leaves & flowers. During leaf formation and stem elongation, some cells separate,
they form the axillary bud. The axillary bud grows into either branch or flower.

INTERCALARY MERISTEM: It is present in permanent tissues. A part of the apical meristem

separates due to the formation of permanent tissues in between during growth. Divides to produce

78
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

primary permanent tissue that increases the length of the plant. It is found present below the nodes in

mint & above the node in grasses.

LATERAL MERISTEM: As it is lateral in position it is referred as lateral meristem. Divides to
produce secondary permanent tissue that increases the thickness of the plant. E.g., interfascicular
cambium, intrafascicular cambium & cork cambium.
Intrafascicular cambium is present in primary plant body but divides to produce secondary permanent
tissue.
ON THE BASIS OF PLANE OF DIVISION:
Meristems are 3 types:
RIB MERISTEM: Cells divide in one plane, forming a row of cells. E.g., formation of lateral roots.
PLATE MERISTEM: Cells divide in two planes, forming a plate like structure. E.g., formation of
lamina in leaves.
MASS MERISTEM: Cells divide in all planes, forming a mass of cells. E.g., formation of cortex,
pith, endosperm.
ON THE BASIS OF FUNCTION:
Meristems are 3 types:
PROTODERM: Forms the Epidermal tissue system.
PROCAMBIUM: Forms the Vascular tissue system.
GROUND MERISTEM: Forms the Fundamental or Ground tissue system.

Indeterminate growth of meristems

Though each plant grows according to a certain set of rules, each new root and shoot meristem can go
on growing for as long as it is alive. In many plants, meristematic growth is potentially indeterminate,
making the overall shape of the plant not determinate in advance. This is the primary growth. Primary
growth leads to lengthening of the plant body and organ formation. All plant organs arise ultimately
from cell divisions in the apical meristems, followed by cell expansion and differentiation. Primary
growth gives rise to the apical part of many plants.
The growth of nitrogen-fixing root nodules on legume plants such as soybean and pea is either
determinate or indeterminate. Thus, soybean (or bean and Lotus japonicus) produce determinate
nodules (spherical), with a branched vascular system surrounding the central infected zone. Often,
Rhizobium infected cells have only small vacuoles.
Induced meristems
Meristems may also be induced in the roots of legumes such as soybean, Lotus japonicus, pea,
and Medicago truncatula after infection with soil bacteria commonly called Rhizobia
THEORIES EXPLAINING THE APICAL MERISTEM:
APICAL CELL THEORY was given by Nageli. According to this theory, apical meristem has a
single apical cell. This is true only in lower plants, i.e., algae, bryophytes & some pteridophytes.
HISTOGEN THEORY given by Hanstein. According to this theory, apical meristem has 3
histogens.
79
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Outer Dermatogen: that forms epidermis.

Middle Periblem: that forms region between epidermis & endodermis.
Inner Plerome: that forms region inner to endodermis.
Calyptrogen is present at the root tip, the cells of which divide to form root cap.
Quiescent center: are a mass of cells present at the root tip in the periblem, that either not dividing or
dividing very slowly. Discovered by Clowe in Zea mays.

TUNICA CORPUS THEORY given by Schmidt. This is the applicable theory. According to this
theory, apical meristem has 2 parts:
Outer Tunica that forms the epidermis.
Inner Corpus that forms the body.

TEST YOURSELF
Q1. Cortex is formed from
A. Cambium
B. Procambium
C. Ground meristem
D. Protoderm
Answer:3
Q2. Plerome is a histogen that gives rise to
A. Pericycle
B. Pith
C. Vascular bundles
D. All the above
Answer: 4
Q3. In case tunica is multilayered, which is likely to happen
A. All layers take part in formation of multilayered epidermis
B. All the layers produce cortex
C. Outer-most layer forms hypodermis, middle layer cortex and inner layer endodermis
D. Only the outer most layer forms the epidermis
Answer: 4
Q4. Apical meristem divides from three meristematic regions
A. Dermal, vascular and ground
B. Calyptrogens, periblem and plerome
C. Protoderm, procambium and ground meristem
D. Lateral, intercalary and subapica
Answer: 3
Q5. Which is true
A. Stem branches are endogenous
80
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

B. Stem branches develop from pericycle

C. Root branches are exogenous
D. Root branches are endogenous
Answer: 4
Q6. Period between initiation of two successive leaves or nodes is
A. Plastochron
B. Diurnal cyclicity
C. Internode
D. Allelopathy
Answer: 1
Q7. An injured plant part is repaired with the help of
A. Lateral meristem
B. Primary meristem
C. Intercalary meristem
D. apical meristem
Answer: 1
Q8. Tunica is
A. Mass meristem
B. Mantle of meristem
C. Internal mass of meristem
D. Central of meristem
Answer: 2
Q9. Corpus is
A. Mantle of meristem
B. Central part of meristem
C. Internal part of meristem
D. Mass meristem
Answer: 3
8. The cells having the ability to divide are
A. Specialized
B. Permanent
C. Meristematis
D. Glandular
Answer: 3
Q10. Meristematic cells are
A. Differentiated
B. Mature and dead
C. Mature and living
D. Immature and living
Answer: 4
Q11. The characteristics of meristematic cell are
A. Thin wall and dense cytoplasm with small amount of endoplasmic reticulum
B. Isodiametric with no intercullar spaces
C. Conspicuous nucleus
D. All the above
Answer: 4
SECTION B
1. Define the term meristem.[2]
2. Explain the types of meristems[12]

81
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 10

PLANT GROWTH

 discuss plant growth and development

 describe the phases of plant cell growth

GROWTH

Growth can be defined as an irreversible permanent increase in size of an organ or its parts or even of
an individual cell. It is an increase in dry weight of bulk of an organism
Genetically, growth is the increase in cytoplasmic and organelle volume (G1 phase), as well as
increase in genetic material (G2 phase) following the replication during S phase.
Generally, growth is accompanied by metabolic processes.

Growth can be measured

Growth, at a cellular level, is principally a consequence of increase in the amount of protoplasm. Since
increase in protoplasm is difficult to measure directly, one generally measures some quantity which is
more or less proportional to it. Growth is, therefore, measured by a variety of parameters some of
which are: increase in fresh weight, dry weight, length, area, and volume and cell number.

Phases of Growth

The period of growth is generally divided into three phases, namely, meristematic, elongation and
maturation. Let us understand this by looking at the root tips. The constantly dividing cells, both at the
root apex and the shoot apex, represent the meristematic phase of growth. The cells in this region are
rich in protoplasm, possess large conspicuous nuclei. Their cell walls are primary in nature, thin and
cellulosic with abundant plasmodesmatal connections. The cells proximal (just next, away from the
tip) to the meristematic zone represent the phase of elongation. Increased vacuolation, cell
enlargement and new cell wall deposition are the characteristics of the cells in this phase.

Further away from the apex, i.e., more proximal to the phase of elongation, lies the portion of axis
which is undergoing the phase of maturation. The cells of this zone, attain their maximal size in terms
of wall thickening and protoplasmic modifications.

82
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Detection of zones of elongation by the parallel line technique. Zones A, B, C, and D immediately
behind the apex have elongated most.

Growth Rates

The increased growth per unit time is termed as growth rate. Thus, rate of growth can be expressed
mathematically. An organism, or a part of the organism can produce more cells in a variety of ways.

The growth rate shows an increase that may be arithmetic or geometrical.

In arithmetic growth, following mitotic cell division, only one daughter cell continues to divide while
the other differentiates and matures. The simplest expression of arithmetic growth is exemplified by a
root elongating at a constant rate.

In geometrical growth. In most systems, the initial growth is slow (lag phase), and it increases rapidly
thereafter – at an exponential rate (log or exponential phase). Here, both the progeny cells following
mitotic cell division retain the ability to divide and continue to do so.
83
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

However, with limited nutrient supply, the growth slows down leading to a stationary phase. If we plot
the parameter of growth against time, we get a typical sigmoid or S-curve. A sigmoid curve is a
characteristic of living organism growing in a natural environment.

FLOWER AND FRUIT DEVELOPMENT

 Explain how vernalisation and photoperiodism influence flower initiation.

 Describe gamete formation in plants.
 Describe pollination mechanisms in plants.
 Explain the concept of double fertilization in plants.

CHAPTER 11

VERNALISATION

Vernalisation in Plants: Site, Requirements, Mechanism and Importance!

Is the induction of a plant's flowering process by exposure to the prolonged cold of winter, or by an
artificial equivalent. After vernalization, plants have acquired the ability to flower, but they may
require additional seasonal cues or weeks of growth before they will actually flower.
Temperature plays significant role in metabolic activities of plants. Temperature is one of the
important factors determining the distribution of plants. Temperature also plays vital role in the
germination of seeds and subsequent flowering of plants.
Plants of temperate zone, as expected, germinate at a relatively low temperature, whereas tropical
plants germinate best at much higher temperature.

Many plants do not come to flower before they experience a low temperature. These plants remain
vegetative during warm season, experience low temperature during winter, grow further and then bear
flowers and fruits. It was found by Lysenko (1928), a Russian scientists that the cold- requiring
84
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

biennial plants can be made to flower in one growing season by providing low temperature treatment
to young plants or moistened seeds. He called the effect of this chilling treatment as vernalisation.
Vernalisation is, therefore, a process of shortening of juvenile or vegetative phase and hastening
flowering by a previous cold treatment.
An example of winter rye may be quoted here. When the seeds of this variety of rye were germinated
at 1°C for four weeks, the plants flowered eleven weeks after planting, but at the same time seeds
germinated at 18°C did not produce flowering shoot in the same duration

Another interesting fact about the effects of vernalisation came from the work on biennial varieties
of Hyoscyamus niger (henbane) by Melchers and Lang (1948). This variety of henbane will flower
only when vernalisation is followed by long day treatment, and vernalisation followed by short-
day treatment fails to induce flowering (see figure 5.6).

Still more interesting is the fact that annual variety of henbane does not require cold treatment for
flowering. The annual variety differs from biennial ones in the possession of a single dominant
gene which functions as a substitute to vernalisation.
It is presumed that the said gene brings about direct production of the precursor of flowering
substance, which in the biennial variety requires cold treatment. These findings indicate the
possibility of conversion of certain hormone precursor into a flower-inducing active form which
under the influence of appropriate day-length induces flowering. Common examples of plants
requiring vernalisation are winter rye, winter wheat, winter oat, winter barley, pea, beet, cabbage,
Henbane, Chrysanthemum, Viola, Clover, etc.
85
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Site of Vernalisation (Site of Perception):

Site of perception of cold stimulus is different in different plants. It can be a germinating seed or
metabolically active embryo (Secale cereale), shoot apical meristem (Chrysanthmum) or
vegetative parts such as leaves (Hyoscyamus niger and other biennials).
Requirements of Vernalisation:
(i) Low temperature:
Vernalisation, unlike photoperiodism, is a cumulative process because plants become gradually
more and more effectively vernalized with time up to as long as about two months. Full
vernalisation requires up to about 50 days of treatment between – 2°C and about 12°C. If
vernalisation is followed by high temperature treatment at about 40°C for a minimum of two days,
the vernalizing stimulus is lost. This is known as devernalisation. Devernalised plants can,
however, be vernalized again.
(ii) Actively Dividing Cells:
Vernalisation does not occur in dry seeds. The seeds must be germinated so that they contain an
active embryo. For this the seeds are moistened before exposing them to low temperature. In a
whole plant, an active meristem is required.
(iii) Water:
Proper protoplasmic hydration is must for perceiving the stimulus of vernalisation.
(iv) Aerobic Respiration and
(v) Proper Nourishment
Mechanism of Vernalisation and Induction of Flowering:
The stimulus received by the actively dividing cells of shoot or embryo tip travels to all parts of
the plant and prepare it to flower. The stimulus has been named as vernalin. Melchers (1936,
1937) demonstrated in henbane plants (Hyoscyamus niger) translocation of vernalisation stimulus
takes place through a graft union (Fig. 5.7). If leaf or stem of a vernalised plant is grafted on an
un-vernalised henbane plant the latter plant will flower.
The stimulus was found to be non-specific, i.e., can pass across a graft between plants of different
species. It was Melchers who, for the first time, suggested that a substance which he called
vernalin was produced during the process of vernalisation. Attempts to isolate and chemically
identify vernalin have not succeeded as yet. However, Lang et. at. (1957) have demonstrated that
treatment with Gibberellic acid (GA), a plant hormone, and substitute for cold treatment in some
species of plants.
The formation of vernalin is not enough to bring about vernalisation. In addition, a suitable day
length is also necessary. It is postulated that in the appropriate photoperiod, either vernalin is
converted into florigen or vernalin regulates the synthesis of florigen from precursors. Florigen
then induces the vegetative meristems to switch over to reproductive development. This initiates
the process of flower differentiation.

86
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Vernalisation simply prepares a plant to flower. It makes the plant perceptive to the stimulus;
however, it itself does not function as a stimulus. Photoperiodism, on the contrary, not only
provides the stimulus for flowering but also induces it.
Advantages/Importance of Vernalisation:
(i) Vernalisation can help in shortening the juvenile or vegetative period of plant and induce early
flowering. It is applicable to not only temperate plants but also to tropical plants, e.g., wheat, rice,
millets, cotton.
(ii) It increases yield, resistance to cold and diseases.
(iii) ‘Kernel wrinkles’ of Triticale can be removed by vernalisation.
(iv) It enables the biennials to behave as annuals.
(v) Plants can be grown in such regions where normally they do not grow.

PHOTOPERIODISM

Photoperiodism is the physiological reaction of organisms to the length of day or night. It occurs
in plants and animals. Photoperiodism can also be defined as the developmental responses of plants to
the relative lengths of light and dark periods.
Plants
Many flowering plants (angiosperms) use a photoreceptor protein, such
is Phytochrome or cryptochrome, to sense seasonal changes in night length, or photoperiod, which
they take as signals to flower. In a further subdivision, obligate photoperiodic plants absolutely require
a long or short enough night before flowering, whereas facultative photoperiodic plants are more
likely to flower under one condition.
In 1920, W. W. Garner and H. A. Allard published their discoveries on photoperiodism and felt it was
the length of daylight that was critical, but it was later discovered that the length of the night was the
controlling factor. Photoperiodic flowering plants are classified as long-day plants or short-day
plants even though night is the critical factor because of the initial misunderstanding about daylight
being the controlling factor. Along with long-day plants and short-day plants, there are plants that fall
into a "dual-day length category". These plants are either long-short-day plants (LSDP) or short-long-
day plants (SLDP). LSDPs flower after a series of long days followed by short days whereas SLDPs
flower after a series of short days followed by long days. Each plant has a different length critical
photoperiod, or critical night length.
Modern biologists believe that it is the coincidence of the active forms of phytochrome or
cryptochrome, created by light during the daytime, with the rhythms of the circadian clock that allows
87
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

plants to measure the length of the night. Other than flowering, photoperiodism in plants includes the
growth of stems or roots during certain seasons and the loss of leaves. Artificial lighting can be used
to induce extra-long days.
Long-day plants
Long-day plants flower when the night length falls below their critical photoperiod. These plants
typically flower in the northern hemisphere during late spring or early summer as days are getting
longer. In the northern hemisphere, the longest day of the year (summer solstice) is on or about 21
June. After that date, days grow shorter (i.e. nights grow longer) until 21 December (the winter
solstice). This situation is reversed in the southern hemisphere (i.e., longest day is 21 December and
shortest day is 21 June).
Some long-day obligate plants are:
 Carnation (Dianthus)
 Henbane (Hyoscyamus)
 Oat (Avena)
Some long-day facultative plants are:
 Pea (Pisum sativum)
 Barley (Hordeum vulgare)
 Lettuce (Lactuca sativa)
 Wheat (Triticum aestivum)
Short-day plants
Short-day plants flower when the night lengths exceed their critical photoperiod.] They cannot flower
under short nights or if a pulse of artificial light is shone on the plant for several minutes during the
night; they require a continuous period of darkness before floral development can begin. Natural
nighttime light, such as moonlight or lightning, is not of sufficient brightness or duration to interrupt
flowering.
In general, short-day ([Link]-night) plants flower as days grow shorter (and nights grow longer) after
21 June in the northern hemisphere, which is during summer or fall. The length of the dark period
required to induce flowering differs among species and varieties of a species.
Photoperiodism affects flowering by inducing the shoot to produce floral buds instead of leaves and
lateral buds.
Some short-day facultative plants are:
 Marijuana (Cannabis)
 Cotton (Gossypium)
 Rice (Oryza)
 Jowar (Sorghum bicolor)
 Green Gram (Mung bean, Vigna radiata)
 Soybeans (Glycine max)
Day-neutral plants
Day-neutral plants, such as cucumbers, roses, and tomatoes, do not initiate flowering based on
photoperiodism. Instead, they may initiate flowering after attaining a certain overall developmental
stage or age, or in response to alternative environmental stimuli, such as vernalisation (a period of low
temperature)

88
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 12

PLANT GROWTH HORMONES

 Plant hormones (also known as phytohormones) are chemicals that regulate plant growth. In
the United Kingdom, these are termed 'plant growth substances'.
 Plant hormones are signal molecules produced within the plant, and occur in extremely
low concentrations. Hormones regulate cellular processes in targeted cells locally and, moved to
other locations, in other functional parts of the plant.
 Hormones also determine the formation of flowers, stems, leaves, the shedding of leaves, and the
development and ripening of fruit. Plants, unlike animals, lack glands that produce
and secrete hormones. Instead, each cell is capable of producing hormones. Plant hormones shape
the plant, affecting seed growth, time of flowering, the sex of flowers, senescence of leaves, and
fruits.
 They affect which tissues grow upward and which grow downward, leaf formation and stem
growth, fruit development and ripening, plant longevity, and even plant death.
 Hormones are vital to plant growth, and, lacking them, plants would be mostly a mass of
undifferentiated cells. So they are also known as growth factors or growth hormones.
 The term 'Phytohormone' was coined by Thimann in 1948
Characteristics
The word hormone is derived from Greek, meaning set in motion. Plant hormones affect gene
expression and transcription levels, cellular division, and growth.
 They are naturally produced within plants, though very similar chemicals are produced by fungi
and bacteria that can also affect plant growth.] A large number of related chemical compounds
are synthesized by humans.
 They are used to regulate the growth of cultivated plants, weeds, and in vitro-grown plants and
plant cells; these manmade compounds are called plant growth regulators or PGRs for short. Early
in the study of plant hormones, "phytohormone" was the commonly used term, but its use is less
widely applied now.
 Plant hormones are not nutrients, but chemicals that in small amounts promote and influence the
growth,development, and differentiation of cells and tissues. The biosynthesis of plant hormones
within plant tissues is often diffuse and not always localized.
 Plants lack glands to produce and store hormones, because, unlike animals—which have two
circulatory systems (lymphatic and cardiovascular) powered by a heart that moves fluids around
the body—plants use more passive means to move chemicals around their bodies.
 Plants utilize simple chemicals as hormones, which move more easily through their tissues. They
are often produced and used on a local basis within the plant body.
 Plant cells produce hormones that affect even different regions of the cell producing the hormone.
 Hormones are transported within the plant by utilizing four types of movements. For localized
movement, cytoplasmic streaming within cells and slow diffusion of ions and molecules between
cells are utilized.
 Vascular tissues are used to move hormones from one part of the plant to another; these
include sieve tubes or phloem that move sugars from the leaves to the roots and flowers,
and xylem that moves water and mineral solutes from the roots to the foliage.

89
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The five classical hormones:

 Auxin
 Cytokinin
 Gibberelin
 Abscisic acid
 Ethylene

These are all small molecules ranging from 28 Da (ethylene) to 346 Da (GA), all synthesized by the
plant, and all active at 10-6 to 10-8 M. They are distributed within tissues from cell to cell, as in the
case of auxin, via vascular bundles (as in the case of cytokinin), or via the intercellular space
(ethylene).

Additional substances gaining recognition as PGRs:

 Polyamines
 Jasmonates
 Salicylic acid
 Brassinosteroids
 Others

What a PGR is not:

 Inorganic compounds such as Ca2+ and K+, even though these compounds cause physiological
responses and can be moved throughout the plant, because they are not synthesized by the
plant.
 Sucrose is not a hormone even though it is synthesized, translocated, and stimulates a response
(growth) because it is only effective at high concentrations. This is also true for other sugars,
amino acids, organic acids, and other metabolites that are usually present in concentrations
ranging from >1mM up to 50 mM or higher.
 Technically, synthetic growth regulatory substances such as 2,4-D even though its structure is
similar to that of auxin.

III. How PGRs Have Been Identified and Characterized

The presence of PGRs in plants was first detected or inferred by observed plant behavior under
various conditions. However, much of what was known about PGRs until recent years was largely
determined empirically because they were very difficult to study.

Why?

 Compartmentalization of PGRs in cells or organelles.

 PGR levels are by definition very low. This has historically made endogenous detection
difficult, although detection methods have vastly improved over the past 15 years due to
improved instrumentation, e.g. GC/MS-detection, use of immunological methods.
 The site of biosynthesis and the site of action of a plant growth regulator may be different.
90
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Because the site of action may be in specialized cells, located physically within a variety of
other cells and tissues, it may be difficult to correlate levels of PGRs with particular
physiological responses.
 A potential PGR may have an effect when applied exogenously, however, this does not mean
that the compound normally acts to elicit the same effect in the plant.
 A PGR may be applied but may not penetrate or be transported like the endogenous PGR, so it
does not get to the site of action.
 High concentrations of PGRs may cause non-physiological effects.
 Different tissues, cells, respond differently to the same PGR.
 Different processes are sensitive to different levels of PGR, even in the same cell.
 Changes in environment or development alter responses to a PGR.

Thus, the problems of working with biosynthetic pathways for hormones, which represent a minor
portion of carbon flux throughout plant, are distinct. Reactions involving intermediates must be
detected using radioisotopes or immunological techniques to get sufficient sensitivity.

When hormones control gene expression, it is likely that gene regulation is affected by certain
enzymes after initial hormone binding. Genes may be altered by secondary and tertiary messengers of
a cellular cascade as well. Hormones may indirectly control gene expression through these enzymes
and messengers by acting during transcription, during mRNA processing, on mRNA stability, at
translation, and/or post-translationally. A single aspect of growth and development can be influenced
by several hormones: a particular response probably results from a changing ratio of hormones rather
than from the presence or absence of an individual hormone. Further, physiological responses elicited
by hormones are strongly influenced by nonhormonal factors like light of a certain wave length,
temperature, etc. PGR actions may be synergistic or antagonistic. However, it should be emphasized
that there is still much to learn about PGRs and their actions.

i. Auxins

Auxin was the first plant hormone to be discovered.

In early plant physiological studies, some by Charles and Francis Darwin (1880), it was found that a
growing grass coleoptile that was illuminated on one side grew towards the source of illumination.
Anatomical studies showed that this was due to an elongation of the cells on the side away from the
light. This phenomenon was termed "phototrophism". The phototropic reaction does not happen if the
coleoptile’s tip is removed, though it can be induced again by replacement of the tip. This indicates
the existence of a substance that moves from tip to bottom (in a basipetal direction) and that causes the
elongation.

In 1985 Salkowski discovered IAA in fermentation media. It would be almost 50 years before the
same compound was identified in plant tissue. The Danish botanist P. Boysen-Jensen in 1913 inserted
a mica sheet into the shielded side of a coleoptile, thus separating the coleoptile’s tip from the tissue
below. This interrupted the phototropic reaction. The bending occurred when the mica sheet was
inserted into the illuminated side or along the coleoptile’s vertical axis.

91
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Went called the effector auxin, derived from the Greek word auxein, which means to grow. We now
know this hormone as indole-3-acetic acid (IAA). Compounds are generally characterized as auxins if
they are synthesized by the plant, they induce cell elongation in stems, and they otherwise resemble
IAA in physiological activity. Auxins typically affect other processes in addition to elongation of stem
cells but this characteristic is considered integral to all auxins and thus helps "define" the hormone.
IAA is chemically similar to the amino acid tryptophan, which is generally accepted to be the
molecule from which IAA is derived, although there are apparent exceptions.

It is now known that auxin is a collective name for several similar compounds. To be active, an auxin
compound has to have three structural properties:

 The molecule has to contain a ring system with at least one double bond.
 The double bond has to be adjacent to a side chain.
 A carboxyl group that is separated by one or two C-atoms is required.

These conditions led to several clues about the structure of the binding site(s); the receptor must have
two separate contact sites.

Transport of auxin within the plant is an active and polar process. There are at least 6 pieces of
evidence for this:

 Transport always occurs in a polar direction, e.g. unidirectionally down the plant stem through
parenchyma cells (appaerently auxin does not move through vascular tissue).
 The transport velocity is higher than is expected from simple diffusion.
 Transport can occur against a concentration gradient.
 Transport is energy-consuming and is drastically reduced in the absence of oxygen.
 The transport system is substrate-specific. It transports certain auxin molecules like IAA or
naphthylacetic acid faster than 2,4-dichlorophenoxy acetic acid, for example.
 The transport system can be blocked by specific inhibitors.

Auxins are believed to affect several different primary processes of the cell, i.e. to:

 Increase the rate of transcription (RNA synthesis).

 Control the activity of certain enzymes.
 Influence ion pumps within the membrane.

Auxin is produced primarily in apical and root meristems, young leaves, and seeds in developing
fruits.

Auxin

 Stimulates cell elongation.

 The auxin supply from the apical bud suppresses growth of lateral buds. Apical dominance is
the inhibiting influence of the shoot apex on the growth of axillary buds. Removal of the apical
bud results in growth of the axillary buds. Replacing the apical bud with a lanolin paste
containing IAA restores the apical dominance. The mechanism involves another hormone -
92
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

ethylene. Auxin (IAA) causes lateral buds to make ethylene, which inhibits growth of the
lateral buds.
 Differentiation of vascular tissue (xylem and phloem) is stimulated by IAA.
 Auxin stimulates root initiation on stem cuttings and lateral root development in tissue culture
(adventitious rooting).
 Auxin mediates the tropistic response of bending in response to gravity and light (this is how
auxin was first discovered).
 Auxin has various effects on leaf and fruit abscission, fruit set, development, and ripening, and
flowering, depending on the circumstances.

Many synthetic auxins have been developed that are used for a variety of purposes, e.g.
indole-3-butyric acid (IBA)(root initiation) and naphthaleneacetic acid (NAA)(apple fruit
thinning). 2,4 dichloro-phenoxy acetic acid (2,4-D) and 2,4,5-trichlorphenoxy acidic acid
(2,4,5-T) proved to be herbicides selectively affecting dicots.

2,4-D increases the rate of DNA, RNA and protein synthesis, so that susceptible plants literally grow
themselves to death. Treated plants display abnormal growth of shoots, breakdown of chlorophyll
(bleaching), and dying of roots. 2,4,5-T is especially toxic to perennial woody plants and is therefore
most often used in forestry. Its is less easily degraded than 2,4-D. IAA, in contrast, is very easily
degraded and is consequently of no use as a herbicide. When these compounds are used in tissue
culture, they are used in very small amounts.

Although it has not been clearly shown that these chemicals cause injury to humans or animals, there
is no doubt that a by-product of the synthesis of 2,4,5-T, tetrachlorodibenzoparadioxin (TCDD) (also
called dioxin), is highly toxic. The Agent Orange used in the Vietnam war as a defoliation agent was
about 50% 2,4-D, but it was the dioxin in the agent that was subsequently shown to cause the illnesses
of individuals exposed.

93
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Auxins are frequently used in tissue culture, for a variety of purposes. We will cover specifics when
we discuss the various tissue culture procedures.

ii. Cytokinins

Researchers tried for many years to culture plant tissue on artificial nutrient medium. The first
approaches go back to the Austrian plant anatomist G. Haberlandt (1854-1945, professor at Graz, later
at Berlin). At first, determining the composition of a suitable nutrient medium posed large problems.
The Dutch plant physiologist van Overbeek discovered in 1941 that plant embryos grow faster when
given coconut milk (the endosperm of the coconut seed). Then, Skoog and Miller, while working with
tissue culture methods for plants, used coconut milk in media formulations. While trying to determine
what substance in the milk was necessary for growth, they added various chemicals they had available
in the lab to media. Herring-sperm DNA appeared to stimulate growth when they first tried it, but on a
second attempt, it produced no effect. They discovered a breakdown product similar to adenine in the
first bottle, which was responsible for the stimulation.

Kinetin was the first cytokinin discovered and was so named because of its ability to promote
cytokinesis (cell division). However, kinetin is not made in plants, and is therefore considered a
synthetic PGR. The most common form of naturally occurring cytokinin in plants is zeatin, which was
isolated from maize (Zea mays) in 1961 (Miller, Letham).

Besides increasing the rate of DNA replication, cytokinins also increase the general rate of RNA and
protein synthesis (binds to ribosomes, regulates protein synthesis). Cytokinins are made via the same
pathway that leads to gibberellin production: the mevalonate pathway. These compounds have a
structure resembling that of adenine. Like auxins, cytokinins can glycosylated or bound to an amino
acid or a protein, thereby being at least temporarily inactive. Cytokinins have been found in almost all
higher plants as well as mosses, fungi (Witch’s broom), and bacteria (Agrobacterium). Today there are
more than 200 natural and synthetic cytokinins that have been identified.

Effects of cytokinins

 Stimulate cell division (cytokinesis).

 Stimulate morphogenesis (shoot initiation/bud formation) in tissue culture.
 Stimulate the growth of lateral (or adventitious) buds - release of apical dominance.
 Stimulate leaf expansion resulting from cell enlargement.
 May enhance stomatal opening in some species.

94
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Promotes the conversion of etioplasts into chloroplasts via stimulation of chlorophyll

synthesis.
 Stimulate the dark-germination of light-dependent seeds.
 Delays senescence.
 Promotes some stages of root development.

Cytokinins are produced mainly in root meristems, young leaves, young fruits, and seeds. They enter
the shoot organs via the xylem, translocated from the roots. Organs that are cut off from a continuous
cytokinin supply like cut shoots age faster than those that are connected to their roots. The addition of
kinetin can stop senescence, as can the development of adventitious roots and thus a new supply
of cytokinins.

Thus, cytokinins like auxins have many modes of action. The variety of hormonally induced
phenomena points to the existence of different receptors.

Commercial uses: applied as Kinetin, BA, or zeatin conjugates

 axillary bud growth in orchids, day lilies

 antioxident (browning preventer) in cut salads
 mixed with GA as a fruit size simulator

iii. Gibberellins

Unlike auxins, which are classified on the basis of function, gibberellins are classified on the basis of
structure as well as function. Gibberellins are diterpenes synthesized from acetyl CoA via the
mevalonic acid pathway. They all have either 19 or 20 carbon units grouped into either four or five
ring systems. All gibberellins are acidic compounds and are therefore also called gibberellic acids
(GA) with a different subscript to distinguish between them. GA3 has historically been called
gibberellic acid but the term is also often used in describing all gibberellins.

In 1926, the Japanese scientist E. Kurosawa was studying a rice disease that is known as ‘foolish
seedling’. The affected plants grow extremely fast, look spindly and pale, and break off easily.
Kurosawa determined that the reason for this abnormal growth was a substance that is secreted by a
parasitic fungus (Fusarium moniliforme = Gibberella fujikuroi). The compound was termed
gibberellin.

During the thirties was gibberellin isolated and crystallized by Japanese scientists from Tokyo (Yabuta
and Sumiki ), and then was almost forgotten in the following years. In 1956, C. A. West and B. O.
Phinney isolated gibberellins from Phaseolus vulgaris and other plants, showing that these compounds
are wide-spread in the plant kingdom. Today are more than 110 (125?) different gibberellins known
(GA1, GA2,….GA3, GA4…..GA110) that differ little chemically but very much in their biological
effects.

Roughly 30 percent of all known gibberellins are biologically active. All higher plants contain
presumably at least one, but usually several active and inactive gibberellins that exist in different
concentrations depending on the respective tissue. They have also been isolated from lower plants
such as mosses and algae, at least two fungal species and most recently from two bacterial species.

95
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Effects of Gibberellic Acid

Young, growing meristematic tissue, embryos, apical root cells, young fruits, as well as unripe or
germinating seeds are all rich in gibberellins.

 Stimulates stem elongation by stimulating cell division and elongation. GA controls internode
elongation in the mature regions of plants. Dwarf plants do not make enough active forms of
GA.
 Flowering in biennial plants is controlled by GA. Biennials grow one year as a rosette and after
the winter, they bolt (rapid expansion of internodes and formation of flowers).
 Breaks seed dormancy in some plants that require stratification or light to induce germination.
 Stimulates a-amylase production in germinating cereal grains for mobilization of seed reserves.
 Juvenility refers to the different stages that plants may exist in. GA may help determine
whether a particular plant part is juvenile or adult.
 Stimulates germination of pollen and growth of pollen tubes.
 Induces maleness in dioecious flowers (sex expression).
 Can cause parthenocarpic (seedless) fruit development or increase the size of seedless fruit
(grapes).
 Can delay senescence in leaves and citrus fruits.
 May be involved in phytochrome responses.

Certain commercial chemicals that are used to stunt growth do so in part because they block the
synthesis of gibberellins. Some of these chemicals are Phosphon D, Amo-1618, Cycocel (CCC),
ancymidol, and paclobutrazol.

iv. Abscisic Acid

The phytohormone abscisic acid (ABA) plays regulatory roles in a host of physiological processes in
all higher as well as lower plants. Abscisic acid mediates stress tolerance responses in higher plants, is
a key signal compound that regulates stomatal aperture and, in concert with other plant signaling
compounds, is implicated in mediating responses to pathogens and wounding. It has therefore been
called the stress hormone.

ABA is a sesquiterpenoid (15-carbon) compound. It is believed that biosynthesis occurs indirectly

through the production of carotenoids in plastids. Carotenoids are pigments produced by plastids that
have 40 carbons. Biosynthesis of ABA is believed to occur as follows: The 40 carbon carotenoid
violaxanthin is isomerized and then split via an isomerase reaction followed by an oxidation reaction.
One molecule of xanthonin is produced from one molecule of violaxanthonin and it is uncertain what
happens to the remaining biproduct. The one molecule of xanthonin produced is unstable and
spontaneously changed to ABA aldehyde. Further oxidation results in ABA.

The transport of ABA can occur in both xylem and phloem tissues. It can also be translocated through
parenchyma cells. The movement of abscisic acid in plants does not exhibit polarity like that of
auxins. ABA is capable of moving both up and down the stem.

96
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Effects of ABA

 Involved in abscission of buds, leaves, petals, flowers, and fruits in many, if not all, instances,
as well as in dehiscence of fruits.
 Production is accentuated by stresses such as water loss and freezing temperatures.
 Involved in bud dormancy
 Prolongs seed dormancy and delays germination (vivipary).
 Inhibits elongation.
 Promotes stomatal closure.
 ABA is implicated in the control of elongation, lateral root development, and geotropism, as
well as in water uptake and ion transport by roots.
 ABA coming from the plastids promotes the metabolism of ripening.
 Promotes senescence.
 Can reverse the effects of growth stimulating hormones.

v. Ethylene

Unlike the rest of the plant hormone compounds, ethylene is a gas. Like abscisic acid, it is the only
member of its class.

Ethylene has been used in practice since the ancient Egyptians, who would gas figs in order to
stimulate ripening. The ancient Chinese would burn incense in closed rooms to enhance the ripening
of pears. In 1864, leaks of gas from street lights were discovered to cause stunting of growth, twisting
of plants, and abnormal thickening of stems (termed the triple response) In 1901, a Russian scientist
named Dimitry Neljubow showed that the active component was ethylene. Ethylene is produced in all
higher plants.

In higher plants, ethylene is produced from L-methionine. Methionine is activated by ATP to form S-
adenosylmethionine through the catalytic actvity of S- adenosylmethionine synthetase. The next step
produces the non-protein amino acid 1-aminocyclopropane-1-carboxylic acid (ACC). It is catalyzed
by ACC synthase with pyridoxal phosphate acting as a co-factor. Formation of ACC is the rate-
limiting step in ethylene biosynthesis. ACC synthase is encoded by a medium-size multigene family.
Various signals, which influence ethylene synthesis, result in increased expression of single members
of the ACC synthase gene family. Production of ethylene from ACC is catalyzed by ACC oxidase.
This reactions is oxygen-dependent. At anaerobic conditions ethylene formation is completely
suppressed. Fe2+ is a co-factor and ascorbate a cosubstrate; CO2 was shown to activate ACC oxidase.
ACC oxidases are encoded by small gene families in plants.

The concentration of ethylene in a plant tissue is dependent on the rate of biosynthesis and on
diffusion of the gas. Ethylene is neither actively transported nor degraded. Induction of ethylene
synthesis by signals such as auxin or wounding usually occurs through activation of ACC synthase.
ACC oxidase activity on the other hand is constitutively present in most vegetative plant tissues. In
some cases, further induction of ACC oxidase by ethylene is observed.

Ethylene is produced in many plant tissues. In maturing fruits, ethylene synthesis is autocatalytically
enhanced, i.e., ethylene induces its own biosynthesis. The self-enhancing synthesis and diffusion of

97
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

the gaseous hormone throughout the fruit accelerate ripening and contribute to a synchronized
ripening process.
Ethylene binds to a protein in the endoplasmic reticulum?

Ethylene action very often involves transcriptional activation. In recent years, much has been learned
about the transduction pathway from ethylene perception to gene activation. Ethylene is bound by a
receptor that is membrane-localized. The N-terminal domain of the receptor protein is responsible for
binding of ethylene. The intracellular portion of the protein is a protein kinase which is activated upon
binding of ethylene. Such receptor protein kinases are termed two-component-systems because they
are generally composed of a sensor, in this case the ethylene binding site, and a response regulator.
Two-component-receptors have first been characterized in bacteria, but are now also known in plants.
The first two-component-receptor to be discovered in plants was the ethylene receptor ETR1 (ethylene
resistant 1) from Arabidopsis thaliana L. Transduction of the ethylene signal is thought to be achieved
through a series of phosphorylations that are carried out by a cascade of protein kinases similar to the
MAP (mitogen activated protein kinase) kinase pathway and probably through other, as yet less well
defined, steps. Finally, a transcription factor that is present in the cell is activated, leading to induction
of one or more early gene(s). One such early gene is itself a transcriptional activator that induces late
genes. These late-induced genes may encode enzymes that degrade the middle lamellae or the cell
wall during fruit ripening and abscission They may encode for proteins involved in pathogen defense.
Or they may encode for proteins required for other ethylene responses.

Ethylene acts as a negative regulator of the signaling pathway. This means that the signal pathway is
turned on in the absence of ethylene and is shut down when ethylene is present. Shutting down of the
pathway induces an ethylene response. As a result of this negative regulation, mutations in the
ethylene receptor are perceived as dominant gain-of-function mutations.

Components of the ethylene signal transduction pathway have been identified by a mutational
approach with Arabidopsis. The initial steps in ethylene signal transduction are shared for all these
responses. Cloning of these genes has started to reveal how ethylene is perceived in plants. ETR1
(along with ERS1, ETR2, ERS2, and EIN4) are all related to the two-component signaling proteins
from bacteria. CTR1 is related to the Raf-like serine/threonine protein kinases from mammals. EIN2 is
related to the Nramp family of metal transporters. EIN3, EIL1, EIL2, and ERF1 are transcription
factors.

Effects of ethylene

 Production stimulated during ripening, flooding, stress, senescence, mechanical damage,

infection.
 Entral regulator of cell death programs in plants (apoptosis).
 Stimulates the release of dormancy.
 Stimulates shoot and root growth and differentiation (triple response)
 Regulates ripening of climacteric fruits.
 May have a role in adventitious root formation.
 Stimulates leaf and fruit abscission.
 Flowering in most plants is inhibited by ethylene. Mangos, pineapples and some ornamentals
are stimulated by ethylene.
 Induction of femaleness in dioecious flowers.
98
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Stimulates flower opening.

 Stimulates flower and leaf senescence.

Commercial Uses - Ethylene Application (applied as ethylene gas or ethephon or ethrel sprays)

 flower initiation (pineapples)

 stimulation of ripening (bananas, tomatoes)
 degreening of citrus
 abscission induction prior to mechanical harvesting (cherries)

Commercial Uses - Ethylene Inhibition (chemical removal or low atmosphere)

 long term CA apple storage

 cut flower maintenance (silver thiosulfate)
 hypobaric storage of many fruits, vegetables, flowers

Silver thiosulfate (0.02 M)is commonly used to block the action of ethylene in plant cell cultures,
where it has been shown to increase due to wounding or the presence of auxins.

vi. Other Potential PGRs

A. Brassinosteroids

There are approximately 60 naturally occurring polyhydroxy steroids known as brassinosteroids

(BRs). They are named after the first one identified, brassinolide, which was isolated from rape in
1979. They appear to be widely distributed in the plant kingdom.

Biological Effects of BRs

 Promote shoot elongation at low concentrations

 Strongly inhibit root growth and development
 Promote ethylene biosynthesis and epinasty
 Interfere with ecdysteroids (moulting hormones) in insects
 Have had contradictory effects in tissue culture. 24-Epibrassinolide has been shown to mimic
culture conditioning factors and to be synergistic with these factors in promoting carrot cell
growth. However, in transformed tobacco cells brassinosteroids in low concentrations
significantly inhibited cell growth.
 Enhance xylem differentiation
 Decrease fruit abortion and drop
 Enhance resistance to chilling, disease, herbicide, and salt stress

Hormones and plant propagation

Synthetic plant hormones or PGRs are commonly used in a number of different techniques
involving plant propagation from cuttings, grafting, micropropagation, and tissue culture.
The propagation of plants by cuttings of fully developed leaves, stems, or roots is performed by
gardeners utilizing auxin as a rooting compound applied to the cut surface; the auxins are taken into
the plant and promote root initiation. In grafting, auxin promotes callus tissue formation, which joins
the surfaces of the graft together. In micropropagation, different PGRs are used to promote

99
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

multiplication and then rooting of new plantlets. In the tissue-culturing of plant cells, PGRs are used to
produce callus growth, multiplication, and rooting.
Seed dormancy
Plant hormones affect seed germination and dormancy by acting on different parts of the seed.
Embryo dormancy is characterized by a high ABA:GA ratio, whereas the seed has a high ABA
sensitivity and low GA sensitivity. In order to release the seed from this type of dormancy and initiate
seed germination, an alteration in hormone biosynthesis and degradation toward a low ABA/GA ratio,
along with a decrease in ABA sensitivity and an increase in GA sensitivity, must occur.
ABA controls embryo dormancy, and GA embryo germination. Seed coat dormancy involves the
mechanical restriction of the seed coat. This, along with a low embryo growth potential, effectively
produces seed dormancy. GA releases this dormancy by increasing the embryo growth potential,
and/or weakening the seed coat so the radical of the seedling can break through the seed coat.
Different types of seed coats can be made up of living or dead cells, and both types can be influenced
by hormones; those composed of living cells are acted upon after seed formation, whereas the seed
coats composed of dead cells can be influenced by hormones during the formation of the seed coat.
ABA affects testa or seed coat growth characteristics, including thickness, and effects the GA-
mediated embryo growth potential. These conditions and effects occur during the formation of the
seed, often in response to environmental conditions. Hormones also mediate endosperm dormancy:
Endosperm in most seeds is composed of living tissue that can actively respond to hormones
generated by the embryo. The endosperm often acts as a barrier to seed germination, playing a part in
seed coat dormancy or in the germination process. Living cells respond to and also affect the
ABA:GA ratio, and mediate cellular sensitivity; GA thus increases the embryo growth potential and
can promote endosperm weakening. GA also affects both ABA-independent and ABA-inhibiting
processes within the endosperm.
TEST YOURSELF
1. Dwarfness can be controlled by treating the plant with
(a) cytokinin
(b) gibberellic acid
(c) auxin
(d) antigibberellin.
2. Which is produced during water stress that brings stomatal closure?
(a) ethylene
(b) abscisic acid
(c) ferulic acid
(d) coumarin.
3. The hormone produced during adverse environmental conditions is
(a) benzyl aminopurine
(b) bichlorophenoxy acetic acid
(c) ethylene
(d) abscisic acid.
4. The regulator which retards ageing/senescence of plant parts is
(a) cytokinin
(b) auxin
(c) gibberellin
(d) abscisic acid.
5. The movement of auxin is largely
(a) centripetal
(b) basipetal
(c) acropetal
(d) both ‘a’ and ‘c’.
100
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 13

GAMETE FORMATION
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the
male gametes (sperm cells). Pollen grains have a hard coat that protects the sperm cells during the
process of their movement from the stamens to the pistil of flowering plants or from the male cone to
the female cone of coniferous plants. When pollen lands on a compatible pistil or female cone (i.e.,
when pollination has occurred), it germinates and produces a pollen tube that transfers the sperm to
the ovule (or female gametophyte).
Individual pollen grains are small enough to require magnification to see detail. The study of pollen is
called palynology.
Pollen itself is not the male gamete. Each pollen grain contains vegetative (non-reproductive) cells
(only a single cell in most flowering plants but several in other seed plants) and a generative
(reproductive) cell containing two nuclei: a tube nucleus (that produces the pollen tube) and a
generative nucleus (that divides to form the two sperm cells). The group of cells is surrounded by a
cellulose-rich cell wall called the intine, and a resistant outer wall composed largely of sporopollenin
called the exine.

POLLEN DEVELOPMENT

a) The anthers comprise pollen sacs(usually four) which contain a mass of diploid pollen mother
cells. Each pollen mother cell undergoes meiosis to form a tetrad of four haploid cells.
b) The cells round off and are called microspores.
c) The nucleus divides by mitosis to give tube nucleus and generative nucleus.
d) The wall thickens and forms an inner layer, intine and an often highly sculptured outer layer,the
exine.
e) When transferred to the stigma of a plant of the same species,the pollen grain germinates to
produce a pollen tube.
f) The tube nucleus moves down the tube first, followed by generative nucleus which soon divides
mitotically to give two male nuclei.

OVULE

In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It
consists of three parts: The integument(s), forming its outer layer(s), the nucellus (or remnant of the
megasporangium), and the female gametophyte (formed from a haploid megaspore) in its center. The
101
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

female gametophyte — specifically termed a megagametophyte— is also called the embryo sac in
angiosperms. The megagametophyte produces an egg cell (or several in some groups) for the purpose
of fertilization.

Ovule parts and development

The ovule appears to be a megasporangium with integuments surrounding it. Ovules are initially
composed of diploid maternal tissue, which includes a megasporocyte (a cell that will undergo meiosis
to produce megaspores). Megaspores remain inside the ovule and divide by mitosis to produce the
haploid female gametophyte or megagametophyte, which also remains inside the ovule. The remnants
of the megasporangium tissue (the nucellus) surround the megagametophyte. Megagametophytes
produce archegonia (lost in some groups such as flowering plants), which produce egg cells. After
fertilization, the ovule contains a diploid zygote and then, after cell division begins, an embryo of the
next sporophyte generation. In flowering plants, a second sperm nucleus fuses with other nuclei in the
megagametophyte forming a typically polyploid (often triploid) endosperm tissue, which serves as
nourishment for the young sporophyte.

Integuments, micropyle and chalaza

An integument is a protective cell layer surrounding the ovule. Gymnosperms typically have one
integument (unitegmic) while angiosperms typically have two (bitegmicThe integuments develop into
the seed coat when the ovule matures after fertilization.

The integuments do not enclose the nucellus completely but retain an opening at the apex referred to
as the micropyle. The micropyle opening allows the pollen (a male gametophyte) to enter the ovule for
fertilization. In gymnosperms (e.g., conifers), the pollen is drawn into the ovule on a drop of fluid that
exudes out of the micropyle, the so-called pollination drop mechanism.[3] Subsequently, the
micropyle closes. In angiosperms, only a pollen tube enters the micropyle. During germination, the
seedling's radicle emerges through the micropyle.

Located opposite from the micropyle is the chalaza where the nucellus is joined to the integuments.
Nutrients from the plant travel through the phloem of the vascular system to the funiculus and outer
integument and from there apoplastically and symplastically through the chalaza to the nucellus inside
the ovule. In chalazogamous plants, the pollen tubes enter the ovule through the chalaza instead of the
micropyle opening.

Nucellus, megaspore and perisperm

The nucellus (plural: nucelli) is part of the inner structure of the ovule, forming a layer of diploid
(sporophytic) cells immediately inside the integuments. It is structurally and functionally equivalent to
the megasporangium. In immature ovules, the nucellus contains a megasporocyte (megaspore mother
cell), which undergoes sporogenesis via meiosis. In the megasporocyte of thaliana, meiosis depends
on the expression of genes that facilitate DNA repair and homologous recombination.[5]

In gymnosperms, three of the four haploid spores produced in meiosis typically degenerate, leaving
one surviving megaspore inside the nucellus. Among angiosperms, however, a wide range of variation
exists in what happens next. The number (and position) of surviving megaspores, the total number of

102
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

cell divisions, whether nuclear fusions occur, and the final number, position and ploidy of the cells or
nuclei all vary.

After fertilization, the nucellus may develop into the perisperm that feeds the embryo. In some plants,
the diploid tissue of the nucellus can give rise to the embryo within the seed through a mechanism of
asexual reproduction called nucellar embryony.

Megagametophyte

In flowering plants, the megagametophyte (also referred to as the embryo sac) is much smaller and
typically consists of only seven cells and eight nuclei. This type of megagametophyte develops from
the megaspore through three rounds of mitotic divisions. The cell closest to the micropyle opening of
the integuments differentiates into the egg cell, with two synergid cells by its side that are involved in
the production of signals that guide the pollen tube. Three antipodal cells form on the opposite
(chalazal) end of the ovule and later degenerate. The large central cell of the embryo sac contains two
polar nuclei.

Zygote, embryo and endosperm

The pollen tube releases two sperm nuclei into the ovule. In gymnosperms, fertilization occurs within
the archegonia produced by the female gametophyte. While it is possible that several egg cells are
present and fertilized, typically only one zygote will develop into a mature embryo as the resources
within the seed are limited.

In flowering plants, one sperm nucleus fuses with the egg cell to produce a zygote, the other fuses
with the two polar nuclei of the central cell to give rise to the polyploid (typically triploid) endosperm.
This double fertilization is unique to flowering plants, although in some other groups the second
sperm cell does fuse with another cell in the megagametophyte to produce a second embryo. The plant
stores nutrients such as starch, proteins, and oils in the endosperm as a food source for the developing
embryo and seedling, serving a similar function to the yolk of animal eggs. The endosperm is also
called the albumen of the seed.

Embryos may be described by a number of terms including Linear (embryos have axile placentation
and are longer than broad), or rudimentary (embryos are basal in which the embryo is tiny in relation
to the endosperm)

OVULE DEVELOPMENT

a) The ovule consists of a mass of cells called the nucellus which is carried on a short stalk called
funicle. The nucellus is completely surrounded by two protective integuments except the narrow
channel at the tip called micropyle. One cell of the nucellus becomes larger and more conspicuous
than the rest. This is called embryo sac mother cell.
b) The embryo sac mother cell divides meiotically to give four haploid megaspore cells.
c) The three cells nearest the micropyle degenerate while the remaining one enlarges to form the
embryo sac.
d) The embryo sac nucleus divides by mitosis and the resultant nuclei migrate to the opposite poles.
e) Each nucleus undergoes two mitotic divisions to give a group of four haploid nuclei at each pole.

103
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

f) One nucleus from each polar group moves to the centre of the embryo sac. These are the polar
nuclei. The remaining nuclei develop cytoplasm around them and become separated by cell walls,
leaving two groups of three cells at each pole.
g) The three cells at the opposite end to the micropyle are called antipodal cells and play no further
role in the process. Of the three cells at the micropyle end,one,the egg remains, the other two,the
synergids, degenerate.

POLLINATION

 Pollination is the process by which pollen is transferred to the female reproductive organs of a
plant, thereby enabling fertilization to take place. Like all living organisms, seed plants have a
single major goal: to pass their genetic information on to the next generation. The reproductive
unit is the seed, and pollination is an essential step in the production of seeds in
all spermatophytes (seed plants).
 For the process of pollination to be successful, a pollen grain produced by the anther, the male part
of a flower, must be transferred to a stigma, the female part of the flower, of a plant of the same
species. The process is rather different in angiosperms (flowering plants) from what it is
in gymnosperms (other seed plants). In angiosperms, after the pollen grain has landed on the
stigma, it creates a pollen tube which grows down the style until it reaches the ovary. Sperm cells
from the pollen grain then move along the pollen tube, enter the egg cell through
the micropyle and fertilise it, resulting in the production of a seed.

 A successful angiosperm pollen grain (gametophyte) containing the male gametes is transported to
the stigma, where it germinates and its pollen tube grows down the style to the ovary. Its two
gametes travel down the tube to where the gametophyte(s) containing the female gametes are held
within the carpel. One nucleus fuses with the polar bodies to produce the endosperm tissues, and
the other with the ovule to produce the embryo. Hence the term: "double fertilization".
 In gymnosperms, the ovule is not contained in a carpel, but exposed on the surface of a dedicated
support organ, such as the scale of a cone, so that the penetration of carpel tissue is unnecessary.
Details of the process vary according to the division of gymnosperms in question. Two main
modes of fertilization are found in gymnosperms. Cycads and Ginkgo have motile sperm that
swim directly to the egg inside the ovule, whereas conifers and gnetophytes have sperm that are
unable to swim but are conveyed to the egg along a pollen tube.
The study of pollination brings together many disciplines, such as botany, horticulture, entomology,
and ecology.

104
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

It is important in horticulture and agriculture, because fruiting is dependent on fertilization: the result
of pollination. The study of pollination by insects is known as anthecology.

Pollination process

 Pollen germination has three stages; hydration, activation and pollen tube emergence. The pollen
grain is severely dehydrated so that its mass is reduced enabling it to be more easily transported
from flower to flower. Germination only takes place after rehydration, ensuring that premature
germination does not take place in the anther.
 Hydration allows the plasma membrane of the pollen grain to reform into its normal bilayer
organization providing an effective osmotic membrane. Activation involves the development
of actin filaments throughout the cytoplasm of the cell, which eventually become concentrated at
the point from which the pollen tube will emerge. Hydration and activation continue as the pollen
tube begins to grow.
 In conifers, the reproductive structures are borne on cones. The cones are either pollen cones
(male) or ovulate cones (female), but some species are monoecious and othersdioecious. A pollen
cone contains hundreds of microsporangia carried on (or borne on) reproductive structures called
sporophylls. Spore mother cells in the microsporangia divide by meiosis to form haploid
microspores that develop further by two mitotic divisions into immature male gametophytes
(pollen grains). The four resulting cells consist of a large tube cell that forms the pollen tube, a
generative cell that will produce two sperm by mitosis, and two prothallial cells that degenerate.
These cells comprise a very reduced microgametophyte, that is contained within the resistant wall
of the pollen grain.
 The pollen grains are dispersed by the wind to the female, ovulate cone that is made up of many
overlapping scales (sporophylls, and thus megasporophylls), each protecting two ovules, each of
which consists of a megasporangium (the nucellus) wrapped in two layers of tissue, the
integument and the cupule, that were derived from highly modified branches of ancestral
gymnosperms.
 When a pollen grain lands close enough to the tip of an ovule, it is drawn in through the micropyle
( a pore in the integuments covering the tip of the ovule) often by means of a drop of liquid known
as a pollination drop.
 The pollen enters a pollen chamber close to the nucellus, and there it may wait for a year before it
germinates and forms a pollen tube that grows through the wall of the megasporangium
(=nucellus) where fertilisation takes place. During this time, the megaspore mother cell divides by
meiosis to form four haploid cells, three of which degenerate. The surviving one develops as a
megaspore and divides repeatedly to form an immature female gametophyte (egg sac).
 Two or three archegonia containing an egg then develop inside the gametophyte. Meanwhile, in
the spring of the second year two sperm cells are produced by mitosis of the body cell of the male
gametophyte. The pollen tube elongates and pierces and grows through the megasporangium wall
and delivers the sperm cells to the female gametophyte inside. Fertilisation takes place when the
nucleus of one of the sperm cells enters the egg cell in the megagametophyte’s archegonium.
 In flowering plants, the anthers of the flower produce microspores by meiosis. These undergo
mitosis to form male gametophytes, each of which contains two haploid cells. Meanwhile, the
ovules produce megaspores by meiosis, further division of these form the female gametophytes,
which are very strongly reduced, each consisting only of a few cells, one of which is the egg.
When a pollen grain adheres to the stigma of a carpel it germinates, developing a pollen tube that
grows through the tissues of the style, entering the ovule through the micropyle. When the tube
reaches the egg sac, two sperm cells pass through it into the female gametophyte and fertilisation
takes place

105
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

TYPES OF POLLINATION
Depending on the source of pollen, pollination can be classified into 2 types - Self-pollination and
Cross Pollination (Xenogamy).
Self Pollination is further divided into Autogamy and Geitonogamy. Depending on agent of
Pollination, pollination can be classified into abiotic pollination and biotic pollination.
Self Pollination is the type of Pollination in which pollen grains are transferred from anther to the
stigma of the same flower (Autogamy) or pollen grains are transferred from anther to the stigma of
different flower of the same plant (Geitonogamy).
Cross Pollination or Xenogamy is the type of pollination in which pollen grains are transferred
from anther to the stigma of a different plant.

DOUBLE FERTILISATION

 This process involves the joining of a female gametophyte (megagametophyte, also called the
embryo sac) with two male gametes (sperm). It begins when a pollen grain adheres to the stigma
of the carpel, the female reproductive structure of a flower. The pollen grain then takes in moisture
and begins to germinate, forming a pollen tube that extends down toward the ovary through the
style. The tip of the pollen tube then enters the ovary and penetrates through the micropyle
opening in the ovule. The pollen tube proceeds to release the two sperm in the megagametophyte.
 One sperm fertilizes the egg cell and the other sperm combines with the two polar nuclei of the
large central cell of the megagametophyte. The haploid sperm and haploid egg combine to form a
diploid zygote, while the other sperm and the two haploid polar nuclei of the large central cell of
the megagametophyte form a triploid nucleus (some plants may form polyploid nuclei). The large
cell of the gametophyte will then develop into the endosperm, a nutrient-rich tissue which provides
nourishment to the developing embryo. The ovary, surrounding the ovules, develops into the fruit,
which protects the seeds and may function to disperse them.
The two central cell maternal nuclei (polar nuclei) that contribute to the endosperm, arise by mitosis
from the same single meiotic product that gave rise to the egg. The maternal contribution to the
genetic constitution of the triploid endosperm is double that of the embryo.
 The generative cells divides by mitosis to produce two male gametes.
Double fertilization gives rise to the zygote and endosperm
 Directed by a chemical attractant, possibly calcium, the tip of the pollen tube enters the ovary,
probes through the micropyle (a gap in the integuments of the ovule) and discharges two sperms
within the embryo sac.
 Both sperm fuse with nuclei in the embryo [Link] sperm fertilizes the egg to form a zygote.
 The other sperm combines with the two polar nuclei to form a triploid nucleus in the central cell.
 This large cell will give rise to the endosperm, a food-storing tissue of the seed.
 The union of two sperm cells with different nuclei of the embryo sac is termed double
fertilization.
 Double fertilization is also present in a few gymosperms, probably via independent evolution.
 Double fertilization ensures that the endosperm will develop only in ovules where the egg has
 been fertilized.

106
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

EVENTS AFTER FERTILISATION

After fertilisation, the zygote divides rapidly by mitosis and develops into the embryo, which then
differentiates into a young shoot called plumule, a young root called radical and seed leaves called
cotyledons. The primary endosperm nucleus also divides mitotically to give a mass of cells, the
endosperm. This forms the food source for the growing embryo. In legumes like beans or peas, it is
quickily absorbed by and stored in cotyledons. The most common food stored is carbohydrates. This is
usually in the form of starch but some seeds store quantities of sugar. After fertilization, the following
changes are observed in a flower:
 There is formation of a diploid zygote and it develops into an embryo, which forms the future
plant.
 The endosperm cells serve as a source of nutrition for the developing embryo.
 The ovule becomes the seed.
 The ovary becomes the fruit.
 In most of the plants the antipodals and synnergids disintegrate before, during or immediately
after fertilization.
 The outer and inner integuments of the ovule become the testa or the seed coat of the seed.
 Petals and sepals fall off.

CHAPTER 14

SEEDS

 describe the internal and external parts of a seed

 describe functions of seed parts
 discuss requirements for seed germination.
 describe the processes of seed germination
 distinguish epigeal and hypogeal germination

The Seed
The seed is a miniature plant in an arrested state of development. This embryonic plant can under
favourable conditions develop, at first drawing raw materials from a food store in the seed, into an
actively metabolizing individual individual that can synthesise its food from simple inorganic
compounds. The seed is not only a source of food for the embryonic plant but is being used to feed

107
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

man and his animals; the seeds of legumes and cereals have proved to be very important in this
respect.
Seed Structure

Figure 8.19 Diagrams showing structure of different seeds

A mature seed consists of the following parts.

Testa:
This is the seed coat and develops from the integuments of the ovule. The integuments may develop
either wholly into the testa or part of them may be absorbed by the other developing structures. The
main function of the testa seems to be that of protecting to varying degrees, the embryo from fungi,
bacteria and pests. The testa of some plants is very impermeable to moisture and this is an important
factor in the delayed germination of these ‘hard’ seeds, e.g. leguminosae.
Endosperm
The endosperm is a product of the fusion of the two polar nuclei and one male gamete. The longevity
of the endosperm and the amount and kind of stored material varies with species. Therefore the seeds
of some plants have no endosperm and are called exalbuminous seeds while those with endosperm are
albuminous. In eexalbuminous seeds food materials are stored in the cotyledons, e.g. Leguminosae,
compositae, cucurbitaceae. Seeds of graminae is usually found starch and proteins. The proteins are
stored either as glutens or as aleurone grains. The aleurone grains are restricted to the outermost layer
of endosperm cells called the aleurone layer. The rest of the endosperm stores starch or fats.
Embryo
The mature embryo of the dicots consists of the embryo axis and the two first foliar structures, the
cotylendons. Below the cotyledons the axis is called the hypocotyls and at its lower end it bears an
embryonic root called the radical. At the top end of the axis is the apical meristem of the future shoot
and sometimes a small shoot develops before the embryo is mature and this bud is the plumule or
epicotyl.
The embryo in the monocots consists of the same parts as that of the dicots but because it lacks one
cotyledon, the embryo is not two-lobed at the distal end. In the caryopsis of the grasses it will be
observed that the embryo is adpressed to the endorsperm by a cotyledon, the scutellum. The
embryonic root and its rootcap are enclosed in a coleorhiza while the shootapical meristem is enclosed
in coleoptiles.
Hilum
This is the scar lefty by the stalk that attached the seed to the ovary.
Micropyle
The micropyle has been identified earlier and is a tiny pore in the testa, usually opposite the radical.
Seed Germination

108
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Germination is the resumption of active growth on the part of the embryo resulting in the rupture of
the testa and the emergence of the young plant. The seed of some plants will germinate soon as they
are ripe and the conditions favourable. e.g. some varieties of groundnuts sprout in the field following
showers of rain late in the season. On the other hand the seeds of many of many plants will not
germinate until after a certain period of time has elapsed – these are dormant.
The process of seed germination

Figure 9.1 Seed Germination and Seedling growth of Allium cepa

Seed Germination is the sum of events beginning with hydration and culminating with root
emergence. There are several stages involved in germination. These include 1) imbibition of water, 2)
activation of enzyme systems, 3) metabolism of storage products and their transport and 5) the
emergence of the radical and growth of the seedling.
Stage 1‐ Imbibition of water: There are two events occurring in water imbibition and they include the
water uptake and increase in respiration. The seed is very dry and has a strong propensity for water.
Remember, water flows from a region of high concentration to one of low concentration or from wet
to dry.
Water absorption has three phases.
In phase I, there is rapid water absorption independent of metabolic activity but dependent on soil
texture, degree of packing, proximity of seed to soil and seed to soil contact.
Phase II shows little water absorption since the seed is saturated. However, there is an increase in
enzyme activation and synthesis that will be used in Phase III.
Phase III is associated with another water uptake where radical emergence and elongation occurs.
This results in cell elongation without cell division. In this stage enzymes activated in Phase II break
down storage materials (fats, proteins,carbohydrates and phosphorous containing compounds) and
transfer them to growing [Link] and are usually transformed into carbohydrates for use.
Proteins are hydrolysed by various proteases into amino acids which are not consumed in respiration
but are utilized in the synthesis of plant organic compounds.

109
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Stage 2 – Increased Respiration: Respiration follows four phases that tracks closely to water
absorption. Respiration involves the uptake of oxygen and the conversion to carbon dioxide and in the
process the generation of energy.
Phase I of respiration involves the activation and hydration of existing enzymes.
Phase II involves a plateau in oxygen uptake caused by the seed coat restricting gas exchange. Phase
III shows an increase in oxygen resulting from the formation of new enzymes in the dividing cells that
leads to the resurgence of respiration.
Phase IV shows a decline in respiration as the seedling emerges and begins to photosynthesize.
The main purpose of the stored materials is to provide a respiratory substrate as a source of energy and
also to provide new materials for growth (e.g. new protein, complex lipids and cellulose). Since the
site of use (sink) of these materials is the developing embryo then the dry weight of the plumule and
radical increases with germination while that of storage organ (source) decreases. However, the total
dry weight of the seedling decreases for time due to respiratory consumption and latter increases with
absorption of materials from the environment and commencement of photosynthesis.

Stage 2 ‐ Activation of enzyme systems: There is great evidence show‐casing the activation of newly
formed enzyme systems that is evidenced by the following;
1) Increased enzyme activity prior to germination.
2) Use of protein synthesis inhibitors which inhibits germination.
3) Incorporation of radioactive precursors into proteins to track the enzyme systems.

110
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

4) Use of immunological and molecular techniques.

Stage 3‐Formation of enzyme systems and metabolism of stored compounds: Metabolism or
stored products and their transport can occur via the activation of pre‐existing enzymes or their
pre‐cursors, activation of pre‐existing but inactive enzymes and the synthesis of new enzymes from
pre‐existing or newly produced mRNA. There are many enzymes involved in this process that include;
lipases (break down lipids to fatty acids and glycerol that produce membranes and
energy), proteinases (break down proteins to smaller compounds) and phosphatases (responsible for
nucleic acid and protein synthesis).
Stage 4‐Radical emergence and seedling growth: This is associated with the second water uptake
leading to radical emergence and seedling autotropism (self dependent). Here cell division (increase of
cell number) in the root and shoot and cell expansion begins. Also the seedling is divided into a
hypocotyl (stem region below the cotyledons) and epicotyls (stem region above the cotyledons) and
respiration drops significantly since the seedling begins responding to sunlight.
Hormones in seed germination
During the embryo development, endogenous plant hormones (Gibberellins (GA), Auxins (IAA) and
Cytokinins (CK)) increase in the embryo while the developing embryo is heterotrophic (dependant on
the mother tissue for support). In later stages of embryo development, Abscisic (Acid ABA) levels
increase that will maintain seed dormancy. There is a great interaction between GA, CK and ABA
during seed germination. Seed germination cannot occur without the presence and activation of GA.
However, if ABA is present at high levels, germination will not occur since ABA blocks GA. But if
CK is in an active form, it suppresses inhibition of ABA on GA and germination can occur.

TYPES OF GERMINATION

Type # 1. Hypogeal Germination:

In this kind of germination, the cotyledons do not come out of the soil surface. In such seeds the
epicotyl (i.e., part of embryonic axis between plumule and cotyledons) elongates pushing the plumule
out of the soil. All monocotyledons show hypogeal germination (Fig. 4.3, 4.4, 4.5). Among
dicotyledons, gram, pea (Fig. 4.2), groundnut are some common examples of hypogeal germination.

In monocotyledons (e.g., wheat, maize, rice, coconut) radicle and plumule come out by piercing the
coleorrhiza and coleoptile respectively. The plumule grows upward and the first leaf comes out of the
coleoptile. The radicle forms the primary root which is soon replaced by many fibrous roots.
(i) Germination of Pea Seed:
111
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The seed imbibes water and swells. The radicle comes out and first penetrates the soil and forms root
system by giving out secondary branches. It is the epicotyls which grows first. It arches out and carries
the plumule above ground. The plumule soon forms the aerial shoot. The cotyledons remain under the
soil throughout (Fig. 4.2).

(ii) Germination of Maize Grain:

The grain imbibes water from moist soil. The coleorhiza pierces the base of caryopsis (fruit) and
appears as a shining knob. After sometimes, the coleorhiza gets ruptured due to growth of radicle.
After sometime coleoptile comes out.
Three seminal roots develop from above the radicle (but variation in number). The radicle and seminal
roots with two branches persist throughout the life of the plant. Adventitious roots are formed from the
lowermost nodes above the mesocotyl (Fig. 4.3).

(iii) Germination of Wheat Grain:

Details of wheat grain germination (Fig. 4.4) are similar to those of maize grain germination. Wheat
grain shows hypogeal germination.

112
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

(iv) Germination of Coconut (Cocos nucifera):

During germination the lower end of the embryo forms the cotyledon which begins to grow as a
spongy structure inside the endosperm. This spongy cotyledon increases in size as it absorbs food
material stored in the endosperm. The upper end of the embryo develops through the ‘eye’ carrying
the radicle and the plumule.
The plumule pierces the fibrous pericarp and emerges like a horn. This develops the aerial shoot even
before the roots have come in contact with the soil. The radicle fails to develop any further but several
adventitious roots grow from the base of the plumule. The seedling becomes established where the
adventitious roots penetrate the soil (Fig. 4.5).

Type # 2. Epigeal Germination:

In seeds with epigeal germination, the cotyledons are brought above the soil due to elongation of the
hypocotyl. In castor, cotton, papay, onion (Figs. 4.7., 4.8), flat green leaf like cotyledons can be seen
in the young seedlings. Here the cotyledons, besides food storage, also perform photosynthesis till the
seedling becomes independent. In some other plants like bean, the cotyledons being thick, do not
become leaf-like; they shrival and fall off after their food reserves are consumed by the seedling.
(i) Germination of Gourd (Cucurbita maxima):
The straight radicle comes out of the seed and fixes the seed to the soil with the secondary roots
developing from the radicle. Next, the hypocotyl grows so quickly that it forms a loop which comes
out of the soil and pulls out the rest of the seed. The seed coat is cast off and the cotyledons open out
113
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

like two leaves, become green, large and thin so that they look and behave like ordinary leaves. The
plumule within the cotyledons becomes exposed and soon grows into the aerial shoot (Fig. 4.6).

(ii) Germination of Castor (Ricinus communis):

The seed imbibes water and the testa bursts near the caruncle and the radicle grows out. After this
hypocotyl grows due to which two papery cotyledons enclosed by endosperm are pulled out of the
soil. Cotyledons come out of the endosperm when it is consumed. The cotyledons become green and
leaf-like, while the plumule slowly develops into leafy shoot. The remanents of endosperm withers
and drop off (Fig. 4.7).

(iii) Germination of Onion Seed:

In this case the radicle as well as the base of the scutellum (cotyledon) grow out of the seed. The
radicle penetrate the soil, while the other end of the cotyledon remains within the endosperm and
sucks the food material. The base of the cotyledon grows further, turns green and pushes the seed out
of the soil. The plumule is not visible so long as it is covered by the base of the cotyledon in the form
of a sheath just above the radicle.
The plumule now pierces the cotyledonary sheath and forms the first cylindrical foliage leaf.
Meanwhile adventitious roots develop from above the radicle and form a fibrous root system (Fig. 4.8)
(In this case the seed is pushed out of the soil by growth at the base to cotyledon and not by growth of
hypocotyl).

114
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Type # 3. Vivipary (Viviparous Germination):

Vivipary is the phenomenon of giving birth to young ones in advanced stage of development. It occurs
in mammals (among animals) and mangrove plants. In mangrove plants (e.g., Rhizophora, Sonneratia,
Heritiera) the seeds cannot germinate on the ground because of the excessive salt content and lack of
oxygen in marshy habitat. In such plants seed dormancy is absent.
The embryo of the seed (present inside the fruit) continues growth while the latter is attached to the
parent plant. Hypocotyl elongates and pushes the radicle out of the seed and the fruit. Growth
continues till the hypocotyl and radicle become several centimetres long (more than 70 cm in
Rhizophora). The seedling becomes heavy.
As a result it breaks its connection with the fruit and falls down in the salt rich muddy water in such a
position that the plumule remains outside the saltish water while the tip of the radicle gets fixed in the
mud. This protects the plumule. The radicle quickly forms new roots and establishes the seeding as a

new plant (Fig. 4.9).

.
Precocious germination

When a seed germinates without undergoing all four stages of seed development, i.e., globular, heart
shape, torpedo shape, and cotyledonary stage, it is known as precocious germination.

115
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Factors affecting germination

Water
Water enters through the micropyle and testa, the relative importance of these paths vary according to
the permeability of the testa. It is the initial imbibition of water that triggers germination. The amount
of moisture in the soil need not be very high. The consequences of water absorption are as follows;
 The testa usually ruptures or its permeability to gases is increased after being moistened. This
will allow gaseous exchange for aerobic respiration to occur at fairly rapid rate.
 Water activates enzymes and provides a medium in which metabolic activities and translocation
can occur.
 Vacuolation can take place so that the embryo can increase its length.
Oxygen
In the dormant seed respiration occurs at a very slow rate and is of the anaerobic kind. Carbon dioxide
(CO2) accumulates in the seed because the testa is dry and impermeable, this gas will inhibit
germination. When the permeability of the testa the gases increase, there is a marked rise in the
respiration rate with the respiration changing from anaerobic to aerobic. The seeds of some species
can germinate under conditions of very low oxygen content. On the other hand the seeds of some
weeds will be buried deep in the soil without germinating. When the land is disturbed by ploughing,
the seeds are brought nearer the surface and will germinate if moisture present. The failure of these
seeds to germinate is because of either low O2 tensions or high CO2 tensions.
Temperature
Dry seeds will remain viable even if exposed to temperatures that are very extreme. Once the seed
imbibes water then the temperature range over which the seed remains viable becomes narrow. The
seed of a specific species will germinate only in a certain temperature range; the temperature below
which no germination occurs is the minimum, the temperature above which no germination occurs is
called the maximum, the temperature that favours maximal germination is the optimum. By maximal
germination is the attainment of the highest germination percentage in as short a period of time as
possible.
The reason for the existence of a maximum temperature is the denaturation of proteins (including
enzymes). Temperature also affects the rate of chemical reactions and water uptake by altering the
viscosity of the water and the kinetic energy of the molecules.

An exposure of seed to low temperatures in the presences of moisture before germination increases the
germination percentage – this is known as stratification. The process decreases the period of after –
ripening in dormant seeds.
Light
Most seeds will germinate in both light and darkness. The seeds of some species will germinate only
in the presence of light (e.g. lettuce, tobacco). In these light sensitive seeds infrared light inhibits
germination and red light promotes germinates. The effect of these wavelengths is connected with a
light sensitive pigment called phytochrome.

Test seed for viability and determine germination percentages.

There are two types of these tests. First type, when germination is done under favourable conditions
(standard laboratory germination, and test of growth intensity). Second type, when seed is exposed to
unfavourable environmental conditions (cold test, accelerated aging test, and Hiltner test);
Biochemical tests – are considered as indirect methods for estimation of seed value. These are
Tetrazolijum test, conductometric measurements, enzyme activity and respiration.
Seed viability

116
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The viability of seed is usually tested by germinating the seed under optimum conditions and
determining the percentage of the seed that produces normal seedlings.
Tetrazolium test
Tetrazolium test is based on reduction of colourless solution 2,3,5 – tripheniltetrazolim chloride or
bromide into insoluble 2,3,5 – triphenilformazan red in colour. This solution acts as an indicator for
detection of reduction processes that take place in living parts of the seed. Inside the seed, tetrazolim
intakes hydrogen from dehydrogenase. By hidrogenization of tetrazolium a red, stable substance
called formazan, which dyes living parts of the seed, is formed in the living cells (ISTA, 2009).
Seed is submerged in water because swollen seed is hard to crack and easy to cut in relation to dry
seed, and dying is more uniform. Tissue of many plant species must be removed to introduce the dye
into the tissue. Tissue removal can be done by pilling the seed coat off, punching, and longitudinal or
cross-cutting of unessential seed parts.
Prepared seed is submerged into 0,5 – 1% tetrazolium solution. Seed must be completely covered with
solution, and not exposed to direct light. After the time needed for dyeing expires (it depends on plant
species) the estimation of dyeing is approached. During testing the viable seed should express its
potential for normal seedling formation through biochemical activity. Non viable seed expresses
malformations which prevent normal seedling formation (ISTA, 2009).
All tissue (necessary for normal seedling development) of a viable seed should be dyed. Except
completely dyed, viable seeds, and completely undyed, unviable seeds, a partly dyed seeds may also
be found. Depending on the species, small undyed spots of some parts of these tissues may be
accepted. Location, size of undyed areas, and sometimes intensity of dyeing, determine whether some
seed is considered as viable or not.
Tetrazolium test has several limitations because it:
- provides too high values of vitality, i.e. within the vigour seeds it cannot be separated seed which
will give typical and abnormal seedlings,
- causes difficulties in the visual identification of abnormal seedling (i.e. split coleoptiles, negative
geotropism etc.),

CHAPTER 15

DORMANCY

Seed dormancy
Non – germination of seeds due to absence of suitable conditions is termed as dormancy.
OR
A physical or physiological condition of viable seed, which prevents germination even in the presence
of favorable conditions

Viable seed of some species does not germinate even if the external conditions are optimal; such seed
is said to be dormant. The dormancy of seed is due to one or more factors.
Types of seed dormancy
In general, there are two types of seed dormancy: seed coat dormancy and internal dormancy. Seeds
with seed coat dormancy usually have a seed coat that is impermeable to oxygen and/or water.
Occasionally the dormancy is caused by an inhibiting chemical in the epidermis or adjacent interior
membranes. Internal dormancy is a general term encompassing a number of
physiological conditions that delay germination. Not all of these conditions are
fully understood or easy to counteract. The most common one is called afterripening.

117
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Seeds that require an after-ripening period, even though harvested when mature, germinate poorly or
not at all until they have been subjected to moisture and either high or low temperatures or both in
sequence
a) Physical dormancy
This type of dormancy is due to structural limitations to germination. The testa may be impermeable
to water and gases as is the case in some legumes. Permeability improves slowly under natural
condition whereby soil micro organisms and fluctuations of moisture and temperature weakens the
testa.
The seed can also be scarified artificially by mechanical abrasion or by the corrosive action of
sulphuric acid. The testa may offer physical resistance to the expansion of the embryo even if water
and oxygen are absorbed. The seed of Amaranthus will remain dormant if saturated with moisture and
on drying certain changes occur in the testa so that on rewetting the seed can germinate.
b) Physiological dormancy
In some plants, the fruit and seed might be ripe when the embryo is still immature. The embryo should
therefore develop further. In other cases the embryo might be mature but dormant. (e.g. apple, apple)
and the embryo requires a period of after – ripening. This period can be shortened by stratification as
has been mentioned. Dormancy can also be controlled by growth regulators (plant hormones).
Inhibitors, such as coumarin, will prevent germination and may be present in the flesh of the fruit (e.g.
tomato) or the seed itself (e.g. irish). The inhibitors can be removed by leaching. Some compounds
have an effect opposite to that of the inhibitors and are called promoters (e.g. gibberellins). The
concentration of a promoter should reach a certain level before the seed can germinate.
c) Secondary dormancy
Some seeds are capable of germinating soon after they are ripe but if exposed to conditions
unfavorable for germination they become dormant for certain period of time. Seeds of white mustard
fail to germinate after exposure to a high concentration of CO2 even if the CO2 is later withdrawn.
Light sensitive seed may go dormant after exposure to darkness. Secondary may include physical
and/or physiological dormancy.

BREAKING SEED DORMANCY

A general summary of methods for breaking seed dormancy,

Methods of Breaking Seed Dormancy
i) Microorganisms present in the soil weaken and decompose the hard seed coat
ii) The digestive juices present in the alimentary canal of the fruit eating birds makes the seed coat soft
iii) Mechanical abrasions weaken the tough and impermeable seed coat
iv) Washing away of inhibitors by rain or irrigated water
v) Inactivation of growth inhibitors by heat and cold
vi) Leaching of solutes in halophytes where dormancy is due to high concentration of salts
vii) Synthesis of growth hormones
viii) Maturation of embryo
Scarification[for some members of legume family]
Mechanical scarification is a technique for overcoming the effect of an impermeable seed coat.
Mechanical scarification can be done by rubbing seeds between two pieces of sandpaper (Schmidt
1980), or using a file, a pin, or a knife to rupture the seed coat. Seed may also be mixed with coarse
sand and shaken vigorously in a jar (Schmidt 1980). Even a vise can be used to squeeze seeds along
the suture until they crack open. Care must be taken not to injure the embryo. It may be necessary to
open a couple of seeds to see where the embryo is located in relation to the micropyle, the former
point of attachment to the fruit. Large seeds like those of the bush lupine (Lupinus) are easily scarified
with a knife; the hot water treatment is easier for small seeds (Emery 1987).
Low Temperature (Chilling)

118
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

In many woody and in certain herbaceous species the dormancy can be broken by chilling treatments,
just above freezing (0 - 5o C).
Alternating Temperatures
Seeds are exposed to alternating high and low temperatures to reduce the concentration of the
inhibitory substances.
Running Water Treatment
Running water washes off the inhibitors from the seeds and breaks dormancy.
Light
In some plants, dormancy can be broken by exposing them to light.
Hormones Treatment
Dormancy in some seeds is broken by treating them with hormones such as gibberellins, cytokinins
and ethylene.
Gibberellic acid
Germination test paper is moistened with a 0.05% solution of Gibberellic acid (GA3), prepared by
dissolving 500 mg of GA3 in 1 l water. Germination is then continued in recommended conditions.
Potassium nitrate
A 0.2% solution of potassium nitrate (KNO3) – prepared by dissolving 2 g KNO3 in 1 l water – is
used to moisten the germination paper at the beginning of the test. Germination is continued in
recommended conditions.

CHAPTER 16

PLANT – WATER RELATIONS

Plants can be classified as hydrophytes – plants that grow in water and usually have a poorly
developed root system; mesophytes – intermediate in water requirements and have a well developed
root systems; and xerophytes – plants which have been adapted to dry conditions.
Importance of water
Water is a constituent of protoplasm and acts as a solvent for numerous solutes and supports the huge
molecules of proteins and nucleic acids. The hydration properties of water permits ready reaction
between molecules in solution and between enzymes and substrates. Water participates as a chemical
itself in a number of reactions such as photosynthesis and hydrolysis and is a product of respiration.
Water also maintains the turgidity of cells and therefore the shape of the plant. Water also acts as a
coolant in plants.

PROPERTIES OF WATER
The molecule of water consists of two hydrogen and one oxygen atom. The two hydrogen atoms are
joined to the oxygen atom by shearing of electrons (covalent bond). Note that although the water
molecule is neutral, the oxygen atom retains a slight negative charge and the hydrogen atoms retain a
slight positive charge. Such molecules are termed polar molecules or dipoles.
The charges on the water molecule ensure the following:-
hydrogen bonds in the process of cohesion.
adhesion.

nuous water columns are required

for the transport of water, mineral and organic nutrients in the plant.
high surface tension. This implies that it is not easy for the surface of water to be
broken or deformed. This is very important for the structural stability of the plant.

119
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Note that the structure of plant parts and organs are maintained because of the forces created by water
in the cells of the plant (turgor).
Other useful properties of water
solvent for polar substances. These include substances like salts, sugars and
simple alcohols.
transport medium as in the xylem and phloem.
medium in which all reactions in the plant take place.
ter requires a lot of energy to increase its temperature (high specific heat). Therefore,
biochemical processes which take place in water can operate over a small temperature range.
high heat of vaporization).
This helps to maintain the temperature of plants at levels where they are not biologically damaged by
sudden increase in temperature.
high heat of fusion). Contents of cells are therefore less
likely to freeze. Ice crystals can be dangerous if they develop inside cells.
density (mass per unit volume) of water decreases below 4ºC and therefore ice tends to float.
Water is the only substance whose solid form is less dense than its liquid form.
Organisms that live towards the bottom of fresh water lakes are therefore protected from freezing.

Water Potential (ᴪ) Concepts

In order for you to understand how water moves in the soil and in the plant, you need to understand a
concept called ‘water potential’. Water potential quantifies the tendency of water to move from one
area to another due to osmosis, gravity, and mechanical pressure and matrix effects. Water potential
is a measure of the energy status of water. Pure water has an energy status of zero. It is the reference
point. A technical definition of water potential makes reference to cell membranes, salts that exist in
plants and creating certain pressure as you will shortly.
‘Water potential is the difference in free energy per unit volume between matrically bound,
pressurized and osmotically constrained water and pure water at the same temperature’.
Remember that that pure water has a water potential of zero. Addition of solutes like salt or sugar to
water reduces its water potential. Since we started our water potential at zero, reducing the water
potential means it attains a negative value. It makes the water potential negative. In other words the
water molecules in this solution are not likely to move out of a membrane system.
The water potential of a plant system is made up of the following components:-
a) Matric potential
b) Osmotic potential
c) Pressure potential/turgor pressure
Below is an illustration of how water potential is calculated:-
ᴪ = ᴪ (Osmotic) + ᴪ(Matric) + ᴪ(Pressure)
(Water potential) (Osmotic potential) (Matric potential) (Pressure potential)
(Negative) (Negative) (Positive)
Now let us explain the three components of water potential listed above.
a) The osmotic potential
This comes about as an effect of solutes. If solutes are added to water, the free energy of water is
reduced as a result of interactions between the solutes and the water molecules. The concentration of
solutes and the type of solutes dilute with the water molecules reducing the water activity. The
importance of osmotic potential is that both the inorganic (potassium, sodium and chlorine) and
organic (sucrose, glycerol and glycine) solutes affect the osmotic potential of plant cells. Inorganic
solutes occur predominantly in the vacuoles while occur in the cytoplasm. These components play an
important role in water regulation of plants. This is defined as ionic adjustment of the cell. This
condition makes it possible for plants to respond to changes in surrounding water [Link]

120
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

amounts of solutes dissolved in water constrain water movement and therefore it’s designated as
negative.
b) The matric potential
This describes the availability of charged surfaces in the cell walls and in the cell system. There is
interaction with water molecules resulting in the reduction of water’s ability to freely move hence free
energy or water potential. This water potential represents suction of water. Water can be held in cell
wall and cell membranes and this is an adsorptive effect. Water can be held in capillaries by surface
tension effect. All these forces reduce the mobility and activity of water within the plant thereby
lowering the water potential. The amounts of matric forces or charged surfaces available bind water
and reduce its movement and therefore it is designated as negative.
c) The pressure potential
This describes pressure exerted on water by the cell wall which forces water to move. Pressure
potential in plant cells is a result of cell walls which push against cell contents especially when the cell
is turgid. The pressure potential of pure water is zero. Thus any increase in pressure increases the
tendency of water to move (out a permeable membrane). Hence it is denoted by a positive value. The
importance of pressure potential is turgidity. It increases turgor pressure in the cell. This exerts
pressure on the cell wall and the protoplasm and the cell volume is increased by 10-30%. It is
important for rigidity; shape and straightness in young plants which have not lignified i.e. cellulose,
hemi-cellulose and lignin have not yet developed. When water surrounding the cell is lower than that
of the plant cell, water moves out of the cell. There will be shrinkage of the cell volume, the
concentration of the cytoplasm increases and then there is separation of the cell wall. This condition is
called plasmolysis. In crop plants it is observed as wilting.
Now let us look at the uptake of water of by plant, but first note the following:-
t contact with water in the soil and with water vapour in the air around
the plant.
(-3, less negative) in the
soil to a region of lower water potential in the atmosphere, down to a gradient of water potentials.

xylem, moves up in the xylem to the leaves and is lost by evaporation from the surface of the leaf
(mesophyll) cells before diffusing out through the stomata.

transpiration stream.

Water movement
Diffusion is the net movement of molecules of molecules from regions of high concentration and this
is a function of their kinetic energy. The tendency of one species of molecules to diffuse (diffusion
pressure, D.P.) is decreased by the presence of another species of molecules, e.g. solutes in water. The
rate of diffusion will depend on the steepness of the concentration gradient (or free energy gradient)
and the permeability of the medium such as membranes to water.
Osmosis is a special kind of diffusion. For instance when an aqueous solution is separated from pure
water by differentially – permeable membrane (a membrane permeable to solvent (e.g. water
molecules and not solute molecules) the net movement of water will be from the solvent into the
solution. Osmosis can be defined as the diffusion of a solvent across a differentially – permeable
membrane. The diffusion pressure of water in a solution is less than that of pure water and the
difference between the two is the diffusion pressure deficit (DPD) or the water potential difference.
The osmotic pressure (OP) of a solution tells us the maximum possible pressure that could be
developed in the solution if it were permitted to come to equilibrium with pure water. The pressure
could be larger in a concentrated solution than in a dilute one.

121
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The actual pressure that develops as a result of osmosis is turgor pressure (TP) and ranges usually
from zero to OP. A negative pressure is exerted by the wall as a result of turgor pressure and is known
as wall pressure (WP).
The parts of the cell that are involved in these relationships are;
 The cell wall – this is completely permeable and has no direct role in osmosis;
 The plasmalemma and tonoplast – These membranes are partially differentially permeable
(solutes can pass through them slowly).
 The vacuole – this contains an aqueous solution called the sap.
If the diffusion pressure deficit of the vacuole is higher than the diffusion pressure deficit of the
surrounding solution then water is taken up by the cell (endosmosis). After equilibrium has been
attained the diffusion pressure deficit will equal. The medium is pure water, then at equilibrium
diffusion pressure deficit equal zero and since DPD = OP = TP, then OP = TP. In such a case the cell
is said to be fully turgid.
Absorption of water
The absorption of water from the soil is through the zone of the root that bears root hairs. Some water
may be absorbed by the zone of cell elongation. Water from the soil passes through the epidermal cells
of the absorbing regions and across the cortex, and then through the endodermis. The route of water
movement through the cortex is the cell wall, and cytoplasm to the plasmodesmata and then into the
cytoplasm of the next cell, and so on. At the endodermis the free passage of water through the cell
wall and cytoplasm is blocked by the presence of casparian strips on the transverse and radial walls of
the endodermis.
The bulk of the water has to move through the vacuole. In some plants thin-walled endodermal cells
are present opposite the xylem tissues and facilitate water movement – these are called passage cells.
Due to the evaporation of water from mesophyll cells a diffusion pressure deficit develops in these
cells. This inturn will cause the development of a diffuision pressure deficit in the xylem tissues and a
tension is built up down the xylem and leads a creation of diffusion diffusion pressure across the root.
When the diffusion pressure water in the root hairs exceeds that of the soil-water, then moves from the
soil into the root. This type of absorption is said to be “passive” and results from the ‘suction’ of the
water upwards.
Another possible method of driving the water would be the pumping action of the root. It has been
demonstrated that water is exuded from the stub of a fresh cut stem. This exudation is due to some
forces which are not well understood and referred to as root pressure.
Factors affecting water absorption
Available soil water
Plants can only absorb soil moisture that is free. When the moisture is held tightly by the soil particles
then absorption is very slow. Such a condition is found when the moisture in the soil is held at below
the permanent wilting point. Absorption is also suppressed when the moisture is in excess of the field
capacity because of reduced aeration.
Aeration of the soil
In most cases the absorption of water by the roots is faster in a well-aerated soil than in a badly-
aerated one. The effects of aeration are obviously felt by the process of aerobic metabolism of the
roots. Poor aeration is usually due to excess soil moisture or soil compaction. The roots of some
species, such as the hydrophytes, absorb water even in anaerobic conditions.
Soil temperature
Very low soil temperatures slow down the rate of absorption. This is probably because root elongation
is retarded, movement of water molecules is slowed, permeability of cell membranes decreased and
metabolic activity of the roots decreased. High soil temperatures are not common but if they exist,
water absorption also decreases.
Concentration of soil solution

122
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

This factor affects the osmotic potential of the soil solution and absorption usually decreases with an
increase in soil solution concentration.

Loss of water to the atmosphere

Plants take in large quantities of water than they require for the life processes. Most of the water is lost
to the atmosphere in the form of vapour – this loss of water by evaporation is known as transpiration.
Sites of water loss
Any cell surface that is not protected by a water-proof covering and in contact with a less saturated
atmosphere loses water by evaporation. Therefore, in order to prevent excessive loss of water, plants
adapted to live on land have developed external layers of cutin and suberin. However, the layers are
perforated to allow for aeration and it is through these perforations that water vapour diffuses into the
atmosphere.
There are three main places where some loss of water can occur;
 The cuticle can lose small amounts of water. The amount evaporated will depend on the thickness
of the cuticle. This is called cuticular transpiration.
 Water loss also takes place through the lenticels of fruits and stems. Water loss by lanticular
transpiration is negligible.
 80 – 90 % of all the water evaporation from the plant passes through the stomata of the leaves and
some stems. Loss of water involves evaporation at the mesophyll cell walls and the subsequent
diffusion of the vapour through the intercellular spaces and out of the stomatal pore into the
atmosphere. The size of the stoma varies with species but is invariably small. Stomata may be
found on both surfaces of the leaf (e.g. wheat, maize, Lucerne), or on the lower surface as in most
woody species (e.g. floating leaves of water lilly).

CHAPTER 17

WATER PATHWAYS
a) Symplast pathway
 Water moves into the cytoplasm of a cortical cell and then to adjacent cells through the
interconnecting plasmodesmata. The symplast pathway is considered the ‘living route’ because
water has to pass through the interior of the cell and get transmitted from cell to cell.
b) Vacuolar pathway
 Water passes from cell vacuole to cell vacuole through neighbouring cells, crossing the symplast
and apoplast and moving through membranes and tonoplasts (vacuolar membrane) (remember the
plant cell structure) by osmosis. Relatively little water moves through this pathway.
c) Apoplast pathway
 Water moves from one cortical cell to another through spaces in the cell wall, without entering the
cytoplasm of the cell. This is movement of water through the non-living continuum of the
epidermal and cortex cell. Note that the apoplast is the major pathway for water movement into the
xylem. This is because the symplast route offers resistance to water movement.

123
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Stomatal mechanism
The most important functional property of the stomata is that they are sometimes open and sometimes
closed. When open then gaseous diffusion can take place and the principal gases that pass through O2
and CO2 and water vapour. The closure of the stomata greatly retards the diffusion of these gases

The closure and of the opening is due of the stomata the changes in the turgidity of the guard cells and
that these changes are due to endomosis or exomosis by the guard cells. It is also a general observation
that the stomata have a diurnal rhythm in that they are open during the day and close at night. The
effect of the changes of turgidity results in changes in the volume of the guard cells. It is this change
in volume and the uneven thickness of the walls of the guard cells that result in the stomata being open
or closed. The stomata opens when the guard cells gain water (high OP) and closes when they lose
water (low OP). The behavior of the stomata was first thought that since the guard cells are the
only epidermal cells that have chlorophyll then the concentration of sugars in their vacuoles rose to
high enough level to create DPD large enough to cause endosmosis. However this placed too great a
demand on the photosynthesis of the guard cells. Later it was discovered that the starch in the guard
cells turned into sugar in light whereas that of other cells was converted in the dark. i.e.
Guard cells Starch to sugar (light)
Parenchyma cells Starch to sugar (dark)
To explain this it was thought that the diastase of guard cells required a different optimum pH from
that of the enzyme in other parts of the leaf. Therefore the guard cells has to change any CO2 that
accumulated from respiration in the dark from photosynthesis and so alter the pH.
However this did not explain the rapidity of stomatal movements.
Another idea arose with the discovery of glucose-1-phosphate and that this compound can be
reversibly converted to starch by a series of dephosphorylation and phosphorylation reaction catalysed
by a ph sensitive enzyme ‘phosphorylase’. This simple conversion did not seem to alter the osmotic
concentration of the cells. It was then suggested that in order to affect osmotic relationships the
glucose-1-phosphate had to be split into glucose and phosphate. However the reverse reaction is not
readily possible since it requires energy from adenosine triphosphate (ATP) and hexokinase. If these
two compounds are adequate then the stomata close.
In the presence of light the guard cells use up CO2 in photosynthesis and this raises the pH favouring
the conversion of starch to glucose-1-phosphate. When light is absent photosynthesis stops and CO2
accumulates lowering the pH and glucose-1-phosphate is converted back to starch.
Factors affecting transpiration
Relative humidity
The air in the intercellular spaces of the leaf is saturated with water vapour while that in the
atmosphere has usually, a relative humidity of less than 100 %. This creates a gradient in vapour
124
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

pressure so that water diffuses out of the leaf into the atmosphere. If relative humidity of the
atmosphere is very low then the gradient will be very steep and the rate of transpiration fast. The
opposite is true.
Temperature
At a constant humidity increasing the temperature results in an increased rate of transpiration. This is
because of the increases in water holding capacity of the air and an increased kinetic energy of the
water molecules.
Light
Stomata are normally open in daylight and closed at night; transpiration is then obviously faster during
the day. Direct sunlight also raises the temperature of the leaf and thus increase the kinetic energy of
water molecules.
Wind
In still air water vapour accumulates around the leaf and this slows down transpiration by offering a
resistance to diffusion. Air currents will tend to sweep away water vapour and thus keep the diffusion
gradient steep.
Soil moisture
Any factor which reduces water supply to the leaves reduces the rate of transpiration because the
stomata tend to close. In some cases this will only occur after wilting.
Plant structure
Some plants, the xerophytes, have morphological modifications that reduce transpiration. For instance,
the surface area/volume ratio of the plant may be reduced; a very thick cuticle may be present; stomata
may be sunk in the epidermis; and, leaves may fold to protect the stomata.
Functions of transpiration
Transpiration may be necessary for uptake of mineral salts from the soil although some experiments
show an independence of the latter process from the former. Evaporation of moisture at the leaf
surface cools the plant; however, the leaf temperature rarely drops by more than 3 – 5 0C. On the other
hand transpiration is undesirable , especially when the plant loses more water than it can absorb.
In brief, transpiration may be taken as a “necessary evil”.

Movement of water up the plant

It has been shown that water is absorbed from the soil by roots and that it is lost through the leaf
surfaces to the atmosphere. In order to bridge these two points, a system to conduct the water up the
plant is necessary. This upward movement of water occurs through the xylem elements of the vascular
tissues.
Root pressure can play a part in “forcing” the water up the water; however, this mechanism cannot
cope with a rise of several meters and the considerable height of some trees. It is known that water
rises to a certain height in fine capillary tubes. There is therefore a possibility that capillary action in
the xylem strands contributes something to the upward movement of water although this would be
insignificant in tall plants.

Water is lost to the atmosphere by evaporating from the surfaces of mesophyll cells. Tension builds up
and water diffuses from the surrounding cells which in turn develop a diffusion pressure deficit higher
than that found in the xylem. Water will the move out of the xylem. This will exert a pull on the water
in the xylem and will tend to “lift” it up. For this to function the water molecules should move up in
mass, i.e. be cohesive; the suction from the leaves should be sufficient to lift the water to great
heights; no air should enter the system and break the continuity of the water column; and the xylem
elements should be rigid enough to resist collapse when the water moves under tension. However this
should not be taken as the final explanation since work on the subject is still going on.

125
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

PHLOEM TRANSPORT
Structure of phloem
 Remember the structure of phloem cells (sieve areas (plates) & sieve pores)

Phloem is a complex permanent tissue, which is specialized for the conduction of food and other
organic substances. Phloem is also a heterogenous tissue, made up of four different types of cellular
elements, namely,
 Sieve tubes
 Companion cells
 Phloem parenchyma and
 Phloem fibres
Of these, the sieve tubes and the companion cells are directly involved in the translocation of the
organic substances. Hence, they are commonly described as essential elements. Phloem parenchyma
and phloem fibres are described as associated elements since they play only a supporting role in the
process.
The sieve tubes, the companion cells and the phloem parenchyma represent the living components of
the tissue while phloem fibres represent the only nonliving component of the tissue.
Phloem is commonly described as a living, complex permanent tissue.

Figure 13.1 Structure of phloem

Contents of phloem cells

 Water
 90 % of solutes are sugars
 Sucrose is primary (30% of phloem sap)
 Sugar alcohols
 Other reducing sugars such as raffinose, stachyose.
Model for phloem transport
Flow from Source to Sink
Food, primarily sucrose is transported by the vascular tissue called phloem from a source to a sink.
Unlike transpiration's one-way flow of water sap, food in phloem sap can be transported in any
direction needed so long as there is a source of sugar and a sink able to use, store or remove the sugar.
The source and sink may be reversed depending on the season, or the plant's needs. Sugar stored in
roots may be mobilized to become a source of food in the early spring when the buds of trees, the sink,
need energy for growth and development of the photosynthetic apparatus.

126
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Phloem sap is mainly water and sucrose, but other sugars, hormones and amino acids are also
transported. The movement of such substances in the plant is called translocation.
The Pressure Flow or Mass Flow Hypothesis
The accepted mechanism needed for the translocation of sugars from source to sink is called the
pressure flow hypothesis. (see diagram below)
As glucose is made at the source (by photosynthesis for example) it is converted to sucrose (a
dissacharide). The sugar is then moved into companion cells and into the living phloem sieve tubes by
active transport. This process of loading at the source produces a hypertonic condition in the phloem.
Water in the adjacent xylem moves into the phloem by osmosis. As osmotic pressure builds the
phloem sap will move to areas of lower pressure.
At the sink osmotic pressure must be reduced. Again active transport is necessary to move the sucrose
out of the pholem sap and into the cells which will use the sugar -- converting it into energy, starch, or
cellulose. As sugars are removed osmotic pressure decreases and water moves out of the phloem.
The Pressure Flow or Mass Flow Hypothesis

Figure 13.2 Diagram illustrating the Pressure Flow of Mass Flow Hypothesis

Phloem Loading

Figure 13.3a Diagram illustrating sucrose loading into the phloem for transport

127
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Apoplast and symplast routes are used during transport at the source. ATP and H+ carrier pump
protons out of sieve tube and K+ are transported in to sieve tube. There is Co transport of H+ and
sucrose into sieve tube which lowers water potential of sieve tube.

Movement in phloem can be bi-directional and rate of flow chiefly depends on strength of sink.

Figure 13.3b Diagram showing Pressure flow in the phloem of flowering plants
Activity
Discuss the various hypotheses underlying phloem transportation.

CHAPTER 18

PHOTOSYNTHESIS

Photosynthesis is the series of reactions by which light energy is utelised to synthesise carbohydrates
(monosaccharide, disaccharides and polysaccharides) CO2 and water in the presence of chlorophyll
and with oxygen as a by-product. This process is the prime source of all organic molecules, animal or
vegetable.
Essential requirements
Raw materials
Water and carbon dioxide are the raw materials in photosynthesis. Water is absorbed from the by the
root hairs and translocation up the plant into the leaves. Carbon dioxide enters the atmosphere through
the stomata and is also obtained from the respiration of the leave mesophyll cells themselves.
Energy
The source of energy for photosynthesis is sunlight. Radiant energy or light consists of a spectrum of
which visible light is a small portion. The visible light in turn consists of light of different colour and
wave lengths. The blue and red light are the most absorbed by chlorophyll and as such are affective in
photosynthesis.
Pigments

128
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The pigment that is necessary for the absorption and conversion of light energy into chemical energy
is chlorophyll. This green pigment is found in the chloroplasts of the mesophyll cells. Two types of
chlorophyll occur in higher plants, that is, chlorophyll a and b. The term chlorosis refers to a condition
when the plant cannot synthesise chlorophyll due to a deficiency of an essential factor in chlorophyll
synthesis, such as magnesium.

Chemistry of photosynthesis
Photosynthesis can be summerised by the equation:
6CO2 + 12H2O ______________ C6H12O6 + 6O2 +6H2O
However the whole process is not as simple as the equation suggest; it is indeed a process that
involves a series of complex enzyme catalysed reactions. Two stages have been identified in
photosynthesis. The first stage occurs only in light and called the light reaction while the second, the
dark reaction. This can be represented as:
A ___photochemical_______ B ___chemical______ C
Under periods of continuous light A B proceeds faster than B C so that B will tend to accumulate. A
period of darkness will allow B C to occur while A B stops. Therefore for a given amount of light
more C will be formed if a dark period is present.
Light reaction
Photosystem I and II
Photosystems are light-absorbing complexes in the thylakoid membranes that are present in
photosynthetic organisms. There are two types of photosystems: Photosystem I and Photosystem II.
Each has one primary photochemical reaction center (either chlorophyll P700 or P680) and a set of
accessory pigments to absorb additional light.
There is a special molecule called chlorophyll a P700, located in photosystem I, which absorbs light
best at 700 nanometers (nm). It also contains other accessory pigments. Another special chlorophyll a
molecule in photosystem II is called chlorophyll a P680 because it absorbs best at 680 nm.
Photosystem II also contains chlorophyll b and other accessory pigments.
Photosystem II
PS II is an extremely complex, highly organized transmembrane structure that contains a water-
splitting complex, chlorophylls and carotenoid pigments, a reaction center (P680), pheophytin (a
pigment similar to chlorophyll), and two quinones. It uses the energy of sunlight to transfer electrons
from water to a mobile electron carrier in the membrane called plastoquinone:
H2O → P680 → P680* → plastoquinone
Plastoquinone, in turn, transfers electrons to cytb6, which feeds them into PS I.
The water-splitting complex
The step H2O → P680 is performed by a poorly understood structure embedded within PS II called
the water-splitting complex or the oxygen-evolving complex. It catalyzes a reaction that splits water
into electrons, protons and oxygen:
2H2O → 4H+ + 4e− + O2
The electrons are transferred to special chlorophyll molecules (embedded in PS II) that are promoted
to a higher-energy state by the energy of photons.
The reaction center
The excitation P680 → P680*of the reaction center pigment P680 occurs here. These special
chlorophyll molecules embedded in PS II absorb the energy of photons, with maximal absorption at
680 nm. Electrons within these molecules are promoted to a higher-energy state. This is one of two
core processes in photosynthesis, and it occurs with astonishing efficiency (greater than 90%) because,
in addition to direct excitation by light at 680 nm, the energy of light first harvested by antenna
proteins at other wavelengths in the light-harvesting system is also transferred to these special
chlorophyll molecules.

129
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

This is followed by the step P680*→ pheophytin, and then on to plastoquinone, which occurs within
the reaction center of PS II. High-energy electrons are transferred to plastoquinone before it
subsequently picks up two protons to become plastoquinol. Plastoquinol is then released into the
membrane as a mobile electron carrier.
This is the second core process in photosynthesis. The initial stages occur within picoseconds, with an
efficiency of 100%. The seemingly impossible efficiency is due to the precise positioning of
molecules within the reaction center. This is a solid-state process, not a chemical reaction. It occurs
within an essentially crystalline environment created by the macromolecular structure of PS II. The
usual rules of chemistry (which involve random collisions and random energy distributions) do not
apply in solid-state environments.
Link of water-splitting complex and chlorophyll excitation[edit]
When the chlorophyll passes the electron to pheophytin, it obtains an electron from P680*. In turn,
P680* can oxidize the Z (or YZ) molecule. Once oxidized, the Z molecule can derive electrons from
the oxygen-evolving complex.

Dark reaction
The dark reaction is sensitive to temperature and not light while the light reaction is sensitive to light
and not temperature. This second stage is essentially a reductive fixation of carbon dioxide and the
elucidation of the course of the reactions was made possible by the use of paper chromatography and
radioactive tracers.
Firstly, CO2 combines with a 5-carbon compound known as ribulose diphosphate. A 6-carbon unstable
intermediate compound is thus formed and this split into two molecules of phosphoglyceric acid. This
acid is reduced by hydrogen from NADPH2 into an aldose (phosphogylceraldehyde) and a ketose
(dihydroxyacetone phosphate). These 3-carbon sugars are interconvertible and a combination of two
molecules will form a hexose such as glucose whose molecules can be converted into other
carbohydrates. The trioses can also form fats and amono acids.
The CO2 acceptor, ribulose diphosphate is regenerated during photosynthesis. This is accomplished by
the conversion of the triose molecules into several intermediate compounds. Eventually ribulose
monophosphate is formed. For this compound to be converted to ribulose diphosphate ATP is
consumed.
Therefore the dark reaction is cyclic except that some molecules are drawn out of the cycle as
carbohydrates, fats, and amino acids.

130
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Factors affecting photosynthesis

Carbon dioxide
At a high light intensity and constant temperature, the rate of photosynthesis is proportional to the
concentration of CO2 within a certain range above which there is no effect. At this point some other
factor limits photosynthesis. Another important aspect is the rate of diffusion of CO2 through the
stomata and into the leaf mesophyll. A rapid rate of diffusion would mean an increased supply of CO2
and therefore a faster rate of photosynthesis.
Light
The light energy stored during photosynthesis is supplied by light. Intensity, quality and duration of
the light are important. In general, an increase in light intensity is accompanied by an increase in the
rate of photosynthesis until another factor becomes limiting. However, the light intensity required for
maximum photosynthesis varies with species since some plants like the shade and others the full sun.
Light intensity also has an indirect effect on photosynthesis due its influence on stomatal behavior and
therefore the diffusion off CO2.
Temperature
When no other factor is limiting, the rate of photosynthesis increases with temperature up to a point
which varies with species. A further increase in temperature results in a rapid decline in
photosynthesis because the protoplasm is injured. The main effects of temperature are on the enzyme
catalysed reactions of the dark stage.
Water
The chloroplasts are usually supplied with adequate water for photosynthesis. In fact, less than 1% of
the water absorbed by the plant is used in photosynthesis so that the effects of water would mainly be
indirect. For instance the results of wilting would be the closure of the stomata and a consequent
reduction in the supply of CO2.

Activity
Photosynthesis comprises a ‘light reaction’ and a ‘dark reaction’. Give an outline of each of these two
stages of photosynthesis. Explain the factors that affect the rate of photosynthesis.
C3,C4,CAM

131
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

C3 CARBON FIXATION
C3 carbon fixation is one of three metabolic pathways for carbon fixation in photosynthesis, along
with C4 and CAM. This process converts carbon dioxide and ribulose bisphosphate (RuBP, a 5-carbon
sugar) into 3-phosphoglycerate through the following reaction:
CO2 + H2O + RuBP → (2) 3-phosphoglycerate
This reaction occurs in all plants as the first step of the Calvin–Benson cycle. In C4 plants, carbon
dioxide is drawn out of malate and into this reaction rather than directly from the air.

Cross section of a C3 plant, specifically of an Arabidopsis thaliana leaf. Vascular bundles shown.
Plants that survive solely on C3 fixation (C3 plants) tend to thrive in areas where sunlight intensity is
moderate, temperatures are moderate, carbon dioxide concentrations are around 200 ppm or
higher, and groundwater is plentiful. Examples include rice and barley.
C3 plants cannot grow in very hot areas because RuBisCO incorporates more oxygen into RuBP as
temperatures increase. This leads to photorespiration (also known as the oxidative photosynthetic
carbon cycle, or C2 photosynthesis), which leads to a net loss of carbon and nitrogen from the plant
and can therefore limit growth. In dry areas, C3 plants shut their stomata to reduce water loss, but this
stops CO2 from entering the leaves and therefore reduces the concentration of CO2 in the leaves. This
lowers the CO2:O2 ratio and therefore also increases photorespiration. C4 and CAM plants have
adaptations that allow them to survive in hot and dry areas, and they can therefore out-compete
C3 plants in these areas.

C4 carbon fixation
C4 carbon fixation or the Hatch-Slack pathway is a photosynthetic process in some plants. It is the
first step in extracting carbon from carbon dioxide to be able to use it in sugar and other biomolecules.
It is one of three known processes for carbon fixation. The C4 in one of the names refers to the 4-
carbon molecule that is the first product of this type of carbon fixation.
C4 fixation is an elaboration of the more common C3 carbon fixation and is believed to have evolved
more recently. C4 overcomes the tendency of the enzyme RuBisCO to wastefully fix oxygen rather
than carbon dioxide in the process of photorespiration. This is achieved by ensuring that RuBisCO
works in an environment where there is a lot of carbon dioxide and very little oxygen. CO2 is shuttled
via malate or aspartate from mesophyll cells to bundle-sheath cells. In these bundle-sheath cells CO2 is
released by decarboxylation of the malate. C4 plants use PEP carboxylase to capture more CO2 in the
mesophyll cells. PEP Carboxylase (3 carbons) binds to CO2 to make oxaloacetic acid (OAA). The
132
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

OAA then makes malate (4 carbons). Malate enters bundle sheath cells and releases the CO2. These
additional steps, however, require more energy in the form of ATP. Using this extra energy, C4 plants
are able to more efficiently fix carbon in drought, high temperatures, and limitations of nitrogen or
CO2. Since the more common C3pathway does not require this extra energy, it is more efficient in the
other conditions.

C4 PATHWAY

NADP-ME type C4 pathway

NAD-ME type C4 pathway

The C4 pathway was elucidated by Marshall Davidson Hatch and C. R. Slack, in Australia, in 1966; it
is sometimes called the Hatch-Slack pathway.
In C3 plants, the first step in the light-independent reactions of photosynthesis involves the fixation of
CO2 by the enzyme RuBisCO into 3-phosphoglycerate. However, due to the
dual carboxylase and oxygenase activity of RuBisCo, some part of the substrate is oxidized rather
than carboxylated, resulting in loss of substrate and consumption of energy, in what is known
as photorespiration. In order to bypass the photorespiration pathway, C4 plants have developed a
mechanism to efficiently deliver CO2 to the RuBisCO enzyme. They utilize their specific leaf anatomy
where chloroplasts exist not only in the mesophyll cells in the outer part of their leaves but in
the bundle sheath cells as well. Instead of direct fixation to RuBisCO in the Calvin cycle, CO2 is
incorporated into a 4-carbon organic acid, which has the ability to regenerate CO2 in the chloroplasts
of the bundle sheath cells. Bundle sheath cells can then utilize this CO2 to generate carbohydrates by
the conventional C3 pathway.
The first step in the pathway is the conversion of pyruvate to phosphoenolpyruvate (PEP), by the
enzyme pyruvate orthophosphate dikinase. This reaction requires inorganic phosphate
and ATP plus pyruvate, producing phosphoenolpyruvate, AMP, and inorganic pyrophosphate (PPi).
The next step is the fixation of CO2 into oxaloacetate by the enzyme PEP carboxylase. Both of these
steps occur in the mesophyll cells:

133
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

pyruvate + Pi + ATP → PEP + AMP + PPi

PEP + CO2 → oxaloacetate
PEP carboxylase has a lower Km for HCO−3 — and, hence, higher affinity — than RuBisCO.
Furthermore, O2 is a very poor substrate for this enzyme. Thus, at relatively low concentrations of
CO2, most CO2 will be fixed by this pathway.
The product is usually converted to malate, a simple organic compound, which is transported to the
bundle-sheath cells surrounding a nearby vein. Here, it is decarboxylated to produce
CO2 and pyruvate. The CO2 now enters the Calvin cycle and the pyruvate is transported back to
the mesophyll cell.
Since every CO2 molecule has to be fixed twice, first by 4-carbon organic acid and second by
RuBisCO, the C4 pathway uses more energy than the C3pathway. The C3 pathway requires 18
molecules of ATP for the synthesis of one molecule of glucose, whereas the C4 pathway requires 30
molecules of ATP. This energy debt is more than paid for by avoiding losing more than half of
photosynthetic carbon in photorespiration as occurs in some tropical plants, making it an adaptive
mechanism for minimizing the loss.
There are several variants of this pathway:
1. The 4-carbon acid transported from mesophyll cells may be malate, as above, or aspartate.
2. The 3-carbon acid transported back from bundle-sheath cells may be pyruvate, as above,
or alanine.
3. The enzyme that catalyses decarboxylation in bundle-sheath cells differs. In maize and sugarcane,
the enzyme is NADP-malic enzyme; in millet, it is NAD-malic enzyme; and, in Panicum
maximum, it is PEP carboxykinase.
C4 Kranz leaf anatomy

Cross section of a maize leaf, a C4 plant. Kranz anatomy (rings of cells) shown. Drawing based on
microscopic images courtesy of Cambridge University Plant Sciences Department.
The C4 plants often possess a characteristic leaf anatomy called kranz anatomy, from the German
word for wreath. Their vascular bundles are surrounded by two rings of cells; the inner ring, called
bundle sheath cells, contains starch-rich chloroplasts lacking grana, which differ from those
in mesophyll cells present as the outer ring. Hence, the chloroplasts are called dimorphic. The primary
function of kranz anatomy is to provide a site in which CO2 can be concentrated around RuBisCO,
thereby avoiding photorespiration.
In order to maintain a significantly higher CO2 concentration in the bundle sheath compared to the
mesophyll, the boundary layer of the kranz has a low conductance to CO2, a property that may be
enhanced by the presence of suberin.
Although most C4 plants exhibit kranz anatomy, there are, however, a few species that operate a
limited C4 cycle without any distinct bundle sheath tissue. Carboxylation enzymes in the cytosol can,
therefore, be kept separate from decarboxylase enzymes and RuBisCO in the chloroplasts, and a
134
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

diffusive barrier can be established between the chloroplasts (which contain RuBisCO) and the
cytosol. This enables a bundle-sheath-type area and a mesophyll-type area to be established within a
single cell. Although this does allow a limited C4 cycle to operate, it is relatively inefficient, with the
occurrence of much leakage of CO2 from around RuBisCO.
The evolution and advantages of the C4 pathway

C4 plants have a competitive advantage over plants possessing the more common C3 carbon
fixation pathway under conditions of drought, high temperatures, and nitrogen or CO2 limitation.
When grown in the same environment, at 30 °C, C3 grasses lose approximately 833 molecules of
water per CO2 molecule that is fixed, whereas C4 grasses lose only 277. This increased water use
efficiency of C4 grasses means that soil moisture is conserved, allowing them to grow for longer in
arid environments.
C4 carbon fixation has evolved on up to 61 independent occasions in 19 different families of plants,
making it a prime example of convergent evolution. This convergence may have been facilitated by
the fact that many potential evolutionary pathways to a C4 phenotype exist, many of which involve
initial evolutionary steps not directly related to photosynthesis. Today, C4 plants represent about 5%
of Earth's plant biomass and 3% of its known plant [Link] this scarcity, they account for
about 23% of terrestrial carbon [Link] the proportion of C4 plants on earth could
assist biosequestration of CO2 and represent an important climate change avoidance strategy. Present-
day C4 plants are concentrated in the tropics and subtropics (below latitudes of 45°) where the high air
temperature contributes to higher possible levels of oxygenase activity by RuBisCO, which increases
rates of photorespiration in C3plants.
Plants that use C4 carbon fixation

Maize (or corn) is a common C4 plant.

About 8,100 plant species use C4 carbon fixation, which represents about 3% of all terrestrial species
of plants. All these 8,100 species are angiosperms. These include the food crops maize, sugar
cane, millet, and sorghum.
Members of the sedge family Cyperaceae, and members of numerous families of Eudicots -
including Asteraceae (the daisy family), Brassicaceae (the cabbage family), and Euphorbiaceae (the
spurge family)- also use C4.

CRASSULACEAN ACID METABOLISM

Crassulacean acid metabolism, also known as CAM photosynthesis, is a carbon fixation pathway
that evolved in some plants as an adaptation to arid conditions. In a plant using full CAM,
the stomata in the leaves remain shut during the day to reduce evapotranspiration, but open at night to
collect carbon dioxide (CO2). The CO2 is stored as the four-carbon acid malate in vacuoles at night,
and then in the daytime, the malate is transported to chloroplasts where it is converted back to CO2,
which is then used during photosynthesis. The pre-collected CO2 is concentrated around the
135
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

enzyme RuBisCO, increasing photosynthetic efficiency. This metabolism was first studied in plants of
the family Crassulaceae. These mainly include succulents. The first time it was studied, Crassula was
used as a model organism. The word "crassulacean" is from the Latin word crassus, meaning “thick;”
succulent plants have leaves that are thick and full of moisture, and they often have a waxy or hairy
coating to help prevent evaporation.
Overview: a two-part cycle

CAM is an adaptation for increased efficiency in the use of water, and so is typically found in plants
growing in arid conditions.
During the night

During the night, a plant employing CAM has its stomata open, allowing CO2 to enter and be fixed as
organic acids that are stored in vacuoles. During the day the stomata are closed (thus preventing water
loss), and the carbon is released to the Calvin cycle so that photosynthesis may take place.
The carbon dioxide is fixed in the cytoplasm of mesophyll cells by a PEP reaction similar to that
of C4 pathway. But, unlike the C4 mechanism, the resulting organic acids are stored in vacuoles for
later use; that is, they are not immediately passed on to the Calvin cycle. The latter cannot operate
during the night because the light reactions that provide it with ATP and NADPH cannot take place.
During the day

During the day, the CO2-storing organic acids are released from the vacuoles of the mesophyll cells
and enter the stroma of the chloroplasts where an enzyme releases the CO2, which then enters into the
Calvin cycle.
Benefits

The most important benefit of CAM to the plant is the ability to leave most leaf stomata closed during
the day. Plants employing CAM are most common in arid environments, where water comes at a
premium. Being able to keep stomata closed during the hottest and driest part of the day reduces the
loss of water through evapotranspiration, allowing such plants to grow in environments that would
otherwise be far too dry. Plants using only C3 carbon fixation, for example, lose 97% of the water they
uptake through the roots to transpiration - a high cost avoided by plants able to employ CAM.

Comparison with C4 metabolism

The C4 pathway bears resemblance to CAM; both act to concentrate CO2 around RuBisCO, thereby
increasing its efficiency. CAM concentrates it temporally, providing CO2 during the day, and not at
night, when respiration is the dominant reaction. C4 plants, in contrast, concentrate CO2 spatially, with
a RuBisCO reaction centre in a "bundle sheath cell" being inundated with CO2. Due to the inactivity
required by the CAM mechanism, C4 carbon fixation has a greater efficiency in terms
of PGA synthesis
Biochemistry

Plants with CAM must control storage of CO2 and its reduction to branched carbohydrates in space
and time.
At low temperatures (frequently at night), plants using CAM open their stomata, CO2 molecules
diffuse into the spongy mesophyll's intracellular spaces and then into the cytoplasm. Here, they can
meet phosphoenolpyruvate (PEP), which is a phosphorylated triose. During this time, the plants are
synthesizing a protein called PEP carboxylase kinase (PEP-C kinase), whose expression can be
136
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

inhibited by high temperatures (frequently at daylight) and the presence of malate. PEP-C kinase
phosphorylates its target enzyme PEP carboxylase (PEP-C). Phosphorylation dramatically enhances
the enzyme's capability to catalyze the formation of oxaloacetate, which can be subsequently
transformed into malate by NAD+ malate dehydrogenase. Malate is then transported via malate
shuttles into the vacuole, where it is converted into the storage form malic acid. In contrast to PEP-C
kinase, PEP-C is synthesized all the time but almost inhibited at daylight either
by dephosphorylation via PEP-C phosphatase or directly by binding malate. The latter is not possible
at low temperatures, since malate is efficiently transported into the vacuole, whereas PEP-C kinase
readily inverts dephosphorylation.
In daylight, plants using CAM close their guard cells and discharge malate that is subsequently
transported into chloroplasts. There, depending on plant species, it is cleaved into pyruvate and
CO2 either by malic enzyme or by PEP carboxykinase. CO2 is then introduced into the Calvin cycle, a
coupled and self-recovering enzyme system, which is used to build branched carbohydrates. The by-
product pyruvate can be further degraded in the mitochondrial citric acid cycle, thereby providing
additional CO2 molecules for the Calvin Cycle. Pyruvate can also be used to recover PEP via pyruvate
phosphate dikinase, a high-energy step, which requires ATP and an additional phosphate. During the
following cool night, PEP is finally exported into the cytoplasm, where it is involved in fixing carbon
dioxide via malate.
Ecological and taxonomic distribution of CAM-using plants

The majority of plants possessing CAM are either epiphytes (e.g., orchids, bromeliads) or
succulent xerophytes (e.g., cacti, cactoid Euphorbias), terrestrial bromeliads; wetland
plants (e.g., Isoetes, Crassula (Tillaea), Lobelia; and in one halophyte, Mesembryanthemum
crystallinum; one non-succulent terrestrial plant, (Dodonaea viscosa) and one mangrove associate
(Sesuvium portulacastrum).
Plants which are able to switch between different methods of carbon fixation include Portulacaria
afra, better known as Dwarf Jade Plant, which normally uses C3 fixation but can use CAM if it is
drought-stressed, and Portulaca oleracea, better known as Purslane, which normally uses C4fixation
but is also able to switch to CAM when drought-stressed

137
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

C3 PLANT C4 PLANT
1 Only C3 cycle found Both C3 and C4 cycles found
2 The efficiency of CO2 absorption at low The efficiency of CO2 absorption at low
concentration is far less and hence they are concentration is quite high and hence they are
less efficient more efficient plants
3 The CO2 acceptor is Ribulose-1.5- The CO2 acceptor is phosphor enol pyruvate
diphosphate
4 The first stable product is phosphoglyceric Oxaloacetate(OOA) is the first stable product
acid(PGA)
5 Plants show one type of Plants show dimorphic type of
chloroplast(monomorphic) chloroplast(mesophyll and bundle sheath)-
Kranzy type of anatomy
6 In each chloroplast,two pigment In each chloroplast of bundle sheath cells,the
systems(PSI and II) are present. PSII is [Link] on mesophyll chloroplasts
for NADPH + H+
7 The Calvin cycle enzymes are present in Calvin cycle enzymes absent in mesophyll
mesophyll [Link] calvin cycle chloroplast but occurs in bundle sheath cells
occurs. chloloplasts
8 The carbon dioxide compensation point is The carbon dioxide compensation point is 0-
50-150ppm CO2 10ppm CO2
9 Bundle sheath cells are unspecialised The bundle sheath cells are high developed with
unusual construction of organelles
10 The optimal temperature for the process is Temperature 30-45oC hence are warm climate
10-25oC plants. Rate double than in C3.
11 18ATPs are required to synthesize one 30ATPs are required to synthesise one glucose
glucose molecule molecule.
12 The light saturation intensityreaches in It is difficult to reach saturation even in full
range of 1000-4000 ft. C sunlight.
13 Photorespiration is present and easily Photorespiration is present only to a slight
detectable. degree or absent.

CHAPTER 20

RESPIRATION

One characteristic of all living things is that they respire. Respiration is basically a process whereby
complex substances are broken down into simpler ones with the release of energy in utilisable forms.
The energy is required for the maintenance of life so that respiration is indeed an important reaction.
Respiration occur without the uptake of oxygen – this is anaerobic respiration of fermandation.
Fermentation is common with fungi and bacteria and may also occur in some higher plants if the
oxygen concentration is low. In most cases respiration is aerobic in that molecular oxygen is utilised
during the process. This type of respiration (aerobic) can be summerised by the equation below.
C6H12O6 + 6O2--------------------- 6CO2 + 6H2O + Energy
The biological combustion of sugars (and at times fats) is accomplished through a series of complex
reactions that involve specific enzymes and energy carriers found in the mitochondria of the cell.
Chemistry of respiration
Aerobic respiration occurs in two steps – the oxidation of carbohydrate to pyruvic acid with small
amounts of energy being produced (glycolisis); and the subsequent oxidation of pyruvic acid through
the Krebs Cycle in which much of the energy is released.
Oxidation is the loss of oxygen whilst reduction is the gain of oxygen.
138
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Glycolysis
The first stage of respiration common to both types is glycolysis. Glycolysis is the first step in the
breakdown of glucose to extract energy for cellular metabolism. Glycolysis consists of an energy-
requiring phase followed by an energy-releasing phase. This first series of metabolic reactions occurs
in the cytoplasm not the mitochondrion
Glucose undergoes phosphorylation, taking up a phosphate group from a molecule of ATP (reducing
it to ADP) and becoming glucose phosphate. This is then rearranged into another molecule – fructose
phosphate, which is again phosphorylated to become fructose biphosphate. This six carbon sugar is
then split into two smaller molecules which are each called triose phosphate (3-carbon)
Finally, two molecules of ADP are phosphorylated for each triose phosphate to become ATP, and the
sugar then has two hydrogen atoms removed using dehydrogenase enzymes to become a molecule
known as pyruvate
The hydrogen atoms are accepted by a coenzyme called NAD (nicotinamide adenine dinucleotide),
reducing the coenzyme to reduced NAD (or NADH2). Overall, as two ATP were used to
phosphorylate the sugar and four were eventually made, it is said that the net ATP production per
glucose molecule in glycolysis is 2 ATP. Two molecules of pyruvate and two molecules of reduced
NAD were also produced.

Krebs Cycle

139
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The next stage involved only in aerobic respiration begins with the pyruvate molecules produced
from glycolysis. This part of aerobic respiration happens in the mitochondrial matrix and the
pyruvate is transported there from the cytoplasm
In the link reaction, named so as it links glycolysis to Krebs, a molecule of pyruvate is first
decarboxylated (has carbon dioxide removed) using decarboxylase enzymes and dehydrogenated
(has hydrogen removed) using dehydrogenase enzymes. The hydrogen atoms are accepted by a
molecule of NAD. The result is an acetyl group which binds with coenzyme-A to form acetyl
coenzyme A. The coenzyme-A leaves the molecule (leaving it as acetate) and is recycled to be used
again
The acetate is a 2-carbon molecule which then joins a molecule of oxaloacetate (a 4-carbon
compound) to form citrate (a 6-carbon compound). Citrate is then decarboxylated and
dehydrogenated to form a 5-carbon acid, which also has hydrogen and carbon dioxide removed,
forming a 4-carbon acid. At this stage, substrate-level phosphorylation occurs as in glycolysis – an
inorganic phosphate is removed from the compound and attached to a molecule of ADP to form ATP
A molecule of FAD is then reduced, as is a molecule of FAD, and the removal of these four hydrogen
atoms forms oxaloacetate, and so the cycle continues
There is one complete turn of the cycle for each molecule of pyruvate. The end products of Krebs
cycle for each glucose include two molecules of ATP produced directly during Krebs, six lots of
reduced NAD and two reduced FAD, as well as four molecules of carbon dioxide, a waste by-product.
The link reaction produces a further two reduced NAD and two carbon dioxide molecules added on to
this total
The molecules of reduced NAD and FAD will enter the electron transport chain and donate their
accepted hydrogen atoms so that ATP can be synthesised via oxidative phosphorylation. Generally,
one NADH2 yields 2.5 ATP and one FADH2 yields 1.5 ATP

140
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Electron Transport Chain

Glycolysis, the conversion of PA to acetyl-CoA, and the Krebs Cycle complete the breakdown
of glucose.
• Up to this point:
4 ATP (2 from glycolysis, 2 from Krebs)
f PA, 6 from Krebs)

• NADH + H+ and FADH2 carry electrons to an electron transport chain, where additional ATP is
produced.
• Steps:
1. NADH + H+ and FADH2 carry electrons to the ETC.
2. As electrons move down the energy gradient in the inner membrane of the mitochondrion, hydrogen
ions are pumped across the membrane into the matrix.
3. An electrochemical gradient forms, and hydrogen ions diffuse through channels that contain ATP
synthetase.
4. The energy generated by the movement of hydrogen ions is used by ATP synthetase to make ATP.

STARCH DEGRADATION

Starch is the major carbon reserve in most plants. It is a mixture of amylose and amylopectin and is
deposited as granules inside plastids (chloroplasts in leaves, amyloplasts in non-photosynthetic
tissues). The initial attack on starch granules in leaves and non-photosynthetic tissues is by a-amylase
and a debranching enzyme. Oligosaccharides released during starch degradation, such as maltose,
maltotriose and maltotetraose, are hydrolysed to glucose by a-glucosidase.

141
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Pathways of starch metabolism. Numbers refer to the following enzymes: 1, ß-amylase; 2, α-amylase;
3, starch phosphorylase; 4, glucosidase; 5, hexose kinase; 6, phosphoglucomutase; 7, glucose 6-
phosphate isomerase. (Original drawing courtesy David Day)

Reactions in plant tissues leading to degradation of sucrose to hexose monophosphates are outlined in
Figure 2.20. The first step is cleavage of the glycosidic bond by either invertase (Equation 2.1)

Sucrose + H2O → D-glucose + D-fructose (2.1)

ATP

The function of ATP is to provide energy for cellular processes such as muscle contraction, cell
division, protein synthesis and DNA replication.
ATP is formed from one molecule of adenosine joined to three phosphate groups (Pi). ATP can
be broken down into ADP and Pi, and reformed in a process called phosphorylation.

142
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Functions in cells
 Metabolism, synthesis, and active transport
ATP is consumed in the cell by energy-requiring (endergonic) processes and can be generated by
energy-releasing (exergonic) processes. In this way ATP transfers energy between spatially
separate metabolic reactions. ATP is the main energy source for the majority of cellular functions.
This includes the synthesis of macromolecules, including DNA and RNAand proteins. ATP also plays
a critical role in the transport of macromolecules across cell membranes,
[Link] and endocytosis.
 Roles in cell structure and locomotion
ATP is critically involved in maintaining cell structure by facilitating assembly and disassembly of
elements of thecytoskeleton. In a related process, ATP is required for the shortening of actin and
myosin filament cross bridges required for muscle. This latter process is one of the main energy
requirements of animals and is essential for locomotion and respiration.
 Cell signalling
Extracellular signalling
ATP is also a signalling molecule. ATP, ADP, or adenosines are recognised by purinergic receptors.
Purinoreceptors might be the most abundant receptors in mammalian tissues.[35]
Intracellular signalling
ATP is critical in signal transduction processes. It is used by kinases as the source of phosphate groups
in their phosphate transfer reactions.
 DNA and RNA synthesis
In all known organisms, the Deoxyribonucleotides that make up DNA are synthesized by the action
of ribonucleotide reductase (RNR) enzymes on their corresponding ribonucleotides. These enzymes
reduce the sugar residue from ribose to deoxyribose by removing oxygen from the 2' hydroxyl group;
the substrates are ribonucleoside diphosphates and the products deoxyribonucleoside diphosphates
(the latter are denoted dADP, dCDP, dGDP, and dUDP respectively.)
 Amino acid activation in protein synthesis
Aminoacyl-tRNA synthetase enzymes utilise ATP as an energy source to attach a tRNA molecule to
its specific amino acid, forming an aminoacyl-tRNA complex, ready for translation at ribosomes. The
energy is made available by ATP hydrolysis to adenosine (AMP) as two phosphate groups are
removed. Amino acid activation refers to the attachment of an amino acid to its Transfer RNA
(tRNA).

CHAPTER 21

PRINCIPLES OF CROP BREEDING AND BIOTECHNOLOGY

 describe the structure of a chromosome

 describe the structure of DNA
 explain DNA replication
 describe protein synthesis starting from DNA
 describe the process of mitosis
 explain the significance of mitosis in crop production
 describe the process of meiosis
 explain the significance of meiosis in crop reproduction
 compare and contrast mitosis and meiosis
 outline Mendelian laws of inheritance
 determine genotype and phenotype ratios
143
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 outline types of gene expression

 describe effects of environment on gene expression
 describe the importance of gene expression
 describe types of mutations
 Explain plant introduction as a breeding method.
 describe types of plant introduction
 discuss advantages and disadvantages of plant introduction
 explain selection as a method of plant breeding
 describe types of selection in plant breeding
 Discuss the advantages and disadvantages of plant selection. explain methods of hybridization
 describe advantages and disadvantages of hybridization
 describe hybrid seed production
 Compare hybrid seed and, traditional and commercial open-pollinated varieties (OPVs)
performances.
 Describe genetic engineering as a breeding method.
 Discuss advantages and disadvantages of genetically modified crops.

CHROMOSOME

THE NUCLEUS, CHROMOSOME AND GENE

So far we have done the functions of some important cell organelles. Now let us look closely at the
most important organelles that are involved in inheritance.

The nucleus

We mentioned that the nuclei contain genetic material which includes the chromosomes and genes.
These control all activity of an individual cell. The nucleus is surrounded by a nuclear envelope and
contains chromatin, one or more nucleoli and nucleoplasma.

Chromatin

Nuclear envelope

Nucleoplasma

Nucleolus

Nuclear pore

The nuclear envelope is composed of two membranes and is perforated by nuclear pores, to allow
exchange of substances between the nuclear and the cytoplasm. Nucleoplasma is a gel-like substance
which contains the chromatin and one or more nucleoli. The chromatin is composed mainly of coils of
the genetic material (DNA) bounded to proteins called histones. During cell division the chromatin
bodies condense to form shorter and thicker threads which are called chromosomes. The nucleolus is a
dense spherical body of variable size which is not enclosed in a unit membrane.

Chromosome

144
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

. A chromosome (from ancient Greek: χρωμόσωμα, chromosoma, chroma means colour, soma means
body) is a DNA molecule with part or all of the genetic material (genome) of an organism.
Chromosomes or chromatin are responsible for the transmission of hereditary information from
generation to generation. The chromatin is contained in the nucleus. Just before nuclear division, the
chromatin coils up into much more compact structures which are shorter and thicker and visible as
separate structures known as chromosomes. Chromosomes are normally visible under a light
microscope only when the cell is undergoing the metaphase of cell division. Before this happens,
every chromosome is copied once (S phase), and the copy is joined to the original by a centromere,
resulting in an X-shaped structure. The original chromosome and the copy are now called sister
chromatids. During metaphase, when a chromosome is in its most condensed state, the X-shape
structure is called a metaphase chromosome. In this highly condensed form chromosomes are easiest
to distinguish and study.
Chromosomes vary widely between different organisms. Some species such as certain bacteria, which
lack histones, also contain plasmids or other extrachromosomal DNA. These are circular structures in
the cytoplasm that contain cellular DNA and play a role in horizontal gene transfer.
DNA condensation of the duplicated chromosomes during cell division (mitosis or meiosis) results
either in a four-arm structure (pictured to the right) if the centromere is located in the middle of the
chromosome or a two-arm structure if the centromere is located near one of the ends.
Chromosomal recombination during meiosis and subsequent sexual reproduction play a significant
role in genetic diversity.
Chromosome number and homologous chromosomes
The number of chromosomes of different species is variable. The number has no specific significance
and does not indicate in any way the revolutionary advancement of the species. Somatic cells of all the
individual of the same species possess the same number of chromosomes. This number is called the
somatic or diploid number and is denoted by 2n.
The total chromosome complement is made up of two matching sets of chromosomes. These pairs of
chromosomes are referred to as homologous chromosomes. Humans have 23 pairs of homologous
chromosomes (46 total chromosomes). One set comes from the female egg (maternal chromosomes).
After fertilization the resulting cell or zygote has two sets of chromosomes.
Advantages of possessing two set of chromosomes is that genetic variation is increased and that if a
gene of one chromosome is faulty, the second chromosome may provide a normal back up.

The morphology of chromosome

A chromosome is made up of two chromatids joined by a region called centromere (figure 2.1). Inside
the chromosome they are two strands of DNA. DNA is made up of strands of segments called genes.

145
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2.4.3 Structure of the chromosome

Chromomere

Centromere

Figure 2.1: Structure of the chromosome

Depending upon the position of the centromere the following chromosome shapes can be
distinguished:

 Telocentric type- In these, the centromere is at the tip of the chromosome

 Acrocentric type- The centromere is near to one end

 Metacentric type- Centromere is in the middle

 Sub-Metacentric/ Subcentric- Centromere lies a little distance away from the middle

o Dicentric type - a chromosome with two centromeres

o Acentric type - a chromosome fragment without any centromere

1. Each chromosome is made up of two chromatids (chromosomal arms) which are joined to each
other at a small constricted region called the centromere. (Primary constriction). These sister
chromatids are conjoined twins the result of DNA replication.
2. The centromere helps the chromatids attach to the spindle fibres during cell division, it is also
concerned with the anaphase movement of the chromosomes, by which the spindle fibers pull the
chromatids to the two opposite poles by their contraction during anaphase.
3. Besides the primary constriction, in certain chromosomes there is a secondary constriction as well.
Because a small portion is pinched off from the chromosomal body; this portion is called a
'satellite' and the chromosome is called an SAT chromosome.
4. The two chromatids are made up of very thin chromatin fibres which are made up of 40% DNA
and 60% histone proteins

146
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

5. Each chromatin fibre consists of one DNA helix coiled around eight histone molecules like a loop;
such a complex is called nucleosome and resembles a bead on a string. These nucleosomes pack
tighter, during condensation required to get to metaphase.
6. The primary constriction cannot take up most stains, so during cell division this region is a gap in
staining.
7. Within the primary constriction there is a clear zone called Centromere.
8. The centromere with the DNA and histone proteins bound to them form a disc shaped structure
called kinetochore.
9. The chromonema is a word that means a chromatid in the early stage of condensation.

Chemical constituents of chromosomes

Chromosomes are composed of DNA and protein molecules (histones), with small amounts of RNA.
These three substances are responsible for storing and utilizing the vast amount of genetic information
and for transmitting the same from one cell generation to the other.

Genes

So far we have covered the structure and morphology of a chromosome. Now we want to look at the
gene which is a segment of a chromosome. A gene is the shortest segment of a chromosome that codes
for a particular characteristic. This means that the gene contains some information and therefore it can
replicate without losing this information. In 1944, Mr. Oswald Avery of the Rockefeller Institute in
New York produced evidence that DNA is the carrier of genetic information. This is some of the
evidence:

 DNA content is the same in all diploid cells of organism of the same species.
 Haploid cell (gametes) contains half the amount of DNA present in the somatic cells.
 DNA is capable of self duplication and that with high accuracy
 Gene is known to be stable; DNA is known to be the best stable compound in a cell.

Activity

1. Draw and label the typical plant cell

2. Write down major differences between the plant cell and the animal cell
3. Illustrate various shapes of the chromosomes

Summary
We mentioned that the cell is the unit structure in all living things. It is made up of the protoplasm
which contains all cell organelles (parts of the cell). The protoplasm is further differentiated into
the nucleus and the cytoplasm. The nucleus contains the genes and the chromosomes. Every
individual has a certain number of chromosomes and they are found in set of two for variation and
also the other set can provide a normal back up when one chromosome is at fault. Genes are
located in the chromosomes and are the ‘units of heredity’ and they control one or more characters
in an organism.

THE STRUCTURE OF DNA

The DNA in chromosomes forms the basis for the genetic code. It alone contains the language of life
that determines how a cell operates and all of your characteristics.
147
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

DNA is made of long chains of nucleotides. DNA has a DOUBLE-HELIX structure where 2 strands
of nucleotides join and twist around to form a spiral staircase or twisted ladder.

Each nucleotide has 3 parts:

1. a phosphate group - helps form the backbone of the DNA molecule

2. a sugar (deoxyribose) - acts as a glue,forms the backbone with phosphate

3. nitrogen bases - 4 types (guanine, cytosine, adenine, thymine)

o the base of one nucleotide forms a hydrogen bond with the base of another nucleotide, thus the
bases form the steps of the DNA ladder.
o the sequence of the bases makes up the genetic code

LAW OF COMPLEMENTARY BASE PAIRINGS:

adenine and thymine always bind to one another (A-T) and cytosine and guanine bind together (C-
G).

Thus the double helix DNA has the complementary bases hidden and protected by the intertwined
sugar-phosphate backbones.

DNA FUNCTION

1. Control of cell activities: to produce characteristics of an individual and species

2. Replication: in order for materials to be passed on from cell to cell, generation to generation,
and for growth, via meiosis and mitosis.
3. Undergo mutations: permanent changes passed onto offspring may advance the species via
evolution

DNA REPLICATION

148
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Why Replicate DNA?

DNA is the genetic material that defines every cell. Before a cell duplicates and is divided into
new daughter cells through either mitosis or meiosis, biomolecules and organelles must be copied to
be distributed among the cells. DNA, found within the nucleus, must be replicated in order to ensure
that each new cell receives the correct number of chromosomes. The process of DNA duplication is
called DNA replication. Replication follows several steps that involve multiple proteins called
replication enzymes and RNA. In eukaryotic cells, such as animal cells and plant cells, DNA
replication occurs in the S phase of interphase during the cell cycle. The process of DNA replication is
vital for cell growth, repair, and reproduction in organisms.

Step 1: Replication Fork Formation

Before DNA can be replicated, the double stranded molecule must be “unzipped” into two single
strands. DNA has four bases called adenine (A), thymine (T), cytosine (C) and guanine (G) that form
pairs between the two strands. Adenine only pairs with thymine and cytosine only binds with guanine.
In order to unwind DNA, these interactions between base pairs must be broken. This is performed by
an enzyme known as DNA helicase. DNA helicase disrupts the hydrogen bonding between base pairs
to separate the strands into a Y shape known as the replication fork. This area will be the template for
replication to begin.

DNA is directional in both strands, signified by a 5' and 3' end. This notation signifies which side
group is attached the DNA backbone. The 5' end has a phosphate (P) group attached, while the 3'
end has a hydroxyl (OH) group attached. This directionality is important for replication as it only
progresses in the 5' to 3' direction. However, the replication fork is bi-directional; one strand is
oriented in the 3' to 5' direction (leading strand) while the other is oriented 5' to 3' (lagging strand).
The two sides are therefore replicated with two different processes to accommodate the directional
difference.

Replication Begins

Step 2: Primer Binding

The leading strand is the simplest to replicate. Once the DNA strands have been separated, a short
piece of RNA called a primer binds to the 3' end of the strand. The primer always binds as the starting
point for replication. Primers are generated by the enzyme DNA primase.

Step 3: Elongation

Enzymes known as DNA polymerases are responsible creating the new strand by a process called
elongation. There are five different known types of DNA polymerases in bacteria and human cells. In
bacteria such as E. coli, polymerase III is the main replication enzyme, while polymerase I, II, IV and
V are responsible for error checking and repair. DNA polymerase III binds to the strand at the site of
the primer and begins adding new base pairs complementary to the strand during replication.
In eukaryotic cells, polymerases alpha, delta, and epsilon are the primary polymerases involved in
DNA replication. Because replication proceeds in the 5' to 3' direction on the leading strand, the newly
formed strand is continuous.

149
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The lagging strand begins replication by binding with multiple primers. Each primer is only several
bases apart. DNA polymerase then adds pieces of DNA, called Okazaki fragments, to the strand
between primers. This process of replication is discontinuous as the newly created fragments are
disjointed.

Step 4: Termination

Once both the continuous and discontinuous strands are formed, an enzyme
called exonuclease removes all RNA primers from the original strands. These primers are then
replaced with appropriate bases. Another exonuclease “proofreads” the newly formed DNA to check,
remove and replace any errors. Another enzyme called DNA ligase joins Okazaki fragments together
forming a single unified strand. The ends of the linear DNA present a problem as DNA polymerase
can only add nucleotides in the 5′ to 3′ direction. The ends of the parent strands consist of repeated
DNA sequences called telomeres. Telomeres act as protective caps at the end of chromosomes to
prevent nearby chromosomes from fusing. A special type of DNA polymerase enzyme
called telomerase catalyzes the synthesis of telomere sequences at the ends of the DNA. Once
completed, the parent strand and its complementary DNA strand coils into the familiar double
helix shape. In the end, replication produces two DNA molecules, each with one strand from the
parent molecule and one new strand.

Replication Enzymes

DNA replication would not occur without enzymes that catalyze various steps in the process. Enzymes
that participate in the eukaryotic DNA replication process include:

DNA helicase - unwinds and separates double stranded DNA as it moves along the DNA. It forms the
replication fork by breaking hydrogen bonds between nucleotide pairs in DNA.

DNA primase - a type of RNA polymerase that generates RNA primers. Primers are short RNA
molecules that act as templates for the starting point of DNA replication.

DNA polymerases - synthesize new DNA molecules by adding nucleotides to leading and lagging
DNA strands.

Topoisomerase or DNA Gyrase - unwinds and rewinds DNA strands to prevent the DNA from
becoming tangled or supercoiled.

Exonucleases - group of enzymes that remove nucleotide bases from the end of a DNA chain.

DNA ligase - joins DNA fragments together by forming phosphodiester bonds between nucleotides.

DNA Replication Summary

DNA replication is the production of identical DNA helices from a single double-stranded DNA
molecule. Each molecule consists of a strand from the original molecule and a newly formed strand.
Prior to replication, the DNA uncoils and strands separate. A replication fork is formed which serves
as a template for replication. Primers bind to the DNA and DNA polymerases add new nucleotide
sequences in the 5′ to 3′ direction. This addition is continuous in the leading strand and fragmented in

150
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

the lagging strand. Once elongation of the DNA strands is complete, the strands are checked for
errors, repairs are made, and telomere sequences are added to the ends of the DNA.

CHAPTER 22

PROTEIN SYNTHESIS

Protein Synthesis Summary

Protein synthesis is one of the most fundamental biological processes by which individual cells build
their specific proteins. Within the process are involved both DNA (deoxyribonucleic acid) and
different in their function ribonucleic acids (RNA). The process is initiated in the cell’s nucleus, where
specific enzymes unwind the needed section of DNA, which makes the DNA in this region accessible
and a RNA copy can be made. This RNA molecule then moves from the nucleus to the cell cytoplasm,
where the actual the process of protein synthesis take place.

What is protein synthesis – The details!

The synthesis of proteins takes two steps: transcription and translation. Transcription takes the
information encoded in DNA and encodes it into mRNA, which heads out of the cell’s nucleus and
into the cytoplasm. During translation, the mRNA works with a ribosome and tRNA to synthesize
proteins.

DNA TRANSCRIPTION RNA TRANSLATION PROTEIN

All cells function through their proteins. Protein function is defined by their molecular function ,
localization within cell and involvement in a particular biological process. All components of protein
function are defined by the exact composition, structure and conformation of the proteins, which is
encrypted within the DNA region (called locus) encoding that protein. With the process of protein
synthesis biological cells generate new proteins, which on the other hand is balanced by the loss of
cellular proteins via degradation or export.

Transcription is the first of overall two protein synthesis steps. During transcription, the information
encoded in the DNA is copied to a RNA molecule as one strand of the DNA double helix is used as a
template. The RNA molecule is sent to the cytoplasm, which helps to bring all components required
for the actual protein synthesis together – amino acids, transport RNAs, ribosomes, etc. In the
cytoplasm the protein polymers are actually “synthesized” through chemical reactions – that is why
the process is known as “protein synthesis” or even more precisely – “protein biosynthesis”.

The RNA copy of the protein genetic information encoded in DNA molecule is produced in the
nucleus and it is called messenger RNA (mRNA). Each mRNA encodes the information for a single
151
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

protein and is much smaller in size compared to the DNA molecule. This makes possible for mRNA
molecules to exit the nucleus through tiny openings called nuclear pores. Once it exits the nucleus and
enters the cytoplasm, the mRNA could interact with a cellular structure known as a ribosome, which
serves as the cell’s assembler within the process of protein synthesis. The ribosome consists of
proteins and ribosome RNA molecules (rRNA), which are organized in two subunits. The mRNA
initially binds to just one of the ribosome sub-units.

When the mRNA interacts with the big ribosome sub-unit, this triggers the approach of another RNA
molecule, called transfer RNA (tRNA). The tRNA molecule possess a specific sequence of 3-bases
(anti-codon), which hast to complement a corresponding sequence (codon) within the mRNA
sequence. When it finds it, it attaches to the mRNA, as the other end of the tRNA is “loaded” with an
amino acid. At this point arrives the other sub-unit of the ribosome and a complete structure is formed.
The first tRNA binds to a so called “start codon”, which is one and the same for all proteins. As the
complete ribosome structure is formed, another tRNA molecule approaches. The next tRNA differ
from the first one and is carrying another amino acid. Again, the tRNA must have an anti-codon that
matches complementary the second codon of the mRNA. The two amino acids carried by the first two
tRNAs are bind together with help from the ribosome and using cellular energy in the form of
adenosine triphosphate (ATP).

The above steps repeats until there are uncoupled codon sequences on the mRNA – thus the chain of
amino acids grows longer. Once the sequence of amino acids is successfully assembled in a protein,
the two ribosome sub-units separate from each other, to be joined again for later use.

The actual sequence of amino acids forms the so called primary structure of the proteins. Depending
on the exact composition and order of the amino acids in the protein sequence, the chain folds into a
three-dimensional shape. When this happens the protein is complete.

The process of protein synthesis takes place in multiple ribosomes simultaneous and all throughout the
cell cytoplasm. A living cell can synthesize hundreds of different proteins every single second.

GENE EXPRESSION
Gene expression is the process by which the instructions in our DNA are converted into a functional
product, such as a protein. OR is the process by which information from a gene is used in the
synthesis of a functional gene product. These products are often proteins, but in non-protein coding
genes such as transfer RNA (tRNA) or small nuclear RNA (snRNA) genes, the product is a
functional RNA.

 When the information stored in our DNA? is converted into instructions for
making proteins? or other molecules, it is called gene expression?.

 Gene expression is a tightly regulated process that allows a cell to respond to its changing
environment.

 It acts as both an on/off switch to control when proteins are made and also a volume control
that increases or decreases the amount of proteins made.

152
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 There are two key steps involved in making a protein, transcription and translation.

Transcription

 Transcription is when the DNA in a gene? is copied to produce an RNA? transcript

called messenger RNA? (mRNA).

 This is carried out by an enzyme? called RNA polymerase which uses available bases from
the nucleus? of the cell to form the mRNA.

 RNA is a chemical similar in structure and properties to DNA, but it only has a single strand
of bases? and instead of the base thymine ?(T), RNA has a base called uracil? (U).

Translation

 Translation occurs after the messenger RNA (mRNA) has carried the transcribed ‘message’
from the DNA to protein-making factories in the cell, called ribosomes?.

 The message carried by the mRNA is read by a carrier molecule called transfer RNA ?(tRNA).

 The mRNA is read three letters (a codon) at a time.

 Each codon specifies a particular amino acid?. For example, the three bases ‘GGU’ code for an
amino acid called glycine.

 As there are only 20 amino acids but 64 potential combinations of codon, more than one codon
can code for the same amino acid. For example, the codons ‘GGU’ and ‘GGC’ both code for
glycine.

 Each amino acid is attached specifically to its own tRNA molecule.

 When the mRNA sequence is read, each tRNA molecule delivers its amino acid to the
ribosome and binds temporarily to the corresponding codon on the mRNA molecule.

 Once the tRNA is bound, it releases its amino acid and the adjacent amino acids all join
together into a long chain called a polypeptide.

153
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 This process continues until a protein is formed.

 Proteins carry out most of the active functions of a cell.

At any given time, the amount of a particular protein in a cell reflects the balance between that
protein's synthetic and degradative biochemical pathways. On the synthetic side of this balance, recall
that protein production starts at transcription (DNA to RNA) and continues with translation (RNA to
protein). Thus, control of these processes plays a critical role in determining what proteins are present
in a cell and in what amounts. In addition, the way in which a cell processes its RNA transcripts and
newly made proteins also greatly influences protein levels.

EFFECTS OF ENVIRONMENT ON GENE EXPRESSION

The expression of genes in an organism can be influenced by the environment, including the external
world in which the organism is located or develops, as well as the organism's internal world, which
includes such factors as its hormones and metabolism. One major internal environmental influence
that affects gene expression is gender, as is the case with sex-influenced and sex-limited traits.
Similarly, drugs, chemicals, temperature, and light are among the external environmental factors that
can determine which genes are turned on and off, thereby influencing the way an organism develops
and functions. Scientists have long appreciated the role that environmental factors play in the
production of traits in animals. Environmental factors such as diet, temperature, oxygen levels,
humidity, light cycles, and the presence of mutagens can all impact which of an animal's genes are
expressed, which ultimately affects the animal's phenotype. For this reason, scientists who study the
genetics of model organisms usually seek to minimize environmental influence by maintaining
constant environmental conditions for the organisms under study. Nevertheless, even genetically

154
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

identical organisms exposed to controlled experimental conditions can have different phenotypes,
pointing to the power of subtle environmental differences on gene expression.

Temperature and Light

In addition to drugs and chemicals, temperature and light are external environmental factors that may
influence gene expression in certain organisms. For example, Himalayan rabbits carry the C gene,
which is required for the development of pigments in the fur, skin, and eyes, and whose expression is
regulated by temperature (Sturtevant, 1913). Specifically, the C gene is inactive above 35°C, and it is
maximally active from 15°C to 25°C. This temperature regulation of gene expression produces rabbits
with a distinctive coat coloring. In the warm, central parts of the rabbit's body, the gene is inactive,
and no pigments are produced, causing the fur color to be white Meanwhile, in the rabbit's extremities
(i.e., the ears, tip of the nose, and feet), where the temperature is much lower than 35°C, the C gene
actively produces pigment, making these parts of the animal black.

Light can also influence gene expression, as in the case of butterfly wing development and growth.

MUTATIONS
Is the permanent alteration of the nucleotide sequence of the genome of an organism, virus,
or extrachromosomal DNA or other genetic elements.

Mutations result from errors during DNA replication or other types of damage to DNA (such as may
be caused by exposure to radiation or carcinogens), which then may undergo error-prone repair
(especially microhomology-mediated end joining), or cause an error during other forms of repair, or
else may cause an error during replication (translesion synthesis). Mutations may also result
from insertion or deletion of segments of DNA due to mobile genetic [Link] may or may
not produce discernible changes in the observable characteristics (phenotype) of an organism.
Mutations play a part in both normal and abnormal biological processes including: evolution, cancer,
and the development of the immune system, including junctional diversity.

Mutation can result in many different types of change in sequences. Mutations in genes can either
have no effect, alter the product of a gene, or prevent the gene from functioning properly or
completely.

What kinds of gene mutations are possible?

The DNA sequence of a gene can be altered in a number of ways. Gene mutations have varying
effects on health, depending on where they occur and whether they alter the function of essential
proteins. The types of mutations include:

Missense mutation

This type of mutation is a change in one DNA base pair that results in the substitution of one amino
acid for another in the protein made by a gene.

Nonsense mutation

155
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

A nonsense mutation is also a change in one DNA base pair. Instead of substituting one amino acid for
another, however, the altered DNA sequence prematurely signals the cell to stop building a protein.
This type of mutation results in a shortened protein that may function improperly or not at all.

Insertion

An insertion changes the number of DNA bases in a gene by adding a piece of DNA. As a result, the
protein made by the gene may not function properly.

Deletion

A deletion changes the number of DNA bases by removing a piece of DNA. Small deletions may
remove one or a few base pairs within a gene, while larger deletions can remove an entire gene or
several neighboring genes. The deleted DNA may alter the function of the resulting protein(s).

Duplication

A duplication consists of a piece of DNA that is abnormally copied one or more times. This type of
mutation may alter the function of the resulting protein.

Frameshift mutation

This type of mutation occurs when the addition or loss of DNA bases changes a gene's reading frame.
A reading frame consists of groups of 3 bases that each code for one amino acid. A frameshift
mutation shifts the grouping of these bases and changes the code for amino acids. The resulting
protein is usually nonfunctional. Insertions, deletions, and duplications can all be frameshift
mutations.

Repeat expansion

Nucleotide repeats are short DNA sequences that are repeated a number of times in a row. For
example, a trinucleotide repeat is made up of 3-base-pair sequences, and a tetranucleotide repeat is
made up of 4-base-pair sequences. A repeat expansion is a mutation that increases the number of times
that the short DNA sequence is repeated. This type of mutation can cause the resulting protein to
function improperly.

GENETIC TERMS
o A gene is a sequence of nucleotide on the DNA strand which codes for a certain peptide chain. It’s
also referred as a unit of inheritance.
o An allele is an alternative form of the same gene responsible for determining, construction of
characteristics. [Link] allele for black skin color and an allele for white skin color.
o A dominant allele is an allele which influences the appearance of the phenotype presence of an
alternative allele. It suppose of expression of a recessive allele. It is represented by capital letters.
o A recessive allele which influence the appearance of the phenotype only in the presence of the other
identical allele. It will not express itself in the presence of the alternative allele of the same gene.
o A phenotype is the outward appearance of an organism or the external expression of a gene or
genotype e.g. black white etc.
o A genotype is the genetic constitution of an organism with respect to alleles under consideration e.g.
BB, Bb, bb.

156
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

o A locus is the position of an allele in the DNA molecule.

o Homozygous is a diploid condition in which the alleles at a given locus are identical e.g. BB or bb.
o Heterozygous is a diploid condition in which the alleles at a given locus are different e.g. Bb.
o A first filial generation (F1) these are off-springs produced by crossing parental genotypes of
organism.
o A second filial generation (F2) these are produced by crossing the parents from the F1 generation.

MENDELIAN LAWS OF INHERITANCE

Law of genes (the "First Law")

The Law of Segregation states that every individual organism contains two alleles for each trait, and
that these alleles segregate (separate) during meiosis such that each gamete contains only one of the
alleles. An offspring thus receives a pair of alleles for a trait by inheriting homologous
chromosomes from the parent organisms: one allele for each trait from each parent. tall & short pea
plants was:

Parents F1
Offspring
Genotype(s) TT x tt 100% Tt
Phenotype(s) tall x 100% tall
short
So, he takes two of the "F1" generation (which are tall) & crosses them. I would think that he is
figuring that he's gonna get all tall again (since tall is dominant). But no! Low & behold he gets some
short plants from this cross! His new batch of pea plants (the "F2" generation) is about 3/4 tall & 1/4
short.

We model this "Law of Segregation" every time. When you "split" the genotype letters & put one
above each column & one in front of each row, you have SEGREGATED the alleles for a specific
trait. In real life this happens during a process of cell division called "MEIOSIS". Meiosis leads to the
production of gametes (sex cells), which are either eggs or sperm. Sometimes the term
"GAMETOGENESIS" is used instead of meiosis.

157
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

You can see from the p-square that any time you cross two hybrids, 3 of the 4 boxes will produce an
organism with the dominant trait (in this example "TT", "Tt", & "Tt"), and 1 of the 4 boxes ends up
homozygous recessive, producing an organism with the recessive phenotype ("tt" in this example).

Law of Independent Assortment (the "Second Law")

The Law of Independent Assortment states that alleles for separate traits are passed independently of
one another i.e from parents to offspring. That is, the biological selection of an allele for one trait has
nothing to do with the selection of an allele for any other trait. Mendel found support for this law in
his dihybrid cross experiments. In his monohybrid crosses, an idealized 3:1 ratio between dominant
and recessive phenotypes resulted. In dihybrid crosses, however, he found a 9:3:3:1 ratios. This shows
that each of the two alleles is inherited independently from the other, with a 3:1 phenotypic ratio for
each.

The phenotypes of two independent traits show a 9:3:3:1 ratio in the F2 generation. Independent
assortment occurs in eukaryotic organisms during meiotic prophase I, and produces a gamete with a
mixture of the organism's chromosomes. The physical basis of the independent assortment of
chromosomes is the random orientation of each bivalent chromosome along the metaphase plate with
respect to the other bivalent chromosomes. Along with crossing over, independent assortment
increases genetic diversity by producing novel genetic combinations.

For example, height (tall or short), seed shape (round or wrinkled), pod color (green or yellow), etc.
Mendel noticed during all his work that the height of the plant and the shape of the seeds and the color
of the pods had no impact on one another. In other words, being tall didn't automatically mean the
plants had to have green pods, nor did green pods have to be filled only with wrinkled seeds,
the different traits seem to be inherited INDEPENDENTLY.

Please note my emphasis on the word "different". Nine times out of ten, in a question involving
two different traits, your answer will be "independent assortment". There is a big ugly punnet square
that illustrates this law so I guess we should take a look at it. It involves what's known as a "dihybrid
cross", meaning that the parents are hybrid for two different traits.

The genotypes of our parent pea plants will be:

RrGg x RrGg

where
"R" = dominant allele for round seeds
"r" = recessive allele for wrinkled seeds
"G" = dominant allele for green pods
"g" = recessive allele for yellow pods

Notice that we are dealing with two different traits: (1) seed texture (round or wrinkled) & (2) pod
color (green or yellow). Notice also that each parent is hybrid for each trait (one dominant & one
recessive allele for each trait).

We need to "split" the genotype letters & come up with the possible gametes for each parent. Keep in
mind that a gamete (sex cell) should get half as many total letters (alleles) as the parent and
158
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

only one of each letter. So each gamete should have one "are" and one "gee" for a total of two letters.
There are four possible letter combinations: RG, Rg, rG, and rg. These gametes are going "outside"
the p-square, above 4 columns & in front of 4 rows. We fill things in just like before --- "letters from
the left, letters from the top". When we finish each box gets four letters total (two "are's" & two
"gees").

This is what it looks like:

RG Rg rG rg
RG RRGG RRGg RrGG RrGg
round round round round
Rg RRGg RRgg RrGg Rrgg
round round round round
rG RrGG RrGg rrGG rrGr
round round wrinkled wrinkled
rg RrGg Rrgg rrGg rrgg
round round wrinkled wrinkled
The results from a dihybrid cross are always the same:
9/16 boxes (offspring) show dominant phenotype for both traits (round & green),
3/16 show dominant phenotype for first trait & recessive for second (round & yellow),
3/16 show recessive phenotype for first trait & dominant form for second (wrinkled & green), &
1/16 show recessive form of both traits (wrinkled & yellow).

So, as you can see from the results, a green pod can have round or wrinkled seeds, and the same is true
of a yellow pod. The different traits do not influence the inheritance of each other. They are inherited
INDEPENDENTLY.

TEST 1 BACK CROSS TECHNIQUE

─ An organism showing dominant characteristics can have two possible genotypes, A tall plant can
have either
homozygous or heterozygous tall. The phenotype will be the same but the genotype is different and is
determined bycrossing the plant with the recessive organism.
─ By crossing this organism with the unknown genotype with homozygous recessive it is possible to
determine the unknown or dominant characteristics genotype e.g. tall pea plant.
─ If half the off-springs are tall and the other half is short this implies that the unknown genotype is
heterozygous tall

CODOMINANCE
─ It is a condition in which two or more alleles fail to show complete dominance or recessiveness to
each other. This causes the alleles to show equally their effects on the phenotype.
─ This is due to the failure of one of the alleles to be dominant in the heterozygous condition. Co
dominance is found in both plants and animals.
─ The heterozygous has a phenotype which is intermediate between homozygous dominant and
recessive condition produced by crossing pure breeding black and splashed white fowls.
─ A cross between pure breeds of red and white cows produces an intermediate color called ROAN.
─ The presence of the black plumage is the result of possession of an allele for black pigment melanin.
Splashed white fowls also lack this melanin. Heterozygous show a partial development of this melanin
which produces a blue sheen in the plumage.
159
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 23

BREEDING

Definition of Plant Breeding

Plant breeding is an applied branch of Botany, which deals with improvement of agricultural crops.
This branch of agricultural science has contributed maximum to the increase in food production all
over the world and therefore, now a day it assuming ever increasing importance in field of agriculture
in every country. Riley, 1978 defined plant breeding as a technology of developing superior crop
plants/ varieties for various purpose. Frankel, 1958 defined plant breeding as the genetic adjustment of
plants to the service man. Plant breeding is a branch of biology concerned with changing the genotype
of plant so that they become more useful. Plant breeding is the science, art, and business of
improving plants for human benefit (Bernardo, 2002;

Methods of breeding – introduction and acclimatization

The following are the methods of breeding autogamous plants.

1. Introduction 2. Selection a) Pure line selection b) Mass selection 3. Hybridization and selection i)
Inter varietal a) Pedigree Method b) Bulk Method. c) Single Seed Descent Method. d) Modified Bulk
method e) Mass - Pedigree Method. ii) Interspecific hybridization 4. Back cross method 5. Multiline
varieties 6. Population approach 7. Hybrids. 8. Mutation breeding 9. Polyploidy breeding 10.
innovative techniques

Plant introduction

Definition

Taking a genotype or a group of genotypes in to a new place or environment where they were not
grown previously. Thus introduction may involve new varieties of a crop already grown in that area, a
wild relative of the crop species or totally a new crop species for that area.

E.g. a) Introduction of lRRl rice varieties..

b) Introduction of sunflower wild species from Russia

c) Introduction of oilpalm in to Tamil Nadu.

Plant introduction may be of two types. 1. Primary Introduction and 2. Secondary Introduction

1. Primary Introduction

When the introduced crop or variety is well suited to the new environment, it is directly grown or
cultivated with out any alteration in the original genotype. This is known as primary introduction. E.g.
IR. 8, IR 20, IR 34, IR 50 rice varieties; oil palm varieties introduced from Malaysia and Mashuri rice
from Malaysia.

2. Secondary Introduction

160
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The introduced variety may be subjected to selection to isolate a superior variety or it may be used in
hybridization programme to transfer some useful traits. This is known as secondary Introduction.E.g.
In soybean EC 39821 introduced from Taiwan is subjected to selection and variety Co 1 was
developed. In rice ASD 4 is crossed with IR 20 to get Co 44 which is suited for late planting.

Objectives of Plant Introduction

 To introduce new plant species there by creating ways to build up new industries.E.g. Oil
 palm
 To introduce high yielding varieties to increase food production. E.g. Rice and wheat.
 To enrich the germplasm collection. E.g. Sorghum, Groundnut.
 To get new sources of resistance against both biotic and abiotic stresses.

E.g. NCAC accessions to have rust resistance in groundnut. Dasal rice variety for saline
[Link] value – ornamentals are introduced for aesthetic value.

Plant Introduction Agencies

At International level International Board of Plant Genetic Resources (IBPGR) with head quarters at
Rome, Italy is responsible for plant introduction between countries.

Procedure for plant Introduction

The scientist / University will submit the requirement to NBPGR. If the introduction is to be from
other countries, NBPGR will address IBPGR for effecting supply. The IBPGR will assign collect the
material from the source and quarantine them, pack them issue phytosanitary certificate suitably based
on the material and send it to NBPGR. The NBPGR will assign number for the material, keep part of
the seed for germplasm and send the rest to the scientist.

Functions of NBPGR

1. Introduction maintenance and distribution of germplasm

2. Provide information about the germplasm through regular publications.

3. Conduct training courses to the scientist with regard to introduction and maintenance of
germplasm.

4. Conduct exploratory surveys for the collection of germplasm.

5. To set up Natural gene sanctuaries.

Merits of plant introduction.

1. It provides new crop varieties, which are high yielding and can be used directly

2. It provides new plant species.

3. Provides parent materials for genetic improvement of economic crops.

161
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

4. Enriching the existing germplasm and increasing the variability.

5. Introduction may protect certain plant species in to newer area will save them from diseases.

E.g. Coffee and Rubber.

Demerits

1. Introduction of new weed unknowingly.E.g. Argemone mexicana, Eichornia and Parthenium

2. Introduction of new diseases: Late blight of potato from Europe and Bunchy top of banana from Sri
Lanka

3. New pests: Potato tuber moth came from Italy

4. Ornamentals becoming weeds: Lantana camara

5. Introduction may cause ecological imbalance [Link].

Acclimatization

When superior cultivars from neighbouring or distant regions are introduced in a new area, they
generally fail initially to produce a phenotypic expression similar to that in their place of origin.

But later on they pickup and give optimal phenotypic performance, in other words they become
acclimatized to the new ecological sphere. Thus acclimatization is the ability of crop variety to
become adapted to new climatic and edaphic conditions.

The process of acclimatization follows an increase in the frequency of those genotypes that are better
adapted to the new environment. The success of acclimatization depends upon two factors

i) Place effect

ii) Selection of new genotypes.

Selection, Mass selection, pure line selection and Johannson’s pure line theory, genetic basis.

Selection in Self-Pollinated Crops

To get successful results by selection there are two pre-requisites.

a) Variation must be present in the population.

b) The variation must be heritable.

Pureline theory

A pure line is the progeny of a single self fertilized homozygous plant. The concept of pureline was
proposed by Johannsen on the basis of his studies with beans (Phaseolus vulgaris) variety called

162
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Princess. He obtained the seeds from the market and observed that the lot consisted of a mixture of
larger as well as smaller size seeds.

Thus there was variation in seed size. Johannsen selected seeds of different sizes and grown them
individually. Progenies of larger seeds produced larger seeds and progenies from smaller seeds
produced small seeds only. This clearly showed that there is variation in seed size in the commercial
lot and it has a genetic basis. He studied nineteen lines al together. He concluded that the market lot of
the beans is a mixture of purelines. He also concluded whatever variation observed with in a pureline
is due to environment only. Confirmatory evidence was obtained in three ways. In line 13 which is
having 450 mg seed wt he divided the seeds on weight basis. He divided the line into seeds having
200, 300, 400 and 500 mg weights and studied the progenies. Ultimately he got lines having weight
ranging from 458 to 475. Thus the variation observed is purely due to environment.

The second evidence was that selection with in a pureline is ineffective. From a pureline having 840
mg selection was made for large as well as small seeds. After six generations of selection the line for
large seed as well as for small seed gave progenies having 680-690 mg.

Thus it was proved that selection within a pureline is ineffective. In third evidence when parent -
offspring regression was worked in line thirteen. It worked to zero indicating that variation observed is
non heritable and it is due to environment only.

Origin of variation in pure lines

1. Mechanical mixtures.

2. Natural hybridization.

3. Chromosomal aberrations.

4. Natural mutation or spontaneous mutation.

5. Environmental factors.

Effect of self-pollination on genotype

Self-pollination increases homozygosity with a corresponding decrease in heterozygosity.

For example an individual heterozygous for a single gene Aa is self pollinated in successive
generations, every generation of selfing will reduce the frequency of heterozygote Aa to 50 percent of
that in the previous generation. There is a corresponding increase in homozygote AA and aa. As a
result, after 10 generations of selfing virtually all the plant in the population will be homozygous AA
and aa.

Genetic advance under selection

Normally selection is practiced based on the phenotype of the individual plant. The phenotype in turn
is the result of joint action of genotype and environment i.e., VP=Vg +VE Where P= phenotype; G =
genotype; E = Environment

The genetic advance is calculated by the following formula.

163
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Genetic advance (GS) = (K) (H) (SD P) or GS = (K) (VP) ½ (Vg / Vp),

Where GS is the genetic advance under selection, K is the selection differential, SD P is the
phenotypic standard deviation of base population and H is the heritability of the character under
selection. The estimates of GS have the same unit as that of the mean.

Pureline Selection

A large number of plants are selected from a self pollinated crop. The selected plants are harvested
individually. The selected individual plants are grown in individual rows and evaluated and best
progeny is selected, yield tested and released as a variety.

Characteristics of purelines

1. All plants within a pure line have the same genotype.

2. The variation with in a pureline is environmental and non heritable.

3. Purelines become genetically variable with time due to natural hybridization, mutation and
mechanical mixtures.

General steps for making a pureline selection

First Season: From the base population select best looking plants having the desirable characters.
Harvest them on single plant basis.

Second Season: The selected single plants are grown in progeny rows and estimate the performance.
Reject unwanted progenies.

Third Season: Repeat the process of second season.

Fourth Season: Grow the selected single plants in replicated preliminary yield trial along with
suitable check or controI variety.

Fifth Season: Conduct regular comparative yield trial along with check variety and select the best
culture.

Sixth Season: Conduct multilocation trial in different research stations along with local check

Seventh Season: Conduct Adaptive Research Trial in farmer's field. Fix the best yielder and release it
as a variety thro' Variety Release committee.

Advantage of pureline selection.

1. Achieves maximum possible improvement over the original variety.

2. Extremely uniform in appearance.

3. Because of the uniformity, a variety is easily identified and seed certification is easy.

164
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Disadvantages

1. It does not have wide adaptability because improvement is made only in the local variety.

2. Time required for developing a variety is more when compared to mass selection.

3. Depending on the genetic variability present in the base population only the improvement is made.
If there is no genetic variability improvement cannot be made.

4. Breeder has to spend more time compared to mass selection.

Mass Selection

Here a large number of plants having similar phenotype are selected and their seeds are mixed
together to constitute a new variety. Thus the population obtained-from selected plants will be more
uniform than the original population. However they are genotypically different.

Steps

First season

From the base population select phenotypically similar plants, which may be 200 2000.

Harvest the selected plants as a bulk.

Second season

The bulk seed is divided into smaller lots and grown in preliminary yield trial along with control
variety. Dissimilar phenotypes are rejected. Higher yielding plots are selected.

Third to Sixth Season

With the selected lots conduct yield trials along with appropriate check or control. Select

the best one and release it as a variety.

Merits of Mass Selection

1. Varieties developed will be having more adaptability since each plant is genotypicaly not similar.
They have buffering action against abnormal environment.

2. Time taken for release of a variety is less.

3. The genetic variability present in the original population is maintained.

Demerits

1. Compared to pure line variety they may not be uniform.

165
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2. In the absence of progeny test we are not sure whether the superiority of selected plant is due to
environment or genotype.

3. May not be as uniform as that of a pureline variety and certification is difficult.

Comparison between pure line and mass selections

Pureline selection Mass selection

The new variety is a pureline The new variety is a mixture of purelines.
The new variety is highly uniform In fact, the variation within a pureline variety
is purely environmental.
The selected plants are subjected to progeny Progeny test is generally not carried out
test
The variety is generally the best pureline The variety is inferior to the best pureline
present in the original population. because most of the purelines included in
pure line selection will be inferior
Maize hybrid production
- They are three types of maize hybrids in Zimbabwe,
1. Single hybrid
2. Double hybrid
3. Three way hybrid
─ Two different types of maize plants with desired qualities are planted on same field.
─ The field must be isolated to prevent pollination from unknown sources.
─ One type of plant is allowed to tassel and produce pollen and this called the male or pollen plant.
─ The second type, the female or cob parent is only allowed to silk and its tassels are removed.
─ -Detasselling is done to prevent self-pollination.
─ The two types of plants then cross pollinate to produce a hybrid.
Production of single hybrid maize cultivars
-pollinating plants for several generations.
cross pollinated to get a single hybrid maize variety.
Inbred line A × Inbred line B
(Seed/cob parent) (Pollen parent)

Single hybrid cultivar (AB)

Production or double hybrid maize cultivars

Single hybrid (AB) × Singled hybrid (CD)

(Seed /cob parent) (Pollen parent)

Double hybrid cross

Production of 3 way hybrid maize cultivars

Single hybrid cross × Inbred line

(Seed /cob parent) (Pollen parent)
Three way hybrid
Advantages of using hybrid seed
1. High yield are obtain because parents with good qualities are used to produce hybrid seed.

166
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2. Resistance to pests and diseases.

3. Good adaptation to the local Environment condition.
4. The quality of yield is high.
Disadvantages of using hybrid seed
1. Only the 1st generation (F1) shows hybrid vigor.
2. They are expensive to produce
Principles of Hybrids
• Maize is normally cross-pollinated, and this maintains the plant vigour
• The principle of maize hybrids is based on the concept of hybrid vigour – achieved by crossing two
unrelated and distinct parents, giving higher yields than the parents.
• The parents are normally developed by inbreeding maize to develop uniform but less vigorous inbred
lines, and then crossing these to re-invigorate the hybrid plant
• Pollination of the maize must therefore be controlled
• For developing the inbred parents, the pollen from the plant is applied to the silks of the same plant
• For creating the hybrid, the pollen from the male must be applied to the silks of the female

GENETIC ENGINEERING

Genetic engineering involves the manipulation of genes within a species, and may also involve the
transfer of genes—and thus the characteristics governed by those genes—from one species to another.
Genetic engineering goals in plants include flower color and in agriculture include improving crop
production and introducing new traits, such as enhanced nutrients, temperature resistance, or the
ability to grow in saltier soils. Certain characteristics may allow a genetically modified (GM) crop to
be grown, harvested, or shipped at lower cost or with less damage and may allow the crop to be grown
using less pesticide. In some cases, GM crops may yield entirely new products that are not normally
derived from traditional crops—such as medicines
What Steps Are Usually Involved in Genetically Modifying a Crop?
How do we transfer genes among species? One way that scientists genetically engineer crops
is with the use of bacterial plasmids discussed earlier. In 1907, scientists found that a common plant
tumor called crown gall was caused by the invasion of a bacterium
called Agrobacterium tumefaciens (Smith and Townsend 1907). In 1977, scientists discovered that the
crown gall tumor was caused when plasmids from the bacteria were taken into the DNA of the host
plant’s cells and expressed (Drummond 1979). By 1983, researchers had modified the plasmids to
remove their ability to induce tumors but left them with the ability to incorporate themselves into plant
cell DNA, thereby creating a strand of DNA that could be used as a transfer plasmid (Barton and Brill
1983). To transfer genes between species, biotechnologists can attach the transgene—along with other
necessary genes, including a promoter that induces gene expression in plant cells—to a transfer
plasmid. The most common promoter used today is from the cauliflower mosaic virus. Marker genes,
such as those for antibiotic resistance, are then used to enable direct selection of the transferred
genetic material. Despite its original popularity, use of antibiotic resistance as a marker is now being
phased out. Europe has outlawed the use of such marker genes in commercial crops, so now it is more
common that, even with antibiotic resistance genes, several techniques are subsequently used to
eliminate marker genes before commercialization.
Transfer plasmid is constructed in a test tube and then reinserted into the A. tumefaciens
bacteria, which are then inserted by syringe or enabled to invade the wounded host
cells. Once inside their host, they release plasmids which the host cell’s DNA takes in to be
expressed as if it were part of the host’s normal genetic makeup. In fact, it is at this point
that they become a part of the plant’s genetic makeup. The new DNA cannot be extracted

167
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

independently of the host DNA and will be passed from generation to generation. This method is
currently considered preferable to the gene gun method (see next paragraph) because fewer copies of
the gene are inserted into a single cell and more cells survive.
Another bioengineering technique does not make use of plasmids. Instead, the transgene copies and
associated genes are painted onto microscopic metal particles and shot into thenucleus of the host cell
with a “gene gun.” Once the metal particles enter the plant cell nucleus, the transgenic DNA washes
off and inserts itself into the host DNA. The host plant cells in these transformation methods are
grown as tissue cul-
Some of the advantages of GM foods:
Genetically modified organisms (GMOs) are a form of scientific farming where chemicals are pumped
to crops to increase product sizes and yield. Although this method is highly debated, it has become
increasingly common in everyday foods. However, just like any other process, it also has two sides.
Here are the advantages and disadvantages of creating genetically modified crops.
What Are the Advantages of GMOs?
1. It allows for more profit.
GMOs are an effective way to provide farmers a larger profit, while making them spend less time on
resources.
2. It introduces the knowledge of genetic alterations.
This is done through mapping genetic material for GMO crops. This way, we would get the ability to
enhance crop genes and make them more beneficial for human production and consumption. Plants
can be engineered to resist temperature or produce higher yields, which is good for regions where
climate limits productivity.
3. It is economically efficient.
Because GMOs are designed to resist pests, there will be no need for pesticides to be used, which
means more savings.
4. It is known to decrease food prices.
Advanced crops and lower costs can lead to cheaper food. This will certainly help families who cannot
afford to buy their needed supply for everyday consumption, so starvation will be prevented.
5. It adds more nutritional value to crops.
The GMO method can put in added nutritional value to crops that lack necessary vitamins and
minerals. Considering that there are places in the world relying on rice or corn as their daily staple,
plant genes may be added to these crops to increase their nutritional value. This would help
malnourished populations receive more nutrients from their diet.
6. Its products are found to be safe.
The precise evaluation and testing of GMOs crops and other products means they are safe for human
consumption. In fact, research shows that they are safer compared with traditional crops
There is a need to produce inexpensive, safe and nutritious foods to help feed the world’s growing
population. Genetic modification may provide:
 Better quality food.
 Higher nutritional yields.
 Inexpensive and nutritious food, like carrots with more antioxidants.
 Foods with a greater shelf life, like tomatoes that taste better and last longer.
 Food with medicinal benefits, such as edible vaccines - for example, bananas with bacterial or
rotavirus antigens.
 Crops and produce that require less chemical application, such as herbicide resistant canola.

Some of the disadvantages of GM foods:

1. It can be dangerous to other insects that are important to our ecosystem.
GMOs are believed to be dangerous to some insects because new crop genes can be deadly to them.

168
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

This is worth noting when it comes to certain insects, such as butterflies, that are not actually
dangerous to crops.
2. It sparks concerns on changing the field of agriculture.
The process of making GMOs includes adding new genetic material into an organism’s genome. In
agricultural ecology, this means introducing new genes in the genome of crops like corn. Research on
the effects of cultivation of GM crops in a large scale has sparked various concerns, specifically those
ideas on ecosystems with GMO strains. As proven by certain studies, GMO strains have the potential
to change agriculture.
3. It can damage the environment.
Genetically modified crops can cause a threat to the environment due to the fact that they are not a
natural way to plant and cultivate plants.
4. It causes unwanted residual effects.
A genetically modified plant can leave unwanted residual substances that can remain in the soil for
extended periods of time. Agricultural regulators were alerted by research that strains from GM crops
would remain in the soil for years after the crops were removed. Its data even reported that despite the
absence of these plants, the strain persisted for up to more than 5 years.
5. It can create more weeds.
Take note that engineered crops can act as mediators in transferring genes to wild plants, which can
create more weeds. To keep these new weeds under control, scientists then invented new herbicides
that were not necessary for non-GMO weeds. These chemicals are also toxic to various mammals and
amphibians, who are feeding on GMO crops. Tests even show that the uptake of these herbicides also
has toxic consequences on aquatic ecosystems.
6. It threatens crop diversity.
There is opposition to introducing GM genes on genetic diversity because these genes can spread to
other organic farm crops and threaten crop diversity in agriculture. And if crop diversity decreases, it
will have a direct impact on our entire ecosystem and would affect the population dynamics of other
organisms. The chance that a single genetically modified crop strain could pollinate an already
existing non-GM crop is unlikely and unpredictable, and there are many conditions that must be met
for cross pollination to occur. However, when a large scale plantation releases a GM strain during
pollination, this risk increases, where the cross pollination to non-GM plants could create a hybrid
strain. This means there is a greater possibility of ecological novelty or new artificial strains that are
being introduced into the environment that could potentially reduce biodiversity through competition.
7. It has trade issues.
In other countries and regions in the world, there may be problems regarding trade matters, such as
tariff and quota.
Food regulatory authorities require that GM foods receive individual pre-market safety assessments.
Also, the principle of ‘substantial equivalence’ is used. This means that an existing food is compared
with its genetically modified counterpart to find any differences between the existing food and the
new product. The assessment investigates:
 Toxicity (using similar methods to those used for conventional foods).
 Tendency to provoke any allergic reaction.
 Stability of the inserted gene.
 Whether there is any nutritional deficit or change in the GM food.
 Any other unintended effects of the gene insertion.
Conventional Breeding Genetic Engineering
 Limited to exchanges between the same  Allows the direct transfer of one or just
or very closely related species a few genes, between either closely or
 Little or no guarantee of any particular distantly related organisms
gene combination from the million of  Crop improvement can be achieved in a

169
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

crosses generated shorter time compared to conventional

 Undesirable genes can be transferred breeding
along with desirable genes  Allows plants to be modified by
 Takes a long time to achieve desired removing or switching off particular
results genes

CHAPTER 24

WEEDS AND WEED CONTROL

 outline the socio-economic importance of weeds

 identify weeds
 classify weeds
 describe mechanisms that make weeds persistent
 explain the mechanisms of crop-weed competition
 describe safe handling of agro- chemicals.
 outline safe storage procedures for agro-chemicals.
 outline safe disposal of agro-chemicals.
 outline the importance of weed management.
 determine effective timing of weeding.
 describe the methods of weed management.
 evaluate weed management methods.
Introduction
Weeds are of economic importance in crop production since they can lead to yield losses in the field,
and down grading of the crop at marketing. They have their own defensive characteristics. To be able
to control weeds one needs to know the behaviour of weeds and then select the ways which can easily
counter their development. This chapter will cover the characteristics of weeds and weed control.

Definition
A weed is a plant that grows where it is not wanted. This definition includes volunteer plants. A weed
can be broad leaved or narrow leaved, annual or perennial.

Special adaptations
Special adaptations which make weeds special competitors.
Seed production
Most weeds multiply and disseminate through seed production. Weed seed can survive in very high
temperatures, drought and frost. Life cycles of weeds can be as short as 6 weeks such that there can be
several generations per season. Most weeds produce a large number of seeds per flowering period e.g.
shamva grass 10000 seeds / plant, Rapoko grass 100000/ plant , Mexican marigold -59000/ plant.
Weeds have very efficient reproductive and disseminative mechanisms which make them strong
competitors with arable crops.

Weed seed dormancy

170
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

A weed may mature and shed thousands of seeds under favourable conditions for germination. Only a
fraction germinates in the first year. Much of the seed will remain dormant and germinate in
succeeding years.
SOCIO-ECONOMIC IMPORTANCE OF WEEDS
1. Reduction in crop yield through:
- Physical Interaction (Allelospoly: competition for growth resources including water, light,
nutrient, air, space.
- Chemical interaction (Allelopathy)

2. Reduction in crop quality through

- direct contamination of cultivated rice and maize grain by wild rice (Oryza longistaminata) and itch
grass (Rottboellia cochinchinensis) respectively.;
- contamination of forage, silage or pasture crop .by C. rotundus seeds ,
- reduction in Sugarcane juice quality by the presence of sida.
- Contamination of cotton lint by dried weed fragments
- Damage of underground tuber of yam and cassava through piercing of Spear grass
rhizomes
3. Interference with field operations (harvest,pesticideapplication,etc.)
4. Some are poisionous to grazing animals e.g. Euphorbia heterophylla, Halogeton glomeratus
contain high oxalate content, it can kill livestock when eaten in dry season.
5. Some are harmful to grazing animals e.g. Amaranthus spinosus, Acanthospermum hispidus
6. increase cost of production; high cost of labour and equipment during harvesting.
7. Presence of weeds can impede water flow in irrigation canals
8. Weeds present in lakes and reservoirs can increase loss of water by evapotranspiration
9. Reduction in quality of pasture land; it reduces the carrying capacity of grazing lands and
pastures through their physical presence and weediness
10. Reduction in quality of animal products;it affects the palatability of pastures, hay, silage etc.
protein content in alfalfa wild garlic (Alliums spp) when eaten by cattle spoils the meat and the
milk.
11. Serve as alternate hosts for many plant diseases and animal pests e.g. insects, rodents, birds.
Cyperus rotundus serve as alternate to nematodes and athropods
12. Impose limitation to the farm size of a farmer
13. Can serve as sources of fire hazards
171
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

14. Reduce erosion problem through the production of protective cover

15. Help in nutrient recycling through decay of vegetative part.
16. Food/vegetables for humans e.g. leaves of Talinum triangulare, and tubers of Colocasia
esculentus .
17. Serve as hosts and nectar for beneficial insects
18. Beautification of the landscape e.g. Cynodon dactylon
19. Feed for livestock and wildlife and aquatic organisms in form of hay, silage and forage / pasture,
fruit seeds and branches and whole plant.
20. source of pesticides e.g. Chrysanthemum cinerariifolium
21. Source of genetic material for useful traits in crop improvement.
22. Medicinal use e.g neem ( Azadirachta indica), Ageratum conyzoides
23. Some serve as trap crop for parasitic weeds.
24. Habitat for wildlife and plant species hence biodiversity conservation.
25. Major role in carbon recycling through carbon sequestration. Field of exposed soil always suffers
a net loss in organic matter and releases carbon dioxide, while a field covered with crops and/or
weeds takes up carbon dioxide. This concept of carbon sequestration is an added advantage of
sustainable and organic farming.

Weed classification
Weeds can be classified based on
(1) Life cycle or history (Ontogeny)
Annual, Ephemeral, Perennial and Biennials weeds

(2) Habitat:
(a) Upland (terrestial) weeds or dry land weeds (Agrestal /Weeds of arable or cultivated crops, and
Ruderal weeds /weeds of disturbed non- cropped area such as rubbish heaps, landfills, paths, roads,
compost heaps
(b)Aquatic weeds (Submerged aquatic, Floating aquatic, Emergent aquatic weeds
(3) Growth habit:
Free living (autotrophic) weeds
ii Parasitic plants(Root parasitic weeds or obligate parasite, Stem parasitic weeds , Hemi parasitic
weeds, Total parasites Floating aquatic Emergent aquatic weeds
(4) Degree of undesirability: ease and difficulty in controlling weeds.
(5) Morphology :
[Link] e.g. Woody Stem e.g Azadirachta indica,
ii. Semi Woody weeds- e.g Chromolaena odorata, Sida acuta.
iiiHerbaceous weeds: e.g Ageratum conyzoides, Talinum triangulare,
b. Leaf Type : narrow leaf: grass like(ii) Broad leaf weeds (Dicotyledons):, Sedges; e.g. Cyperus
rotundus, C. esculentus, Mariscus alternifolius

172
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

(6) Scientific classification (Binomial nomenclture): based on their taonomy (family,, genera and
specific epithet
(7) Ecological affinities

 dryland weeds,
 gardenland weeds and
 wetland weeds
(8)Origin:
- native or introduced.

Weed control
Weed control should be done within the first 8 weeks of plant establishment (referred to as the critical
period). Weeds are easy to control in the early stages.

Techniques used in weed control

1. Hand hoeing
Hoes are used to remove or dislodge the weeds resulting in death.
Advantages of hand hoeing
1. No chemical damage or toxicity to the crop.
2. There is enough aeration to the soil.
3. Organic matter is also added to the soil.
4. Cut weeds can act as a mulch to conserve moisture

Disadvantages
1. There is high labour demand during the process of weeding.
2. Hand hoeing is very difficult in wet weather.

Mechanical cultivation
Ox drawn cultivators or tractor drawn implements can be used also to dislodge the weeds.
Herbicide use
Stage and method of application of herbicides
1. Pre-planting herbicides – Herbicides are applied before planting and incorporated into the soil.
Preplanting herbicides are applied 2-3 weeks before planting. If applied earlier they lose efficiency
due to volatilization, sunlight destruction or photodecomposition. An example is Trif in cotton.
2. Pre- emergence herbicides – Herbicide is applied after planting before the emergence of the crop.
Should be applied before the emergence of weeds (3-4 day of planting). Delay in application result
in poor weed control and crop damage. An example is Atrazine in maize.
3. Post emergence herbicide – Is a herbicide applied after the crop has emerged over the top or
directed to affect only weeds under the crop. Application late in the season is referred to as lay-by
treatment. Post emergence herbicide may have contact action or translocated therefore should be
applied after weed emergence. Soil acting post emergence herbicides are applied before weed
emergence.

Selective herbicides
173
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Selective herbicides retard or kill the weeds while the crop is tolerant. The herbicide is selective to a
particular crops within certain limits. Over application of such herbicides can affect the crop.
Basics of selectivity of herbicides
With herbicides, several mechanisms are involved in the selectivity to crops. Selection depends on one
or more of the following:
1- Genetic vigour of the crop
2- High crop seed viability and low weed seed viability under herbicides.
3- Physiological resistance or inherent crop tolerance with the herbicides
4- Application of herbicide to avoid excessive contact with the crop.
Mechanisms can be divided into physiological and non physiological mechanisms.

Non physiological mechanisms selectivity (positional selectivity)

a. Positional selectivity depends on the position of the growing point of the crop and the weed. Broad
leaved weeds have well exposed meristematic tissue while the meristem of grass is intercalary.
Each node in grass has meristematic tissue protected by leaf tissue. Herbicides affect growing
point and discriminate between the crop and broad leaved weeds which are killed by virtue of their
exposed apical meristems.
b. Leaf angle -Broad leaved weeds have flat leaf surface areas therefore intercept more herbicide
than grasses or cereals which have upright leaves.
c. Surface nature- Hairiness, waxiness of the leaf affects the effect of herbicide by preventing the
herbicide from adhering to the leaf therefore protecting the crop from the herbicide.
d. Different behaviour and movement of herbicide in the soil.

Depth control
Depth control is the principle on with pre-emergence herbicide work. Selectivity depends on the depth
of the crop and weed seed in relation to the depth to which the herbicide would penetrate. Crop seed is
planted deep and herbicide with low leachability is prayed and light irrigation is applied. Herbicide
stops weed germination in superficial layers of the soil.
Fig 1. Depth control

Herbicide
Crop seed protected by depth, seal
Therefore no contact with the
herbicide
In this case weed seed which germinate from a depth e.g upright star bar, salt weed are difficult to
control because pre- emergence herbicide do not penetrate to that depth. Examples of pre-emergence
herbicide which work by depth control are: Metachlor (dual) , Cotogard , Sencor , Alachor ( lasso)
,Planavin ,Trifluralin, Igran.

Depth control in orchards

Deep rooted crops such as fruit trees are depth protected by using a low solubility herbicide which is
highly adsorbed by soil colloids and will form a layer which will prevent weed growth in that layer.

Fig 2. Diagramatic Illustration of Depth control in orchards

174
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Herbicide layer, low solubility and

high adsorption, e.g. Diuron.

Weed seed will not germinate

Selective and non selective herbicides can be directly applied to hit weeds and avoid excessive
contact with the crop using flat jet nozzles or shielded sprayers. The crop must be taller than the weeds
within the rows.

Physiological selectivity mechanisms

Differential absorption of herbicides
Weeds with certain morphological and anatomical characteristics e.g. permeable cuticles, and large
stomatal apertures absorb folia sprays, thus the weed absorb more herbicide than the crop and is
controlled.

Differential Translocation of herbicide

Differences in the ability of translocation of herbicide in the phloem and xylem between the weed and
the crop forms the basis of selectivity.

Enzymatic activation
In some plants enzymes are capable of converting non toxic compounds into toxic compounds as is
the case with many weeds while monocots are unable to do so e.g m.c.p.a.
Persistence of herbicides in the soil
Persistence is the period a herbicide remains active in the soil to affect a subsequent crop. Persistence
determines the period of weed control achieved. A herbicide should not persist in a subsequent crop.
Herbicides like atrazine are generally used in maize. It is selective and cheap but persists for a long
period. It affects any broad leaf crops which follow the maize. Persistence affects rotations.

Factors that affect persistence of herbicides

Physical factors and chemical properties of herbicides.
a- Water solubility
Water solubility influence movement of herbicides in the soil, and very soluble herbicides move faster
downwards than less soluble herbicides.

b- Alkalinity
Alkalinity determine the degree to which the herbicide is positively charged and therefore its likely
wood to be adsorbed to the cation exchange sites. Under alkaline conditions more of the cations are
adsorbed to soil colloids. Most herbicides emit positively charged hydrogen ions to soil colloids and

175
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

the positively charged herbicides is coupled by soil colloids and is leached therefore persistence is
reduced.
c- Volatility
Herbicides evaporate at certain temperatures and lost as a volatile gas. Herbicides with high vapour
pressure are highly volatile therefore must be protected immediately after application to avoid loss due
to vapourisation. Highly volatile herbicides include perbulate , litraline, and trif.
Soil type
Adsorption on soil colloids
a. Strongest adsorption occurs in soil of high organic matter or high clay content. For herbicides to
be effective they must be in soil solution to be taken up by the plant. Herbicides adsorbed in clay
particles is not readily available to plants. Dosage rates are higher in clay soils than sand soils to
compensate for the higher adsorption power of clays. Persistence is higher in strongly adsorptive
soils e.g. clays.
b. Leaching is the down ward movement of chemical substances in solution through the soil. The
degree of leaching is determined by the chemical solubility, movement of water, and amount of
herbicide adsorbed by soil colloids.
c. Photo-decomposition
When a soil applied herbicide is exposed to light gradual photo-decomposition takes place. This effect
is minimized if rain or irrigation follows within a few days after application. In prolonged dry weather
shallow incorporation will reduce photo-decomposition.

d. Chemical decomposition
Organic compounds when applied to the soil are decomposed by micro-organisms and reserve the
energy. Many herbicides are organic compounds and are broken down by micro-organisms. This
process is facilitated by warm moist conditions in well aerated soils
Examples of persistence of herbicide
Herbicide persistence (weeks)
Simazine, durone,flomethrone 12-25
Atrazine, trif , nitraline 8-18
[Link],alachloa, linurone, cynazine ,terbutryne,
naphthalin,DNCP. 6-12
24D,MCPD 4-8

Summary
The use of herbicides is an important and easy approach towards the control of weeds; this method is
less labour intensive as compared to hand weeding. The different modes of action of different
herbicides in conjunction with different means of selectivity help in the control of different weed
species.

Activities

 How does weed dormancy influence the survival of the weed?

176
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 With the aid of diagrams explain the herbicide control mechanisms employed in the control of
weeds.
 Define herbicide persistence. Explain how persistence affects crop succession in the lands?

CROP WEED COMPETITION(allelospoly):

Concept
Competition is nothing but the struggle for existence and superiority. Competition exerts a powerful
force in the plant community,irrespective of the mechanism,tending towards limitation or extinction of
the weal competitors. Competition is maximum when available resources for crop growth become
limited.
Competition is a negative interaction where individuals make simultaneous demands that exceed
limited resources and while both suffer,one individual suffers less.
So, crop weed competition indicates competition between crop and weed in a natural ecosystem in
response to resources struggle for their existence and superiority.
Crop weed competition occurs in two broad aspects:
1. Direct competition- for nutrients,moisture,light and space.
2. Indirect competition- through exudation and or production of allelopathic chemicals.

LIST OF CHARACTERISTICS ASSOCIATED WITH COMPETITIVE PLANTS

Shoot characteristics
- Rapid expansion of tall,foliar canopy
- Large leaves
- A C4 photosythetic pathway and low leaf transmissivity of light.
- A climbing habit
- A high allocation of dry matter to build a tall stem
- Rapid extension in response to shading
- Leaves forming a mosaic leaf arrangement for best light interception.
- Horizontal leaves under overcast conditions and obliquely slanted leaves (plagiotropic) under
sunny conditions.

Root characteristics

177
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

- Early and fast root penetration of a large soil area.

- High root density/soil volume.
- High roo-shoot ratio.
- High root length per root weight
- High proportion of actively growing roots.
- Long and abundant root hairs.
- High uptake potential for nutrients and water
INTERSPECIFIC AND INTRASPECIFIC COMPETITION

Interspecific competition, in weed science, is a form of competition in which individuals

of different species compete for the same resources in an ecosystem (e.g. food or living space). Weeds
and crops are different species which [Link] can be contrasted with interspecific cooperation, a
type of symbiosis. Competition between members of the same species is called intraspecific
competition. For example weeds themselves may compete for resources.

Persistence of Weeds

It is the measure of the adaptive potential of a weed that enable it to grow in any environment. Weeds
are both hardy and persistent. Therefore, they are difficult to control.
Weeds are the best examples of successful struggle for existence on earth advocated by Charles
Darwin in the year 1859.

The persistence of weeds or weed adaptability is largely influenced by edaphic ( Soil), climatic and
biotic factors which affect its occurrence, abundance , range and distribution.

1) Soil Factors of Persistence of Weeds:

The soil factors which influence persistence of weeds are soil, moisture, aeration , temperature , PH
fertility level and also a cropping system.

Many weeds have adaptability to a wide range of soil environment. Many weeds can adopt and grow
in soils of low fertility , very high PH ( alkali soils) or low PH ( acidic soils) waterlogged soils etc.
however, crops can’t tolerate such abnormal conditions of soil. The distribution of weed species
depends upon conditions of the soil as below:

E . g 1. Moist Soil Condition: Kena ( commelina spp), Maka etc.

2. Water Logged Soils: Typha, Cyperus
3. Light Soils: Cock’s comb, Piwala dhotra, Euphobia hirta ( dudhi) , Aghada etc.
4. Heavy soils: Hariali, kunda.
5. Basophiles: Weeds growing on alkali soils ( PH 7.4 to 8.5 ) E.g Alkali grass, Quackgrass.
6. Acedophiles: Weeds growing on acidic soils ( PH 4.5 to 6.5 ) E.g Hariali, Digitaria spp.
7. Neutrophiles : Weeds growing on neutral soils ( 6.5 to 7.4 ) several weed species grow on neutral
soils.

2) Climatic Factors of Persistence of Weeds:

The important climatic factors that effect persistence of weeds are sunlight, temperature, rainfall,
wind, etc.
178
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

a) Sunlight:
Light intensity and duration are important in influencing growth, reproduction and distribution of
weeds. The photoperiod governs affects flowering and time of seed setting and maturation.
It has also important bearing on evaluation of different ecotypes within weed species. Tolerance to
shading is major adoption that enables weeds to persist.

b) Temperature:
Temperature of atmosphere and soil affects seed germination and dormancy which is major survival
mechanism of weeds. Many weeds are hardy and tolerate extreme condition of temperature.
c) Rainfall:

It has significant effect on distribution and persistence of weeds. The weed species found in desert and
low rainfall areas. E. g Acacia spp. Zizyphus spp, cactus, cock’s comb, piwala dhotra etc. are different
from those of aquatic environment or high rainfall area ( E. g Water hyacinth, hydrilla, cyperus iria,
maka etc.)

d) Wind:

It is the major factor in dissemination of weeds. It modifies the transpiration and evaporation losses
and balance of oxygen and carbon dioxide in the atmosphere. Wind can limit or restrict the occurrence
and persistence of weeds.

Climate has profound effect on persistence of weeds which can adopt to wide variety of climates.
Many weeds are hardy and can tolerate adverse climatic conditions. Many of the weed are C4 plants
e.g Nustedge, Hariali, Vasatvel, Portulaca spp. Amaranthus spp, wild oats, wild rice, chandvel etc.
The C4 weeds can withstand low or high CO2 levels, high intensity and high temperatures.

3) Biotic Factors of Persistence of Weeds:

Plants, animals, birds, insect and disease causing organisms are the biotic factors that modify the
growth of weeds and seed production in variety of way that affects weed persistence directly and
indirectly. Weeds produce large number of seeds has dormancy and viability for longer period and
also have dis-agreable taste and odour. All these characteristics of weeds enable them to survive even
under the various effects of biotic factors.

ALLELOPATHY
Allelopathy is the detrimental effects of chemicals or exudates produced by one (living) plant species
on the germination, growth or development of another plant species (or even microorganisms)
sharing the same habitat.
Allelo chemicals are produced by plants as end products, by -products and metabolites
liberalised from the plants; they belong to phenolic acids, flavanoides, and other aromatic
compounds viz., terpenoids, steroids, alkaloids and organic cyanides.
Allelopathic Effect of Weeds on Crops
(1) Maize
•Leaves & inflorescence of Parthenium sp. affect the germination and seedling growth

179
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

•Tubers of Cyperus esculentus affect the dry matter production

2)Sorghum
•Stem of Solanum affects germination and seedling growth
•Leaves and inflorescence of Parthenium affect germination and seedling growth
3) Wheat
•Seeds of wild oat affect germination and early seedling growth
•Leaves of Parthenium affects general growth
•Tubers of C. rotundus affect dry matter production
•Green and dried leaves of Argemone mexicana affect germination & seedling growth
4) Sunflower
•Seeds of Datura affect germination & growth
Allelopathic Effect of crop plants on weeds
(i) Root exudation of maize inhibits the growth of Chenopodium album
(ii) The cold water extracts of wheat straw when applied to weeds reduce germination and growth of
Abutilon sp.
Allelopathic effect of weeds on weeds
•Extract of leaf leachate of decaying leaves of Polygonum contains flavonoides which are
toxic to germination, root and hypocotyls growth of weeds like Amaranthus spinosus
•Inhibitor secreted by decaying rhizomes of Sorghum halepense affect the growth of Digitaria
sanguinalis and Amaranthus sp.
Factors influencing allelopathy
a. Plant factors
i. Plant density: Higher the crop density the lesser will be the allelo chemicals it encounters
ii. Life cycle: If weed emerges later there will be less problem of allelochemicals
iii. Plant age:
The release of allelochemicals occurs only at critical stage. For eg. in case of Parthenium, allelopathy
occurs during its rosette & flowering stage.
iv. Plant habit: The allelopathic interference is higher in perennial weeds.
v. Plant habitat: Cultivated soil has higher values of allelopathy than uncultivated soil.
b. Climatic factors:
The soil & air temperature as well as soil moisture influence the allelo chemicals potential
c. Soil factors:
Physico-chemical and biological properties influence the presence of allelochemicals.
d. Stress factors:
Abiotic and Biotic stresses may also influence the activity of allelochemcals
Mechanism of action of allelochemicals
Interfere with cell elongation
Interfere with photosynthesis
Interfere with respiration
Interfere with mineral ion uptake
Interfere with protein and nucleic acid metabolism
Use of Allelopathy in biological control of weeds:
1. Use of cover crop for biological control
2. Use of allelopathic chemicals as bio-herbicides
Effect of weed competition on crop growth and yield
1. Crop growth and yield is affected
2. Crop suffers from nutritional deficiency
3. Leaf area development is reduced
4. Yield attributes will be lowered
5. Reduce the water use by the crop

180
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

6. Affect the dry matter production

7. Lowers the input response
8. Causes yield reduction
9. Pest and disease incidence will be more
Crop mimicry
Crop mimicry is an example of the extent to which weeds have adapted themselves to survive in that
frequently disturbed site.
Crop mimicry is defined as the phenomenon whereby weeds develop morphological and or
biochemical close resemblance to some phases in the life history of a crop as to be mistaken for the
crop and thus evade eradication.
Types of crop mimicry
Vegetative mimicry:A situation where close similarity in appearance occurs between weeds and crops
at seedling and vegetative stages.e.g. wild rice (Oryza longistaminata) in cultivated rice ; wild
sorghum (Sorghum halepense) in cultivated sorghum, wild sugarcane (Saccharum spontaneum) in
sugarcane.
Seed mimicry:This is a situation whereby the similarities between weeds and crops is observed in
seed, weight, size and appearance. e.g. similarity in seed size between seeds of upland rice and those
of itch grass (Rottboellia cochinchinensis).
Biochemical mimicry:This is a situation in which a weed develops resistance to a herbicide that has
been used previously for selective control in a given crop

METHODS OF WEED CONTROL

Weed control is the botanical component of pest control, which attempts to stop weeds,
especially noxious or injurious weeds, from competing with domesticated plants and livestock. Many
strategies have been developed in order to contain these plants.

The original strategy was manual removal including ploughing, which can cut the roots of weeds.
More recent approaches include herbicides (chemical weed killers) and reducing stocks by burning
and/or pulverizing seeds.

A plant is often termed a "weed" when it has one or more of the following characteristics:

Little or no recognized value (as in medicinal, material, nutritional or energy)

Rapid growth and/or ease of germination

Competitive with crops for space, light, water and nutrients

Methods

Pesticide-free thermic weed control with a weed burner on a potato field in Dithmarschen

Coverings

In domestic gardens, methods of weed control include covering an area of ground with a material that
creates a hostile environment for weed growth, known as aweed mat.

Several layers of wet newspaper prevent light from reaching plants beneath, which kills them. Daily
saturating the newspaper with water plant decomposition. After several weeks, all germinating weed
seeds are dead.
181
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

In the case of black plastic, the greenhouse effect kills the plants. Although the black plastic sheet is
effective at preventing weeds that it covers, it is difficult to achieve complete coverage. Eradicating
persistent perennials may require the sheets to be left in place for at least two seasons.

Some plants are said to produce root exudates that suppress herbaceous [Link] minuta is
claimed to be effective against couch and ground elder, whilst a border of comfrey is also said to act
as a barrier against the invasion of some weeds including couch.

Gravel can serve as an inorganic mulch.

Irrigation is sometimes used as a weed control measure such as in the case of paddy fields to kill any
plant other than the water-tolerant rice crop.

Manual removal

Many gardeners still remove weeds by manually pulling them out of the ground, making sure to
include the roots that would otherwise allow them to resprout.

Hoeing off weed leaves and stems as soon as they appear can eventually weaken and kill perennials,
although this will require persistence in the case of plants such as bindweed.

Goat grazing

Companies using goats to control and eradicate leafy spurge, knapweed, and other toxic weeds have
sprouted across the American West.

"Stale seed bed"

Another manual technique is the ‘stale seed bed’, which involves cultivating the soil, then leaving it
fallow for a week or so. When the initial weeds sprout, the grower lightly hoes them away before
planting the desired crop. However, even a freshly cleared bed is susceptible to airborne seed from
elsewhere, as well as seed carried by passing animals on their fur, or from imported manure.

Irrigation

Drip irrigation involves bringing water directly to the roots of the desired plants, thereby limits weed's
access to water.

Biological pesticide

Vinegar kills the visible part of the weed. They will wrinkle and die next day, although the root will
still be in place to continue growing.

Tilling

Ploughing includes tilling of soil, intercultural ploughing and summer ploughing. Ploughing uproots
weeds, causing them to die. In summer ploughing is done during deep summers. Summer ploughing
also helps in killing pests.

Mechanical tilling can remove weeds around crop plants at various points in the growing process.

182
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Crop rotation

Rotating crops with ones that kill weeds by choking them out, such as hemp, Mucuna pruriens, and
other crops, can be a very effective method of weed control. It is a way to avoid the use of herbicides,
and to gain the benefits of crop rotation.

Thermal

Several thermal methods can control weeds.

Hot foam (foamstream) causes the cell walls to rupture, killing the plant. Weed burners heat up soil
quickly and destroy superficial parts of the plants. Weed seeds are often heat resistant and even react
with an increase of growth on dry heat.

Hybrid

One method of maintaining the effectiveness of individual strategies is to combine them with others
that work in complete different ways. Thus seed targeting has been combined with herbicides. In
Australia seed management has been effectively combined with trifluralin and clethodim.

"Organic" approaches

Organic weed control involves anything other than applying manufactured chemicals. Typically a
combination of methods are used to achieve satisfactory control.

Soil solarization in some circumstances is very effective at eliminating weeds while maintaining grass.
Planted grass tends to have a higher heat/humidity tolerance than unwanted weeds.

Herbicides

The above described methods of weed control use no or very limited chemical inputs. They are
preferred by organic gardeners or organic farmers.

However weed control can also be achieved by the use of herbicides. Selective herbicides kill certain
targets while leaving the desired crop relatively unharmed. Some of these act by interfering with the
growth of the weed and are often based on plant hormones. Herbicides are generally classified as
follows:

Contact herbicides destroy only plant tissue that contacts the herbicide. Generally, these are the
fastest-acting herbicides. They are ineffective on perennial plants that can re-grow from roots
or tubers.

Systemic herbicides are foliar-applied and move through the plant where they destroy a greater
amount of tissue. Glyphosate is currently the most used systemic herbicide.

Soil-borne herbicides are applied to the soil and are taken up by the roots of the target plant.

Pre-emergent herbicides are applied to the soil and prevent germination or early growth of weed
seeds.

183
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

In agriculture large scale and systematic procedures are usually required, often by machines, such as
large liquid herbicide 'floater' sprayers, or aerial application.

Resistance

Resistance occurs when a target adapts to circumvent a particular control strategy. It affects not only
weed control,but antibiotics, insect control and other domains. In agriculture is mostly considered in
reference to pesticides, but can defeat other strategies, e.g., when a target species becomes more
drought tolerant via selection pressure.

Farming practices

In wheat fields, introduced varieties of ryegrass, while good for grazing sheep, are intense competitors
with wheat. Ryegrasses produce so many seeds that, if left unchecked, they can completely choke a
field. Herbicides provided excellent control, while reducing soil disrupting because of less need to
plough. Within little more than a decade, ryegrass and other weeds began to develop resistance.
Australian farmers evolved again and began diversifying their techniques.

CHAPTER 24

CROP PESTS

Introduction
Pests are of great importance in the crop production as they reduce crop productivity. In this unit you
will general information about their classification and control.

Objectives

 outline the socio-economic importance of pests

 identify pests
 classify pests according to feeding habits
 describe the life-cycle of pests.
 explain the importance of pest management.
 explain economic threshold and economic injury levels of pests.
 describe methods of pest management.
 compare and contrast pest management methods

Definition
Pest – Any form of plant or animal life or any pathogenic agent injurious or potentially injurious to
plants, plant products, livestock or man.
Following this definition pests include insects and other arthropods, vertebrates, weeds and micro-
organisms e.g. fungi, fungi, bacteria and viruses.

Categories of pests
1. Key or major pests

184
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

This refers to the insect pests which occur perennially and cause serious and persistent economic
damage in an ecosystem in the absence of effective control measures. An example is the cabbage
webworm in brassicas in Zimbabwe.
2. Minor pests
These are pests whose populations under normal conditions do not cause economic damage.

3. Occasional pests
These are insects whose population build up would reach an economic injury level to the plants. An
example is the increase in the Lepidoptera populations which lead to heavy leaf defoliation to affect
the crop yield.

4. Potential pests
Potential pest refers to insect species with the potential to develop into major pest depending on the
situation present.

5. Migrant pests
These are insect pests which move from one area (zone) to another to cause an economic damage.
Examples are the red locust, armyworm.

6. Disease vectors
These are insects that transmit diseases to plants making them important to agriculture. Examples are
white fly which transmits tobacco and a cotton leaf curl virus, aphids transmits groundnut mosaic
virus, and leaf hopper which transmits maize streak virus. It is important to note that very low
populations of disease vectors can lead to serious crop damage.
Pest damage
Pests can cause plant damage in different ways. This can be through biting and chewing, piecing and
sucking and can be vectors of pathogens.

Pests with biting and chewing mouthparts

These are insects which feed by biting pieces of plant material and chewing. Very good examples of
such pests are: locusts, crickets, caterpillars and beetles.

Damage caused by biting and chewing pests

Losses in plant yield can be brought about in different ways by biting and chewing pests as indicated
below:
1. During feeding some pests produce substances which irritate the host plant leading to proliferation
and gall formation.
2. Loss of photosynthesis tissue due to the eating away of the leaf lamina of the plant and leaf drop,
e.g. by leaf worm
3. There is destruction of seedlings and young plants, e.g. vegetable seedlings can be destroyed by
cutworm.

185
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

4. Feeding of pests from the growing points lead to the destruction of buds and shoots e.g. the bud
worm in tobacco.
5. Some pests feed by boring and tunneling the stems interfering with the movement of sap up and
down the plant and weakening of the plant, e.g. maize stalk borer in maize.
6. During feeding the pests destroy flowers, seed and fruits there by significantly reducing fruit
production, e.g. cape mounted rifle beetle in sugar bean.
7. Tubers and roots can be bored or eaten by pests in the soil leading to reduced yields, e.g. potato
tuber moth in potatoes.
Pests with piercing and sucking mouthparts
Pests in this category have part or all of the mouthparts modified into piercing proboscis. They suck
sap from the xylem or phloem tissues of the plant. Examples of such pests are from the order Acarina,
homoptera and thrips.

Pests that are vectors of pathogens

These are insects that transmit diseases to plants making them important to agriculture. Examples are
fruit fly which transmits tobacco and cotton leaf curl viruses, aphids transmits groundnut mosaic virus,
and leaf hopper which transmits maize streak virus. It is important to note that very low populations of
disease vectors can lead to serious crop damage.

Pests life cycles

Pests go through different developmental stages during their life. Some go through complete
metamorphosis and some incomplete metamorphosis.
With most insects growth is limited to immature stages. Insects develop from stage to stage and the
change from one stage to the next is through moulting (ecdysis). The change in appearance of the
insect due to moulting is referred to as metamorphosis.
Complete metamorphosis
In complete metamorphosis insects go through the four stages of development which are as in the
figure 4 below.

Fig 4 Complete metamophosis

(Embryonic development) (Internal wing pads)
Egg Larva

Adult Pupa (External wing pads)

(Reproduction dispersal
and sometimes feeding)
186
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The four stages are mostly exhibited by holometabobla insects like the American boll worm (Heliothis
armigera), maize stalk borer (Bassiola fusca). In the complete metamophosis group of pests there are
four instars, a modified larval or pupal instar and the adult phase.
Incomplete metamorphosis
In this case insects go through three stages of development which are: as illustrated in the in the figure
5 below.
Fig 5 Incomplete metamophosis
Embryonic development
Egg

Adult Nymph (Larva)

Sexually mature 1st instar no wings

and winged 2nd to 5th instar is sexual immaturity
External wing pads
Insects falling under this group belong to the Homimetabola.
In incomplete metamorphosis group of pests, wings appear as external wing pads in the second instar
and they are fully formed after the fifth moult e.g. grass hoppers and locusts.

VIVIPARY
Viviparity (from the Latin vivus ("living") and parere ("to beget"), literally "giving live birth"),
both in zoology and in botany refers to certain classes of modes of reproduction.
In botany vivipary is a common alternative form of the term, and botanical definitions also vary. One
usage refers to reproduction via embryos, such as shoots or bulbils, as opposed to germinating
externally from a dropped, dormant seed, as is usual in plants; however, vivipary also may refer to
plants arising from buds or seedlings that have formed or germinated while still on the parent plant,
and are dropped when ready to establish themselves in the substrate.
In zoology viviparity refers to development of the embryo inside the body of the parent, eventually
leading to live birth, as opposed to reproduction by laying eggs that complete their incubation outside
the parental body. There are several forms of reproductive processes that answer to that description
however, and they differ so radically, both in their nature and evolutionary origin, that no definitive
and exclusive terminology is universally accepted, and the usage of the term is largely a matter of
convenience and preference.

Pest control methods

Pest control can be achieved using different means. The approaches to pest control are: cultural
method, natural control, mechanical, and chemical control.

187
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CULTURAL METHODS OF PEST CONTROL

These approaches to pest control reduce pest populations rather than completely destroying the pests.
The advantage of using cultural control measures is that there are no hazards to the user unlike with
chemicals. The different cultural approaches to pest control are explained below:

Crop rotation
Rotating crops of different families one after the other breaks the lifecycles of some pests. For
example if a maize crop is grown following a cabbage crop, pests like cabbage webworm (Helulla
hundalis) and diamond back moth (Plutella zylostella) lifecycles are broken leading to a reduction in
the pest populations. This implies that if a four year rotation is strictly followed the method becomes
effective. Another example is the growing of Katambora Rhodes grass after tobacco leading to the
control of the root knot nematode. This is because the grass is non host to the nematode hence it dies
from starvation resulting in a decrease in the populations of the nematode.

Tillage
Tillage mechanically damage the pests and their stages, burry or expose the developmental stages of
the pests, increase the growth vigour of the crops, eliminates the alternative host plants, and changing
the physical conditions of the soil e.g. moisture, oxygen concentration and soil pH resulting in the
death of most pest hence the reduction of pest populations.

Trap crops
With this approach, a susceptible crop is grown close to the major crop to attract the pests and protect
the major crop. The accumulated pests are then destroyed either chemically or by tillage or any other
suitable means.

Growing tolerant varieties

Some plants have the ability to recover from pest damage due to some tolerance in them. Plants with
low or no levels of tolerance cannot recover pest damage as a result it leads to economic damage.
Use of manure and fertilizer
Healthy plants are produced through proper manure application and fertilization. Healthy plants can
better withstand pest attack.

Pruning and thinning

Healthy and pest resistant new shoots can develop on the plant if the plant is occasionally pruned;
hence it is advisable to remove old branches from the plants.

Crop location
Pests may not move from one field to the other in adjacent fields if the crops in the adjacent fields are
deferent. This minimizes the spread of pests and keeps the pest populations lower.
Crop residue destruction
Burning of crop residues destroy the pests which overwinter in the crop residues e.g. maize stalk
borer, and cotton ball worms.

Alternating the planting times

188
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

By changing the planting periods one can avoid the infestation times of a particular pest there by
reducing pest infestation to the crop.
Natural control
a) Climatic factors
Populations of pests and their host plants are directly affected by humidity, temperature, and day
length. The changes in climate also limits the dispersal of pests.

b) Natural barriers
Features like mountain ranges, large water bodies and deserts limits the spreading of pests through
migration. Hence reduce the spread of some pest species.

c) Natural enemies
Most of the vertebrates feed on insects and keep insect populations low. Within the insect group are
predator species that capture and feed on insect pests or lay their eggs inside on or in the body of other
insects (parasitoids) for the newly born parasitoids to get readily available food.
d) Diseases
Plant pests like any other living organisms are attacked by diseases e.g. the semi-looper is affected by
the virus , plusia nuclear poly hydrosis virus.

Mechanical methods
Where manual and devices and machines are used to control pests this is referred to as the mechanical
methods of pest control. Following are some of the mechanical practices used in the control of insect
pests.

Shaking or beating of branches

Kerosinised water in a tub may be placed under the plant which is shaken or beaten with a stick to
drop the pests into the tub.
Wire gauge screens
Borers can be prevented from attacking plants by surrounding the stems and fruits of the plants with
wire gauge screens.

Hand picking
Insects can be handpicked and crushed to death e.g. beetles and caterpillars which are easily picked by
hand.

Banding
Trees can be banded to prevent insects from climbing up the plant to damage the top parts of the plant.
E.g. sticky materials can be painted on the tree or on waxed paper.

Traps
Different traps can be used to trap different insects due to different behaviour. Examples of traps used
in the control of pests are:
189
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Light traps, pheromone traps, vacuum traps, sticky traps and sound traping.
General Pesticide Safety
Describe safe handling of agrochemicals.
The following sections provide general or specific guidelines for handling pesticides. To help reduce
the hazards associated with pesticides:
 Do not transport, mix, or use agricultural chemicals unless you can summon help, if needed.
 Keep an ample supply of water nearby to flush exposed areas, if a spill occurs.
 Check all pesticide equipment before you use it to ensure proper working condition.
 Read pesticide labels carefully. Follow the label directions when mixing, applying, storing, or
disposing of pesticides.
 Wear personal protective equipment to prevent dermal, inhalation, and mucous membrane
exposure.
 Do not eat, drink, or smoke when handling pesticides.
 Launder clothing and bathe after working with pesticides to ensure that all chemicals are
removed from clothing and skin.
 Do not use agricultural pesticides around the home or office.
 Observe assigned reentry intervals. Always wear the appropriate protective clothing when
entering fields before the reentry date.
 Always handle pesticides downhill from wells, cisterns, sink holes, ditches, or standing water.
 Do not apply pesticides when rain is imminent or if wind could affect the spraying area.
 Triple-rinse spray equipment and empty containers. Apply the rinse water to the treated field.
 Properly dispose of empty containers.
Preparing to Apply Pesticides
Preparation is essential for chemical safety. Follow these steps to properly prepare for pesticide
application:
1. Plan Ahead
Always read chemical labels before attempting to work with pesticides. Prepare for a possible
emergency by maintaining a personal decontamination site, a chemical spill kit, and by knowing the
proper first aid procedures associated with your pesticide.
2. Move Pesticides Safely
Careless chemical transportation can cause spills and contamination. Do not carry pesticides in an
enclosed area, such as a car. Be sure to secure the pesticides to prevent shifting or bouncing. In
addition,never leave your vehicle unattended when transporting chemicals.
3. Select Appropriate Personal Protective Equipment
Regardless of the pesticide's toxicity, always wear a long-sleeve shirt and pants when working with
pesticides. Wear additional protective equipment, as necessary.
4. Select Application Equipment
Choose suitable equipment to properly apply pesticides. Before using the equipment, inspect it for
good working order.
5. Provide Prior Notification
Before applying pesticides, inform all people in or around the application area. Notification allows
people to protect themselves from harmful chemicals.
6. Mixing Pesticides
Always read and carefully follow label directions when mixing pesticides. Even if you are familiar
with a particular chemical, reread the label to ensure that you have the latest safety information. In
addition, follow these guidelines for mixing pesticides:
Wear Personal Protective Equipment
Always wear protective gear when handling hazardous chemicals.
Work in a Safe Area
190
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The pesticide mixing and loading area should be well ventilated, well lighted, and downhill from any
water sources. Concrete slabs are ideal for mixing chemicals since they allow for easy clean up.
Measure Chemicals Correctly
Measure and mix pesticides carefully. Never mix different pesticides except as directed by the label or
chemical manufacturer. Do not use more chemical than prescribed by the pesticide label. The overuse
of pesticides is illegal, and may result in the following:
 Higher pest control costs
 Pesticide residue in food
 Groundwater pollution
 Pesticide resistance
Pour Pesticides Carefully
Always wear a face shield and take care not to splash chemicals when pouring pesticides. Never use
your mouth to siphon pesticides.
Applying Pesticides
When you apply pesticides, you are responsible for protecting yourself, other people, and the
environment.
Follow these guidelines when applying pesticides:
Minimize Exposure
Even mildly toxic chemicals can harm you if you use them daily. Take care to minimize your
exposure to any chemical. Avoid working in pesticide spray, mist, or runoff. Always work with
another person when working with hazardous chemicals.
Avoid Applying Pesticides in Sensitive Areas
Avoid spraying pesticides near beehives or areas that humans normally occupy (e.g., schools,
playgrounds, hospitals, etc.). If you must apply pesticides in sensitive areas, do so when the weather is
calm and when people are not around.
Avoid Pesticide Drift, Runoff, and Spills
Pesticides that fall outside the targeted application area can be very hazardous. Choose weather
conditions, equipment, and chemicals that do not lend themselves to these hazards.
Avoid Equipment Accidents
Equipment accidents are often caused by poor maintenance and improper work habits. Avoid
equipment accidents by following all operating instructions.
Pesticide Storage and Disposal
Outline safe storage procedures for Agrochemicals
Always try to use all the pesticide in your application tank. If pesticides remain, use them on other
target locations. After emptying the tank, clean and store the equipment.
The following summary of EPA storage criteria should be followed for pesticides labeled with the
signal words DANGER, POISON, or WARNING, or the skull and crossbones symbol. These
procedures and criteria are not
necessary for the storage of pesticides classed as less toxic (CAUTION word on the label) or for those
registered
for use in the home or garden.
Site Storage

system.
Storage Facility

chemical fire extinguisher).

DANGER, POISON, and PESTICIDE

STORAGE).
191
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

decontaminated.

disposed of as excess pesticide; contaminated runoff collected and treated as excess pesticide.
Operational Procedures

egregate pesticides by formulation.

permit
access and inspection.

ck containers regularly for corrosion and leaks.

o Inspect pesticide containers for leaks before handling them.

o Do not allow unauthorized personnel in the storage area.
o Do not store pesticides next to items intended for consumption by animals or humans.
o Do not eat, drink, smoke, or chew tobacco where pesticides are present.
o Do not store beverages, food, eating utensils, or smoking material in the storage or loading areas.
o Wear rubber gloves while handling containers of pesticides.
o Wash hands immediately after handling pesticides. Remove contaminated protective clothing
immediately; extra sets of clean clothing should be nearby.

o Where large quantities are stored, inform the Facilities department.

o The Facilities Department will furnish the fire chief with home telephone numbers of responsible
persons.

o Unused or outdated pesticides must be disposed as hazardous chemicals.

o See the Hazardous Waste Disposal chapter for more information.
Pesticide Cleanup
Always thoroughly clean all pesticide equipment as soon as you are through with it. Leaving pesticide
residue in mixing, loading, or application equipment can result in accidental injury or death to
livestock or people or unwanted contamination of plants or soil.
Clean the inside and outside of pesticide equipment, including nozzles. Dispose of contaminated rinse
water as
directed on the chemical label.
Disposal
Outline the safe disposal of agrochemicals
-over insecticide suspension can be disposed of safely by pouring it into a specially dug hole
in the
ground or a pit latrine
ponds or
rivers.
Some insecticides, such as the pyrethroids, are very toxic to fish.

be
on the lower side of such areas.
-off water from hand washings and spray washings into the hole, and bury containers,
boxes

192
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

and bottles used for pesticides in. Close the hole as soon as possible.
- mitted, far away
from
houses and sources of drinking-water. For reuse of cleaned containers, see box .

degraded and do not cause environ-mental [Link] solution can be used to kill insect pests
such as ants and cockroaches.

similar
breeding places.
of pyrethroids for the treatment of mosquito nets and other fabrics can be used for a few
days after preparation.

from biting from below. Where bedbugs are a problem, mattresses can be treated. Do not stir liquids
or
scoop pesticide with bare hands. Use the pressure-release valve of the pump or a soft probe to clear
blockages in the nozzle). Wash the hands and face with soap and water each time the pump has been
refilled. Eat and drink only after washing the hands and face. Take a shower or bath at the end of the
day
Summary
Pest management is involving quite a good number of approaches for successful crop production. It
was revealed in this chapter that different pests damage crops differently depending on the mode of
feeding of individual pests, hence different modes of control become applicable.
Activities
i. Define a pest.
ii. Make a list of pests and classify them in any of the categories explained before
iii. Compile a list of pests in your locality and classify them according to their mode of feeding.
iv. Differentiate between complete metamorphosis and incomplete metamorphosis giving
examples of pests.
v. Explain the approaches to pest control with reference to particular crops.

CHAPTER 24

INTEGRATED PEST MANAGEMENT (IPM)

Introduction
IPM is much a clean approach to pest and disease control which is environmentally friendly. The
implementation of IPM produces crops which contain the least chemical residues. Its implementation
to pests and diseases reduces the costs of production significantly, hence the need for consideration.

Objectives
 Define IPM.
 Define the components of IPM
 List the cultural practices used in IPM
193
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Identify the precautions one should observe when handling chemicals

Definition
Integrated pest management is an integrated approach to the management of pests (insect pests,
diseases, animals and weeds) in a sustainable manner. The approaches include biological, cultural,
physical and chemical approaches.

IPM practices leading to sustainable pest conntrol

Identify pests and asses pest levels in the field.
Know the pest history, development and the factors which influence its availability.
Keep low levels of pests in the field as this ensures that natural enemies are always available.
The main objective of IPM is to keep and maintain low levels of pests and make sure that they are
always below the economic injury level.
Principles of integrated pest management
Ensure healthy plants at all times
Conserve natural enemies
Chemical control should be taken as the last option in pest and disease control

COMPONENTS OF IPM

 Monitoring
Involve operations like scouting to detect, identify and determining the pest levels in the fields.
 Forecasting
Is the prediction on the development of the pest to economic injury level depending on the
environmental conditions.
 Thresholds
Thresholds refer to the pest levels at which economic losses are realized. Threshold levels are
also used to estimate the severity of the pest observed. Some control measures should be taken
at this level.

MANAGEMENT TACTICS IN IPM

1. Cultural
- Crop rotations are planned in such a way that they reduce the buildup of pests and diseases.
- Avoid planting susceptible plants in areas where particular pests are known too be problematic.
- Destroy plant residues in order to remove overwintering conditions for the pests.
- High relative humidity should be avoided at all costs to minimize the development of some pests
and diseases.
- Winter ploughing exposes some pest species to natural enemies and reduces pest pressure.
- Planting times should be such that they avoid the peak periods of particular or known pests during
the vulnerable stages of the crops.
- Proper fertilization of crops ensures high growth vigour to avoid stress that may predispose the
crop to diseases.
- Farmers should plant disease free seed to avoid early disease attacks.
- practicing good sanitation.
194
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

- Good weed control practices to avoid seeding reduce pressure to crops and promote plant growth
and it also reduces host plants to pests and diseases.
- Tap crops are also very useful in reducing pest pressure on the major crop.
- Proper irrigation scheduling reduces continuously humid conditions which favour the development
of most fungal diseases.
2. Physical and mechanical control
- Hand picking of pests can be used on some pests e.g. cape mounted riffle beetle.
- Use of traps for flying insects and sticky boards for crawling pests.
- Barriers prevent pests from reaching the crop.

3. Biological control
- Use of predators reduces pest populations significantly.
- Conserve natural enemies to keep pest populations low.
- Use of cultured bacteria to control some pest species is also effective.

4. Chemical control
Chemical control should be regarded as the last option in choosing control measures for pests and
disease control.
The right chemical should be chosen for the right pest at the right time, applied at the correct rate,
distributed correctly to ensure efficiency of the chemical used.
[Link]
Legislated planting and destruction dates should be observed as they minimize the spread of pests e.g.
in cotton and tobacco.

SAFE USE OF PESTICIDES

Get any chemical containers with amber, green, red and purple triangles to identify the precautions to
follow when using each chemical.
1. Pesticide transport
During packaging avoid packing pesticides together with food staffs and avoid transporting pesticides
together with human food.
Also avoid transporting pesticides in the passenger compartment in any vehicle.
2. Storage of pesticides
The store room should be well ventilated, kept under lock.
Only enough pesticides for the season should be bought.
Pesticides should be stored out of reach of children and apart from feed and food staffs.

3. Chemical hygiene
After applying chemicals remove protective clothing, wash hands and face with soap and clean water,
wash the used equipment avoiding the contamination of public water and destroy empty containers.
Do not recycle or put the empty containers to other uses.

[Link]

195
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Chemicals selected for use in any one crop should be registered. They should posses the registration
number and a colour triangle. It is advisable that one should buy originally packed chemicals from the
suppliers and avoid buying repacked chemicals.
What should you do to use chemicals safely?
 First of all read and understand the instructions and all other information given on the label
 All chemicals should be equally regarded as potentially dangerous
 Chemical users should wear protective clothing whenever chemicals are to be used.
 Pesticides should be used only when necessary.
 Recommended rates should always be used to maintain the efficiency of the chemicals.
 Over application of the chemical may lead to crop damage and over accumulation of crop residues
beyond the limit and under application may lead to ineffective pest control.
 Pre-harvest intervals should be observed where crop consumption is direct.

Factors to consider when choosing a pesticide for use

 The use of chemicals should always be the last option after all other possible measures have been
exhausted.
 The choice of the chemical should depend on the following
 Choose the right chemical for the right pest to be controlled.
 Chemicals should be used when the pest level reaches the economic threshold level.
 Choose the least dangerous chemical, but the chemical should be effective against the problem.
 Choose a chemical that can be safely applied with the available equipment.
 Choose affordable and available chemicals.

Summary
The way a farmer protects the crop from planting to harvesting from all the possible factors which
lead to the reduction in yield can lead to a sustainable yield. Integrated pest management practices
involve different means of pest control. The use of different methods of pest control reduces the risk
of chemicals to human beings.

Activity
i. Define IPM and make a list of the cultural practices used in IPM in your area.
ii. What are the precautions one should observe during the use of pesticides?

CHAPTER 25

CROP DISEASES

Introduction
Crop diseases cause heavy losses in crop production if left unchecked. There are many causes of
different diseases e.g. fungi, virus, and bacteria. Different causative agents cause different forms of
damage to the crop resulting in different symptoms to different diseases.
Objectives
By the end of this unit you should be able to:

 outline the socio-economic importance of diseases

196
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 classify diseases into bacterial, fungal and viral.

 describe signs and symptoms of diseases.
 describe mode of transmission of diseases
 explain importance of disease management
 outline the different methods of disease management
 compare different methods of disease management
 calibrate a knapsack sprayer

Definition of disease
Any deviation from the normal functioning of the plant structure.

GROUPS OF DISEASES
Parasitic diseases
These are caused by fungi, virus, or bacteria.

Signs of parasitic diseases

 Spot diseases
Are generally caused by fungi or bacteria. They characterized by spots. E.G bacteria spots in
sunflower and ground nuts. Cercospora leaf spot, alternaria, wildfire and angular spot in tobacco.

 Blight diseases
Blights kill large areas of tissue e.g. potato blight, leaf blight in maize

 Wilts
These are characterized by total wilting of leaves due to infection of roots or vascular system e.g.
bacterial wilt in potatoes and tomatoes.

 Canker
Is a disease of woody tissue which causes shrinking and cracking of bark e.g. bacterial canker in
tomatoes.
SYMPTOMS CAUSED BY FUNGI ON PLANTS
Fungi cause local or general symptoms on their hostsand such symptoms may occur separately or
concurrently
or may follow one another. In general, fungi cause local or general necrosis of plant tissues, and they
often
cause reduced growth (stunting) of plant organs or entire plants. A few fungi cause excessive growth
of infected plants or plant parts. The most common necrotic symptoms are as follows.
Pythium (DAMPING-OFF)
-off diseases of seedlings occur worldwide in valleys and forest soils, in tropical and
temperate climates, and in every greenhouse.
plants.

either before or after emergence.

197
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

seedlings in seedbeds are completely destroyed by damping-off or they die soon after they are
transplanted.

dampingoff infections in the preemergence stage.

 Rots
Are characterized by rapid disintegration of tissues of plant organs e.g. cob rots in maize.

 Mildews
Powdery mildew – is characterized by white powdery coating on leaves and stems e.g. in wheat and
peas.
Downy mildew – is caharacterised by bluish gray mouldy growth usually on the underside of the leaf
e.g. in brassicas, sorghum etc.

 Rust
This disease appear on the leaves and stems and produce pustules which are rust coloured (red brown
to orange) e.g. rust in wheat.

 Smuts
Appear as powdery masses on cereals e.g. head smut on maize.

 Mosaics
Mosaics are viral diseases. Indicated by paten of light and dark coloured portions on the leaves e.g.
cucumber mosaic virus.

 Stunting disease
Caused by virus e.g. Rosette virus in ground nuts.

Transmission of Plant Diseases

All parasitic as well as viral diseases are transmissible, the parasites or issues being infectious to
suitable host plants with ability to spread from host to host and from one area to another. The
microscopic parasites or sub-microscopic infectious agents viz. Viruses causing plant diseases are
technically termed as ‘pathogens’.

Wherever a disease is established in a particular area or country, transmission of the pathogen from
host to host or from one place to another is termed as ‘dissemination’ or ‘dispersal’ of the pathogen.
Dissemination of plant diseases is recognized in relation to different phases of diseases as under:

1. Primary infection: Contact of a pathogen with a suitable host plant and initiation of the
disease first time in the season of a crop is called ‘primary infection’. Often a few or several
plants in the crop are likely to get primarily infected.

198
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2. Secondary spread: When a plant or few plants are primarily infected, rapid multiplication of
the pathogen sets in under favorable climatic conditions, which helps ‘secondary spread’ of the
disease.

Modes of primary infections:

According to the general mode of primary infection plant diseases are recognized as:-

1. Soil borne

2. Seed borne, including diseases carried with planting material.

3. Wind borne

4. Insect borne etc.

Continuous and discontinuous transmission: -

Transmission of disease is termed as ‘continuous’ when it occurs naturally by way of growth,

multiplication and spread of the pathogen in an area or country where the disease is established. At
times, however, in an area or country where a particular disease has never occurred, it may get
introduced through the agency of man carrying diseased material to a new locality or to a distant
country for the purpose of introduction of new plants, crops, varieties etc. such transmission, of
course, is unnatural and regarded as ‘discontinuous’ transmission. Altogether different control
measures viz. Quarantine Regulations are enforced to check such discontinuous transmission.

Direct and indirect transmission:-

For classifying the methods of disease transmission in relation to the methods of suitable control
measures, the following two groups can be conveniently recognized.

1. Direct transmission: - Disease transmission where the pathogen is carried externally or

internally on the seed or planting material like cuttings, sets, tubers, bulbs etc.

2. Indirect transmission:- The pathogen spreading itself by way of its persistent growth or
certain structures of the pathogen carried independently by natural agencies like wind, water,
animals, insects, mites, nematodes, birds,implements and tools etc. are the different methods of
indirect transmissions.

Direct transmission: -

1. Internal transmission through seed or planting material:- False smut disease as well as
Helminthosporin Blight disease of wheat are the common examples of fungal diseases carried
internally through apparently healthy seed. Ring rot and Brown rot of potato caused by
bacteria are carried internally through the tubers. The well known whip smut and red rot of
sugarcane are fungal diseases carried internally in the planting sets. Mosaic and leaf roll of
potato which are viral diseases are also carried inside the infected tubers.
199
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2. External transmission through seed or planting material:- In this mode of transmission the
pathogen is carried externally over the surface of seed or vegetatively propagated plant parts
like sets, tubers, bulbs etc. or may even be carried as a physical mixture of fungal structures
with the seed. Indirect transmission: -

1. Autonomous transmission:- It takes place by continuous and persistent growth of the threads
or ‘hyphae’ of the causal fungi in soil, characteristic of several wood rotting fungi attacking
forest trees and some fruit plants. Some root rotting fungi infecting certain seasonal crops also
are transmitted by this method. The autonomous dispersal of such soil fungi may range from
few cm. To several (8 to 10) meters in a single season. Some plant parasitic nematodes also
exhibit active but limited mobility in the soil.

2. Wind dispersal: - Fungal spores produced externally on host surfaces are most easily carried
by wind currents and this is the most dangerous mode of transmission of plant pathogenic
fungi like those causing powdery and downy mildews, leaf spots, blasts, blights and rust
diseases. Spores may be carried from low to very high altitudes of 12,000 to 14,000 feet and
from short distances to very long distances of several hundred kilometers.

3. Water dissemination: Disease transmission through the agency of water in different ways is
comparatively less important as compared to the wind transmission. Splashing rain drops
mostly transmit the foliar diseases from leaf to leaf, from shoot to shoot and even from plant to
plant in case of closely spaced crops. Such transmission is usually accompanied by wind
dispersal as well. Plant pathogens requiring high humidity conditions like the fungi causing
downy mildew diseases or bacteria causing canker of citrus are well adapted to this kind of
short distance water dispersal.

Certain soil inhabiting pathogenic fungi and bacteria causing root and collar rots, wilts, foot,
rots, etc are likely to be transmitted to much longer distances through the agencies like
irrigation water, streams and rivers, etc. It is also an important agency in transmission of seeds
of higher flowering parasites like dodder and striga.

4. Animals: Farm animals serve as disease transmitting agents in some cases. They are likely to
carry the pathogen externally on their body surface, particularly on legs and hoofs, etc. or
internally through their intestinal tract. Commonly, the soil inhabiting fungi causing rots and
wilts are carried externally while certain smut fungi causing diseases to grain crops are
transmitted through the intestinal tract.

5. Birds: Although birds play a very minor role in disease transmission, in cases of dispersal of
seeds of higher flowering parasite. Loranthus sp. Parasitising certain trees like mango, etc.
their role is of great significance. They transmit loranthus both externally and internally.

6. Implements and Tools: Farm implements used for cultivation of soil are often likely to
transmit plant pathogens from one place to another. The pathogens in this case are usually
200
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

carried in the form of bits of plant disease debris lying in the soil. Similarly tools used for
carrying out operations like cutting, pruning, budding, grafting, thinning,etc. also help in the
transmission of certain diseases from plant to plant. Several viral diseases are disseminated
through the budding and grafting operations.

7. Insects: Most of the viral diseases of plants are transmitted through the agency of different
insects. Both types of insects viz. sucking and chewing or/biting are capable of transmitting
viral diseases. The transmission may be simply `mechanical’ or it may be `biological’. In the
latter case the specific insect and the specific viral pathogen have some kind of association or
relationship between the two. Insects in such cases are called the `vectors’ for the particular
viral pathogen. In case of mechanical transmission the pathogen is simply carried externally or
internally by the insect.

Viruses carried `biologically’ by the insect vectors are of two types:

1. Non-persistent-viral pathogen requiring no latent or incubation period in the insect body.

2. Persistent: viral pathogens requiring certain incubation period inside the vector body before
they are inoculated or transmitted to healthy [Link] insects responsible for transmission of
viral diseases belong to the species of aphids, jassids (leaf hoppers), white flies, mealy bugs,
etc. Certain bacterial and several fungal pathogens are also known to be carried by insects.

1. Mites: Mites in contrast to insects are wingless anthropods resembling ticks and having four
pairs of legs and no antennae. It is suspected that some viral diseases of chillies, tomato,
brinjal, etc. have vector relationship with mites.

2. Nematodes: Nematodes have been observed to transmit viral, bacterial and fungal plant
diseases. Nematodes feeding externally on host plant roots cause injuries to roots which
become the avenues for entrance of fungal and bacterial pathogens infecting plant roots. The
Fan-leaf virus of grapevine is a well known example of transmission through a species of
nematodes.

3. Biological transmission: Dodder which is higher flowering parasite is known to transmit

certain viral diseases which remain `persistent’ in the dodder plant. The flowering parasite
after acquiring the virus from infected plant does not show any symptom itself but remains
capable of transmitting the virus to healthy hosts.

4. Human dispersal: Man is often responsible for transmission of plant diseases in two ways viz.

1. Workers handling seedlings, other planting material or fruits are likely to get personally in
contact with plant pathogens like fungi or bacteria. While handling the diseased material and
unknowingly and indirectly transmit the pathogens to healthy seedlings or plant parts through
his contaminated hands. This is a kind of `continuous’ mode of transmission.

201
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

2. The other or`discontinous’ mode of transmission for which only man is responsible is the most
efficient and equally dangerous phenomenon of transmission of plant diseases between distant
geographical areas often separated by physical barriers like oceans, mountains or deserts, etc.
Such long distances transmission of a disease to an area or country hitherto free from the
disease is usually accomplished by the transport of infected seed, nursery stock or timber, etc.
Thus it is a kind of direct transmission through propagating material.
Diseases can also be caused by nutrient deficiencies, Dodders and cuscutas (parasitic plants)

DISEASES CONTROL METHODS

 Cultural control method

 Rotations – break cycle of diseases
 Ploughing in and destroy trash except soil borne pathogens
 Weed control – relate to insect transmitted diseases
 Use of certified seed
 Time of planting – diseases avoided by growing crops when conditions are unfavourable to
diseases e.g. bush top is not common in early planted crops
 Hygiene- avoid taking the disease by washing equipment
 Selection of healthy seedlings
 Spacing
 Fertilizer application - apply correct levels , over application stimulate rank growth which affect
micro-climate
 Use resistant varieties
 Legislation- e.g. cotton and tobacco
a. Legislation that stipulates eradication with varieties e.g. moniclar potatoe
b. Notifiable diseases e.g. virus Y in potatoes
c. Time of planting is stipulated
d. Quarantine – applied to imports
3. Agricultural services – Use research findings on the crop. Use set light or photo to identify the
diseases.

Agrochemicals
Pesticide formulation
Dusts;Granules;Sprays;Aerosols & Fumigants

 Dusts- active ingredient is mixed in inert carrier e.g. lime- not mixed with water. Apply when
calm to avoid breathing in.

 Granules- formulated as granules to lessen health hazards. – are very easy to apply , need no
water e.g. Dipterex

 Sprays – Wetable powders are sprayed with a wetting agent but still need constant agitation.
Disposal powders have finer particles than WP used with ULV spray.

202
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Emulsions are suspensions of oil droplets in water. Pesticide dissolves in oily organic solvents
together with emulsifying agent. Agrochemicals are sold as concentrates therefore have to be diluted
before use. Agrochemicals are very expensive. Emulsions break down quicker than powders, less
easily washed by rain and suitable for ULV spray.

 Aerosoles – smokes and fogs. Are not for agricultural use Used on shelter places e.g. houses

 Fumigants - act in gaseous form. Fumigants are usually used for soil pests or stored products.
Used on confined space

Types of pesticides
Plant derivatives – pyrethrum
Inorganic - lead arsenate
Organo clorine -DDT , Aldrin
Organo phosphate - malathion
Carbonates - larvin , carbaryl
Benzilic acid compounds
Amadines - kelthin , acricide ,morocide
Pyrethroids – karate , marvirik , agrithrin
Colour coding - show the level of toxicity of a chemical
Green - is a safe chemical
Amber - dangerous
Red - very dangerous
Purple- extremely dangerous (lethal)

Modes of functions of pesticides

Four ways
Stomach poisons – have to be ingested
Contact poisons – act by direct contact
Systemic chemicals – are stomach poisons absorbed through the plant.
Fumigants - act by inhalation contact

Use of pesticides
Used as seed dressings
Used as baits – most suitable for small scale orchards
Used as sprays –most common and effective method.

203
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Full cover or wet spray – used for non mobile insects e.g. scales
Soil sprays – are systemic pesticides – do not affect innocent insects

Sprayer Calibration

Steps in calibration
There are a number of ways to calibrate sprayers. This method is one that may be easier with
backpack sprayers. [Link] need a tape measure, and you may find a calculator convenient:
1. Select the spray tip or boom that provides the desired coverage.
2. Add water and spray the ground or dry pavement as if you were spraying your field. Now check the
spray pattern for uniformity (and proper spray pattern overlap if [Link] using a boom). Adjust nozzle
spacing and/or height until you achieve the desired pattern. Be certain [Link] getting uniform
3. If all is well, add exactly 2 gallons of water to the tank. (Note: You can use any amount of water,
but remember to substitute your figure whenever you see .2 gallons. in the example that follows step
9.)
4. Mark your starting spot.
5. Spray the water as if you were actually spraying your field. Remember, you must maintain
• constant pressure,
• constant walking speed, and
• consistent height of the nozzle or boom over your spray target.
6. When the water is gone, stop and mark the spot.
7. Measure the area you sprayed and calculate square feet (length of swath × width).
8. Calculate how much of an acre you covered:
number of ft2 you sprayed
43,560 ft2/acre = acre sprayed
9. Calculate how many gallons/acre you sprayed:

2 gal sprayed
acre sprayed = gal/acre
Example. Let’s say that we sprayed 3 rows of Christmas trees with 2 gallons. Rows were 5 feet apart,
and each row was 387 feet long. The area sprayed was:
3 rows × 5 ft/row × 387 ft = 5,805 ft2
Now calculate gallons/acre, in two steps:
First, to find the acres we sprayed with 2 gallons, we divide the square feet we sprayed by the number
of square feet in an acre:
5,805 ft2 sprayed
204
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

43,560 ft2/acre = .13 acre

Next, divide our 2 gallons by .13 acre, to find our gallons/acre rate:

2 gal
.13 acre = 15 gal/acre
With the above walking speed, nozzle(s), pressure, and coverage, we sprayed 15 gallons/acre. That.s a
very important number. [Link] need it to determine the amount of pesticide to add per gallon. You can
also see that if you fill the sprayer with 5 gallons each time, it will require 3 tankfuls of pesticide and
water to cover 1 acre completely.
Summary
The use of agrochemicals in the control of pests should come in as the last option in pest control. For
you to choose a chemical to use on a particular pest you should be able to identify the pest, know the
mode of feeding of the pest and be able to choose the correct chemical for the pest to be controlled.

Activity
i. Identify the crop diseases in the farms around your area and categorize them accordingly in
any one of the above categories.
ii. Briefly explain how each of the diseases listed previous can be transmitted from one crop to
the other and how best it can be controlled.
iii. Using the pests identified in unit two select the chemicals suitable for the control of each pest.

CHAPTER 26
CROP PRODUCTION

AGRONOMIC PRINCIPLES

 explain the basis for dividing Zimbabwe into agro-ecological zones.

 design suitable cropping programmes for each agro-ecological zone
 distinguish primary from secondary tillage
 explain the significance of primary and secondary tillage in crop production
 calculate plant population per unit area
 discuss factors that influence plant population
 discuss implications of plant population in crop production
 explain the principles of crop rotation
 discuss the advantages and disadvantages of crop rotation
 design a 4-crop rotation cycle

AGRO-ECOLOGICAL ZONES OF ZIMBABWE

Introduction
Zimbabwe has five natural regions. These regions are divided according to the rainfall pattern of their
respective areas. In this chapter, we are going to discuss the five natural regions of Zimbabwe.

Terms
205
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Ecology- is the scientific study of interactions of organisms with one another and with the physical
and chemical environment
Agro-ecology – loosely it incorporates ideas about a more environmentally and socially sensitive
approach to agriculture, one that focuses not only on production, but also on the ecological
sustainability of the production system (Altier, 1998). “At its heart is the idea that a crop field is an
ecosystem in which ecological processes found in other vegetation formations such as nutrient
cycling, predator/prey interactions, competition, commensalism, and succession changes occur”
Alstier, 1998.
Agro-ecology is about agricultural productivity and ecological sustainability the main aim being to
understand the agro-ecosystem (soil, animals and vegetation) and manipulate it to produce higher
yields using fewer external inputs whilst minimising negative environmental and social aspects
Agro-ecological classification – land use categorisation in terms of farming systems best suited to
the natural characteristics
Natural region – a relatively large area where land use is common throughout and the agricultural
activities are conditioned by one or a few natural characteristics.
Diversified farming – a system with several lines of production not directly interlinked
Agro-ecosystem - a type of ecosystem or physical habitat where soils, animals and vegetation
interact. The soils and water (environment) are well managed to minimise biophysical constraints so
that the crop realises its full potential.
Objectives
By the end of this chapter, you should be able to;

 Define the following terms: Natural Region, Natural area, pentad, dry spell, and drought.
 Discuss the characteristics of the natural regions of Zimbabwe.
Definition of terms

 Natural region is a relatively large area where agricultural development is and will be conditioned
by one or a few natural characteristics. The natural characteristic dominant in Zimbabwe is rainfall
hence natural regions in Zimbabwe are largely demarcated on the adequacy and efficiency of
rainfall.
 Natural areas are areas in the same natural region with varying characteristics largely soil. The
rainfall considered here is effective rainfall.
 Pentad is a five day period. A rainy pentad is a centre pentad in three consecutive pentads which
should receive at least 40mm and two of which must receive at least 8mm of rainfall. Pentads give
a clear picture of rainfall reliability by dividing the year into 73 pentads. The Zimbabwean rainfall
season runs from 1st October to 30th April and this coincides with pentad 56 to pentad 24 to give a
40pentad period.
 Dry spell a period of 11-20 days without significant rainfall, that is, less than 25mm in heavy soils
and less than 15mm in light soils.
 Drought is a period of 21 days or more without significant rainfall.
Basis for dividing the country into Agro-ecological zones
Agro-ecological classification also known as natural region (NR) classification of Zimbabwe, divided
the country into five regions based on mean annual rainfall and was done in the 1960s (Vincent and
Thomas, 1960).

1. Rainfall.

206
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Monthly averages and the yearly averages for each station were computed. From the yearly averages,
the long- term mean annual rainfall was calculated.
2. Temperature.
Daily temperature records were obtained for the same period (1972-2006) also from the Department of
Meteorological Services. The long-term mean averages were then computed and used to describe the
temperature ranges for the new regions. Temperature was used to calculate the length of growing
period and to describe the new regions, just like rainfall.

3. Length of cropping period.

Length of growing period data were obtained from the FAO New Clim model (FAO, 2005). To obtain
the length of growing period ranges, the days to maturity and corresponding harvest index of maize
varieties in Zimbabwe were analysed as adapted and modified from FAO (1978). Numerical values
were allocated to the length of growing period for each meteorological station and these were used to
obtain a length of growing period suitability map.

4. Soils.
The chemical and physical properties of the soil that are relevant in crop production were used in
rating of the productivity of the soils. These parameters included the basic soil requirements of major
crops grown in Zimbabwe, which are water holding capacity, soil structure, the nutrient status of the
soil and its depth. Soils were rated with the assumption that the productivity of certain soils can be
improved if certain economically feasible improvements are made.

Agro-ecological zones of Zimbabwe

Natural Region I
It is a specialised and diversified farming region. Altitude is 1700m or more above sea level. The
rainfall is >1 000mm per year. Temperature is cool and is normally around 20oC on average. The
intensive safe use is afforestation (plantation crops) like pine and wattle trees. There is also intensive
livestock production especially dairy as well as deciduous fruit tree production. In frost free areas,
there is production of macademia nuts, potatoes, coffee and tea production. It covers less than 2% of
the country. The area stretches from Nyanga to Tamandai.
Natural Region II
It is a very intensive farming region. Most of the country’s maize and wheat production occurs in this
region. Most of the rainfall is confined to summer and it averages 750-1 000mm/year. It has two-sub-
regions namely IIa and IIb.
NR IIa receives an average of 18 rain pentads per season. It rarely experiences any severe dry weather.
The region is suitable for intensive crop production and also livestock production.
NR IIb receives an average of 16-18 rain pentads per year and it normally experiences severe dry
spells in the rain season and at times it may have short rain periods. Crop production may be affected
in some of the seasons but the effect is not sufficient enough to change utilization from intensive
system.

Natural Region III

207
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

This is a semi-intensive farming area. It is involves mixed farming. It receives rainfall of about 650-
800mm a year. The region has an average of 14-16 rainy pentads per year. The region is subject to
mid season dry spells. It is suitable for livestock and fodder production. There is marginal production
of tobacco and maize. The region covers about 19% of the country.
Natural Region IV
The intensive safe use is semi-extensive farming. It receives 450-650mm of rainfall per year. The
region is prone to drought and severe dry spells hence not suitable for crop production. Production is
mostly livestock, natural forest and some fodder crops. It is recommended to grow small grains which
are drought resistant. The area covers about 38% of the country.

Natural Region V
This is an extensive farming region which receives less than 450mm rainfall per year. The rainfall is
too low and very erratic and therefore is not suitable for crop production. Crops that are grown in this
region are grown under irrigation. The region is suitable for cattle ranching and wildlife management.
It covers about 27% of the country.

Cropping Programmes for Agro-ecological zones Summary

Natural Farming region Crops

Region 1  Afforestation(wattle and softwoods)
 Plantation crops(tea,coffee)
 Fruits and vegetables(apples,bananas,potatoes,peas,cabbages)
 Protea flower production
 Seed potato production
Region 2  Cash and food crops (maize,soyabeans,tobacco,potatoes and
cotton)
 Irrigated wheat and barley
Region 3  Drought tolerant grain crops(sorghum,millet,cotton,maize SC
513,SC501,groundnut and sunflower)
 forestry
Region 4  Unsuitable for dryland farming
 Drought tolerant crops such as sorghum,pearl millet,finger millet
and maize
 Irrigation is necessary eg for sugarcane
Region 5  Drought resistant crops are grown for household food security

Summary.
The five agro-ecological regions of Zimbabwe are natural regions I, II, III, IV and V. Natural region I
receives the highest amount of rainfall greater than 1 000mm and natural region V receives the least
rainfall of less than 450mm. this rainfall amount therefore dictates the type of production in each
particular region.

Activities.
a) Explain the significance of Natural regions to agriculture.
b) Describe the agro-ecological zones of Zimbabwe.
c) Define pentad, natural area and natural region.
208
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

d) Differentiate a dry spell from drought.

References

 Department of Meteorological Services (1981) Climate Handbook of Zimbabwe, Government

Printers, Harare.
 Makawa J. e.t.l (1987) School Atlas For Zimbabwe, Esselte Map Services, Stockholm, Sweden.

CHAPTER 27

TILLAGE

Tillage is the modification of soil physical structure for establishment and growth of crops. It involves
the manual or mechanical stirring actions carried on a soil for the purposes of growing crops. From the
above two definitions, it is clear that tillage refers to any disturbance of the soil surface.

Characteristics of good tilth

1. Good soil tilth should be mellow or friable (mellowness or friability of soil is that property of the
soil by which the clods when dry, become more crumbly; crumbling of soil = gradual disintegration of
clods)
2. Soil with good tilth is quite porous and has free drainage up to the water table. The capillary and
noncapillary pores are about equal so that sufficient amount of water is retained in soil as well as free
[Link] aeration, activity of microorganisms and chemical reactions in the soil increase with increase
in noncapillary [Link] porosity and pore size distribution are important to improve moisture
retention and availability of oxygen in the rooting zone.
3. Powdery soil encourages surface crusting when the soil dries after wetting
4. Soil aggregate should be stable enough to withstand water and wing erosion
Aims of tillage

 To loosen and break up the soil in order to increase aeration and water infiltration.
 To prepare a seedbed of suitable tilth for easy planting, good germination and emergence of crops.
 To break hard and capped soils at the end of dry season.
 To break soil compaction cause by traffic during field operations.
 To promote better root development and increase yield through deeper and more thorough
cultivation.
 To minimise soil erosion.
 To destroy weeds. For example perennial grasses like cynadon dactylon and cyperus spp.
 To enhance organic matter decomposition.
 To conserve moisture.
 To expose soil pest to their predators and the heat of the sun. The heat of the sun can also desiccate
soil pathogens.

Timing of land preparation.

Early land preparation is done soon after harvesting around March to May. It has several advantages
which include the following:

 A good tilth is achieved since the soil will be still moist.

209
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 There is early and good burial of plant debris in moist soil which is conducive for decomposition
thereby increasing soil fertility.
 There is reduced draft power requirement since soil will still be moist compared to working on dry
soil.
 Wear and tear of implements is reduced hence reduced cost of preparing the land.
 It allows spread of operations over the entire farming season or year.
 It enhances early plantings giving crops an early start when conditions are more suitable for plant
growth.
 Assists in weeds, pests and diseases control.
Late ploughing is normally done towards or during the planting time during the period September to
November. Large clods are normally brought up and these clods need to be broken up or else the
planting, germination and emergence of crops will be affected. Disease weed and pest load is usually
higher for crops planted in late ploughed lands. There is poor moisture conservation, ploughing costs
are high, wear and tear of implements is high and labour bottlenecks are common.

Tillage methods
Tillage methods are large divided into two namely conventional and conservation methods.

 Conventional tillage: This involves a series of tillage operations done to achieve a desired soil
tilth, for example, rip, plough and harrow. There are two tillage categories namely primary and
secondary which are determined by the target tillage depth, implement used and goal of tillage.
Primary tillage is tillage which cuts, inverts or shatters the soil to a depth of 15 to 36cm. It usually
leaves the soil rough. The implements used tend to be heavier and stronger than secondary tillage. his
can be immediately after the crop harvest or at the beginning of the next wet season. When there is
sufficient power available some soil types are ploughed dry.
The objectives of primary tillage are:
 to attain a reasonable depth (10-15 cm) of soft soil with varying clod sizes;
 kill weeds by burying or cutting and exposing the roots
 soil aeration and water accumulation; depending on the soil type and the plough the soil
will normally be inverted aerating the deep layers and trapping water during a rainfall
event
 chop and incorporate crop residues.
The implement most commonly used with an animal powered system is the moldboard plows. In
clay soils, the fields often have to be fully saturated before tillage can be undertaken. In lighter
texture soils such as loam or sand, tillage can be undertaken at moisture levels below field capacity.
In 2-wheel powered systems both moldboard and disc ploughs (one-way disc)are used. The disc is
usually the preferred system as it takes less power and can handle obstacles much easier.
In a 4-wheel tractor system, three-disc, seven-disc and offset ploughs are the most common. Tined
plows are preferable in the upland systems
These include rippers, subsoilers, mouldboard and disc ploughs and tandem disc harrows.
The aims of primary tillage include loosening and aerating the soil, covering plant residues, and
incorporating fertilizers and lime.
Disc plough (action):
Action is similar to that of the mould board plough and the soil inversion is only partial. It can be used in
situations where the mouldboard may not function well properly/satisfactorily e.g.
i ) hard and dry soil ii ) sticky soils iii) highly abrasive soils
210
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

iv) hard pan soils v) new ground with remaining stumps, roots etc.
Chisel plough:
It does not plough. The desired action of the chisel plough is to break and shutter the soil while leaving
enough residue on the surface to help control soil erosion.
It is a heavy cultivating implement
It is also used to breakdown plough pans or plough soles caused by mouldboard ploughing at the same depth
for years
Useful in hard conditions and in reclaiming rough lands and orchards
It is heavy structured
Stubble mulch tiller
Combines the operations of chiselling and disking. It consists of disk blades and in front to cut and loosen
trash
and a chisel plough in the rear section to break and shatter the soil
Powered rotary tiller
Require engine power to rotate the blades for cutting/lifting and loosening the soil. Depth of cut is up to 12-15
cm.
Effective in chopping and incorporating trash and preparing a fine tilth bed. They are often used for secondary
tillage at shallower depths
Note: excessive use may lead to soil pulverization- limiting water infiltration and increasing the erosion
hazard.
Rotary tillers are suitable for light soils

Secondary tillage is the operations that follow primary tillage in an effort to achieve a fine tilth. It
works the soil to shallower depth of 5-15cm. Secondary tillage is any working completed after
primary tillage and is undertaken for
 reducing clod size,
 weed control,
 incorporation of fertilizers,
 puddling and
 leveling soil surface.

Secondary workings are usually shallower and less aggressive than primary tillage. In the animal
powered system, the second working is normally undertaken with the moldboard plough when the
field is fully saturated. The final workings are then completed using peg tooth harrows to puddle the
soil and leave the surface level and ready for planting.
In 2-wheel tractor systems, the moldboard, the disc and the rotovator are used for second working.
In some instances peg tooth harrows are also used if rotovators are not available. Cage wheels on the
tractor are needed for traction in all soil types and these also help puddle the soil.
In 4-wheel tractor systems, tined cultivators, seven-disc ploughs, offset disc ploughs and rotovators
are the most commonly used equipment for second workings. In this system, fields are either
mechanically puddled with tractors using a rotovator and leveling board or by tractors fitted with
large cage wheels and harrows.

The implements used include disc harrow, spike toothed harrow, roller, rigid cultivator and spring
loaded cultivator.

The aims of secondary tillage include the attainment of fine tilth, weed control, firming the soil and
levelling the land.
[Link] harrow
o e.g. tandem disk harrow(can be used for primary tillage depending on size)
211
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

o Action: pulverize soil clods, levels and firms soil

2) Field cultivators (usually have conditioners).
─ Resembles chisel plough except that it is lighter constructed and designed for shallower tillage.
─ They are widely used for seedbed preparation and weed control.
3) Spring, spike and tine-tooth harrows
─ These implements vary in design but they have similar purpose of levelling, pulverising,
firming soil and sometimes weed control. They may be operated separately or pulled behind
ploughs, disc harrows or field cultivators
4) Roller packers and roller harrows?
Can be made from stone, iron or wood
Implement consists of heavy rollers designed to crush soil clods and the soil surface- good for small-seeded
grasses and legumes
General principle involved is the rotation of massive material round the axle
It should not be used on wet soils otherwise severe crusting results
5) Row-crop cultivators

 Conservation tillage
It is a system of farming which involves managing the land with principles of conservation. Such a
system would include contour farming, crop and pasture rotation, and soil and water conservation.
It also involves any practice which leaves at least 30% crop residue cover on the soil surface after
planting.

Advantages of conservation tillage

 It reduces soil erosion as the crop residues cushion the rain drop action.
 There is maximum use of available moisture as water run-off and evaporation are reduced thereby
improving penetration.
 There is moderation of soil temperature extremes by crop residues.
 Reduced tractor operating costs as less fuel and labour will be used, and there is reduced wear and
tear of implements.
 It enhances timely operations since the time consuming operations can be avoided.
 It improves soil structure since the built up organic matter tends to bind soil particles together and
compaction of the soil is minimised. Minimum soil disturbance results in less pulverisation of the
soil.

Disadvantages of conservation tillage.

 There is built up of pests and diseases.

 There is poor incorporation of trash and as a result the decomposition process is slowed down
which may result in high C:N ratio that affects nutrient use by plants.
 Nutrients tend to be concentrated at the top layers of soil. Roots will not go deeper hence lodging
is common and crops are susceptible to droughts. There is difficulty in monitoring the levels of
phosphorus and soil reactions.
 Requirements of conservation tillage
Weed pressure: It is advisable to start with a lower weed pressure. Control late weeds in the
previous crop to reduce the weed seed bank for the next crop.
Soil fertility: Manage the soil fertility by ensuring that the soil has sufficient phosphorus and the
pH is well adjusted at the onset. The pH should be 5.5 and phosphorus level should be 25ppm
(25mg/kg of soil)
Compaction: Dig inspection pits and correct compact layers by ripping and sub-soiling.
Otherwise there could be problems of root penetration and water movement in the soil.
212
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Types of conservation tillage

 Strip tillage: It is performed through or into crop residue. A line into which seed is planted is
opened up. The most common practice is rip-on row which ensures loosening the soil to a depth of
25-30 cm. Fertilizer is banded into the rip line in the same operation. Planting may be done by
hand or machine.
 Direct planting: Involves planting into previous crop residues without prior tillage operation
 Zero tillage: Involves the broadcasting of seed usually cereals over previous crop residue.
 No- till tied ridges: Planting is done on tied ridges year after year. This method conserves
moisture and soil.

Summary
Land selection and preparation are important aspects in crop production. The factors considered in
land selection include soil fertility, soil depth, soil texture and structure, soil permeability, salinity and
pH as well as the slope of the land, erosion and wetness. Depending on available equipment and other
resources as well as the crop to be established the farmer has a choice of doing conventional or
conservation tillage. Every effort should be made to get high crop yields using the most sustainable
means of production especially conserving the soil.
Activities
a) State and explain the factors considered when selecting land for the cultivation of crops.
b) Suggest the reasons why farmers should take-up conservation tillage at the expense of
conventional tillage.
c) List the challenges likely to be met in practicing conventional tillage.
References

 Arnon I (1972) Crop Production in Dry Regions, Volume 1, Leonard Hill, London.
 Chapman S.R and Carter L .P. (1976) Crop Production Principles and practices. Freeman and
Company, USA.

CHAPTER 28

PLANTING

Introduction
Planting is the placement of propagative material (seed/cuttings or suckers) into the rooting media
(soil) to enhance germination and emergence. This is a very important part of crop production. It is
important to note that for high yields to be obtained, the farmer must plant at the right time, using the
correct plant population, depth and method, and applying the correct fertilizer type and rate as well as
using the right type of seed. In this chapter, we are going to discuss these important aspects of
planting.

Objectives
By the end of this chapter, you should be able to:

 State the importance of timely planting.

 Highlight the importance of achieving an ideal plant population.

213
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Discuss the various methods of planting.

Time of planting
Planting should be done when the environment is ideal in terms of such factors as temperature,
moisture and day length. Time the planting so that the crop will be ready when the conditions will be
conducive for ripening or harvesting, for example, soyabean should not be ready for harvest when it is
still raining. If it does the seed will germinate or may rot on the mother plant. For most plants of
tropical origin, one must avoid frosty periods because crops easily suffer from frost damage and
chilling injury. Other factors that determine the planting time are pest and diseases incidences, market
prices, weeds pressure and the onset of rain for summer crops. For summer crops, it is ideal to do early
planting (October- November) depending on crop and the Natural Region where production is done.
The use of irrigation can advance the planting time prior the fall of the summer rains. For most
summer crops there is yield decrease with late plantings. For most summer crops, plant early when the
weeds, pests and disease pressure is still low. If it is dryland production, the earliest you can plant
crops like soyabean is when you have receive the first effective rainfall.

Plant population
Plant population is the total number of plants per given area. It is generally expressed as plants per
hectare (plants/ha).
Factors that determine the plant population to use include:

 Type of crop and variety, for example in groundnuts the population is high when growing the
bunch type than the spreading type.
 The level of management, for example a maize crop under irrigation requires a higher plant
population compared to dryland cropping.
 Type of season, for example a predicted low rainfall season would require lower plant population
so that plants will not compete for the limited moisture available.
 The region in which the crop is grown. Low rainfall regions e.g. Region III and IV one to use low
plant populations.
 The intended use of the crop, e.g. silage maize crop should be grown at a higher plant population
compared to commercial maize.
 Time of planting. Reduce the population with delayed plantings and
 Expected yield. Go for optimum population for high yields.
 Biological factors

 Interspecific competition is a major limiting factor on population size. Seral stage plants can often
grow very well in the conditions afforded during a succession climax - they are just out competed
by the climax community, whose branches reach higher to trap the light and whose roots reach
lower to gain the nutrients and water
 Intraspecific competition can limit growth - which is a reason why many plants develop seed
dispersal mechanisms based on the wind, insects or animals. Intraspecific competition is density
dependent - the more offspring there are, the more limiting the competition becomes.
 Herbivores may reduce a plant population - this is particularly true if an invertebrate herbivore
undergoes a population boom
 Disease can also cut a population - particularly if spread by an insect
 Introduced (alien) species can undergo uncontrolled population growth

214
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

For crops in rows, plant population is calculated as follows:

Plant population = 10 000 m2
Inter-row spacing (m) X In-row spacing (m)
For example: A maize crop planted spaced at 900mm X 200mm
Plant population = 10 000m2
(0,9 X 0.2)m2
= 55 555plants/ha
For crops in twin rows,
Plant population = 10 000 m2
Average inter-row spacing (m) X In-row spacing (m)
For example tobacco spaced at 1.2m x o.56m and 1.1m x0.56m
Plant population = 10 000m2
(1.15 x 0.56)m2
= 15 527plants/ha
For crops with no planting pattern, a quadrant of known dimensions is used. Choose a minimum of
three sites at random and calculate the average plants per site per quadrant.
For example: In wheat, three sites had the following plants per quadrant which is 1m2: 10; 15and 8
Therefore, plant population = 10 000 m2 X 1 m2
(10 + 15 + 8)/3
For plants on hills, sample 10 hills at random. Average the plants per hill, and calculate plant
population as for single rows.
For example, maize planted 3kernels per station spaced at 0.9m 0.3m,
Plant population = 10 000 x 3
0.9 x 0.3
= 111 111plants/ha
Advantages of an optimum plant population per hectare.

 To get optimum yields due to low competition among crops for available resources and land not
wasted.
 To get high quality produce from a crop filly benefited from resources available.
 To provide good soil cover especially in row crops like tobacco and cotton.
 To reduce disease prevalence. This will be achieve through reduce transfer of pathogens from
plant to plant and also through good ventilation within the field.

215
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 To help in the control of weeds especially in wheat and barley crops. Use the recommended seed
rate per hectare to achieve the desired plant population.
 To enhance efficiency execution of other operations like spraying and movement of implements
especially in crops like cotton and tobacco. Adhere to the correct inter-row spacing.
IMPLICATION OF PLANT POPULATION
Larger populations of plants are likely to be more attractive to pollinators resulting in higher visitation
rates and therefore pollination success (Sih & Baltus 1987; Ågren 1996), whereas small populations
may suffer from insufficient pollen transfer and consequently lower seed set (e.g. Jennersten
1988a; Lamont et al. 1993; Ågren 1996; Fischer & Matthies 1998).
In addition, the level of inbreeding may be higher in small, isolated populations (e.g. Barrett & Kohn
1991; Falconer & Mackay 1996) because of the higher rate of selfing and more frequent matings
between close relatives. The resulting inbreeding depression can reduce the fitness of these plants
compared with those in larger populations (Menges 1991; Aizen & Feinsinger 1994a; Ouborg & Van
Treuren 1994; Heschel & Paige 1995).
High population density (32 plants/m2) planting of soybeans [Glycine max (L.) Merrill] resulted in
small plants but high dry weights per unit area of ground, while low population density (four
plants/m2) produced larger plants with a smaller dry weight per unit area of ground. Intermediate
population densities (16 and 8 plants/m2) produced intermediate plant sizes and yields. Shoot/root,
bean/shoot, and leaf area ratio were all decreased by increasing population density..
Conditions necessary to achieve optimum plant population

 Spacing: use the recommended spacing and seed rates.

 Planting depth: plant seed at the correct planting depth.
 Good seed soil contact, to ensure the seed will absorb moisture from the soil.

Types of planting

 Dry planting: is the planting of seed well in advance of the planting season. Land is fully prepared
and seed planted 2-3 weeks prior the coming of the first rain to ensure an early start to the season
and to release labour for other farm operations when the rains start. The seed must be planted in
dry soil. The dangers of dry planting include the swelling of seed and death should there be
showers before the first effective rainfall and also pest damage. The planting depth will depend
on seed size and crop.
 Water planting is a method of lengthening the growing season especially in sandy soils. The aim is
the establishment of the crop early before the first effective rains. Early plough the land and leave
it rough to facilitate water intake. Destroy all weeds activated by the last rains. Supply adequate
planting water to link with the residual soil moisture. Allow the planting water to sock away
before covering with dry soil which will act as mulch. Preferably plant socked seed.
 Rain planting: is planting after receiving good effective rains.

Planting methods

 Hand method involves the placement of seed in prepared planting holes, furrows or land in the
case of broadcasted small grains.
Its advantages include correct placement of seed, good assurance of seed placement as there are no
blockages of seed outlets, simple and easy to implement, no restrictions to seed type and size and
is possible under varied soil conditions and land forms.

216
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

However, hand method require more labour force, strict supervision on seeding and covering and
best suits small areas.
 Mechanical planting is the use of machinery in the planting operation. Equipment commonly used
include, planters, seed drills and vicons.
Mechanical method is fast and best suits large areas. However, it tends to be affected by weather
conditions and soil conditions especially in wet soils they may skid and clog.
 A combination of the hand and mechanical, for example, furrows and fertilizer application can be
opened mechanically and the planting is down by hand.
Summary
Planting is the placement of seed in a rooting media to enhance germination and emergence. The time
of planting is determined by the climate, pest and disease incidences, weeds pressure and market
prices. It is important to achieve a desirable plant population for every crop grown. The best possible
method of planting should be opted for depending on available resources and the type of planting
pursued.

Activities
a) Outline the factors that determine a plant population to adopt.
b) Calculate the plant population of soyabeans planted at 450mm x 75mm.
c) Compare and contrast hand and mechanical methods of planting.
d) Describe the method of water planting crops.

References

 Arnon I (1972) Crop Production in Dry Regions, Volume 1, Leonard Hill, London.
 Commercial Grain Producers Association (CGPA) ,(1996) Grain Handbook, Screenlitho (Pvt) Ltd,
Harare , Zimbabwe.
 Seed Co (2002) , Agronomy Manual, Seed Co Ltd, Harare, Zimbabwe.

ROTATIONS

Crop rotation is a cropping system in which two or more crops are grown in affixed sequence on the
same field. Alternatively, it is the growing of different crops in a more or less definite sequence on the
same field. The rotation desired is the one that gives the greatest net return over a period of time. The
most widely recommended rotations consist of a tilled crop, a small grain and a legume or grass crop
in that order so as to get the benefits of crop rotation to be discuss in this unit.
Basic principles of developing a crop rotation.
The following are considered when developing a crop rotation:

 Select crops well suited to the soil, topography of land and climate.
 Include legumes to maintain or increase nitrogen and organic matter of the soil especially on land
grown row crops such as potatoes, tobacco, maize and cotton.
 Select crop sequences which produce the highest yields and provide the best control of weeds,
insects and diseases.
 Select crops that provide sufficient feed for livestock on the farm.
 Select crops that provide good distribution for labour.

217
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Grow a maximum area of the most profitable crops, provided they do not sacrifice the future
productivity of the soil. The area of the crop should be nearly the same year after year unless need
for change arises.
 Other factors include soil conservation, goals and managerial ability of the farmer, and capital
available as well as effective use of the nutrients in the soil profile.
 Duration of the rotation normally varies from two to four years.

Types of rotations.

 Grass ley rotation: Is a rotation which includes a period in grass which is used for grazing and
conservation.
 Continuous cereal and break crops. For example; maize- soya bean – maize, cotton – maize-
cotton, tobacco – maize- maize, maize- wheat- soya bean maize, tobacco- grass- grass- grass-
tobacco.

Benefits of crop rotations.

 Increased yield from conserved soil and improved fertility. The supply of nitrogen is increased by
the inclusion of legumes.
 The supply of organic matter is maintained or increased which is important in maintaining the
physical properties of the soil, for example water holding capacity and soil texture. The supply of
organic matter also encourages growth and development of beneficial bacteria in the soil.
 Soil nutrients are utilized more efficiently. Various crops differ in nutrient absorption and also take
up nutrients at different rooting depth.
 Erosion losses are reduced. Various crops differ in protecting soil from wind or water erosion. The
amount of nutrients removed in cropping is often small compared to that lost by erosion. Legumes
and grass are highly effective in reducing soil and nutrient losses by erosion.
 Manure and commercial fertilizers are more effectively utilized. These could be applied to high
value crops or crops that respond best to such treatments.
 Weeds, pests and diseases are better controlled. Rotation of crops will call for the use of different
herbicides to control weeds, and pesticides to control pest there by reducing resistance built-up in
pest. Also the inclusion of different plants will reduce the seed bank and pest population or
diseases as they are crop specific.
 Labour is more uniformly distributed throughout the year as the cropping will be done throughout
the year.
 Keeps the land occupied a greater part of the year with crops as one crop will be planted soon after
the other crop.
 Hazards are reduced. The risk of complete failure or poor production is usually greater with one
crop than when several crops are grown.
 Improves crop quality through timely operations like planting, reduced weed challenge and also
reduced pest and disease damage.
 Systematizes farming. The labour force would be aware of the crop to follow after harvesting
hence preparations for the next crop will be done timely. The structures established at the farm
will be used more efficiently.
 Saves on labour by growing two or more crops on one soil preparation where conservation
farming will be practiced.
Disadvantages of crop rotation.

 It requires diverse knowledge for management of different crops.

 Requires more capital investment as same crops do not share the same equipment or structures.
218
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Crop relationship in a rotation.

When crops are grown in a rotation, they can have at least three relationships as follows:
 Competitive: Two crops are said to be competitive when one crop is grown at the expense of the
other.
 Complimentary: is when an increase in total output of one crop results in an increase in total
output of another crop from a fixed area or given resource.
 Supplementary: is when the crops neither add nor subtract from the output of each other.
Factor affecting crop rotation
 Climate
Climate is the one of most important factor which is effect the crop rotation either by wind, rain or
other factors.

 Type and nature of soil

Type and nature of soil is also important factor which effects the crop rotation some soil are fertile and
some are low in fertility

 Availability of inputs
Availability of inputs at the place is also effects the crop rotation like fertilizer, pesticide etc

 Availability of labor
Availability of labor is effect the crop rotation. The labor is required at the critical stages of crop if the
labor is not available at that time the crop may cause loss

 Situation of farm
The farm location is also very important factor which is effect the crop rotation.

 Size of Farm
The size of farm is effects the crop rotation. Small land holding is major problem in Pakistan that’s
why crop rotation is effect by the farm size

 Type of farming
Type of farming is also effect the crop rotation

Summary.
Cropping systems include monoculture, intercropping, fallow system as well as rotation. Each system
has its own advantages and disadvantages. However all systems target have the highest possible yields
in crop production in a sustainable way.
Activities
a) Explain the disadvantages of monoculture.
b) Discuss the effects of practising intercropping in commercial farms.
c) Outline ten reasons for practising crop rotations.
References
 Arnon I (1972) Crop Production in Dry Regions, Volume 1, Leonard Hill, London.
 Chapman S.R and Carter L .P. (1976) Crop Production Principles and practices. Freeman and
Company, USA.

219
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

CHAPTER 29

MAIZE

Common name: Maize

Scientific name: Zea mays L.
Shona: Chibage/Magwere
Ndebele: Umumbu
Rainfall
Favours long seasons (110-140 days) in which there is adequate moisture.
Severe moisture deficit at tasseling and silking causes crop failure due to poor pollination.
If rainfall is inadequate during taselling/silking, it is advisable to irrigate the crop.
Water logging also has negative effects on maize.
Temperature
Mostly night temperatures tend to affect maize yields.
Temperature also affects time taken to maturity. Lowveld area crops tend to mature much faster due to
high photosynthesis.
Temperatures above 38ºC destroy pollen viability.
Soil types
Sandy clay loams and heavier soils are most suitable.
Soils should be well drained and the optimum pH range is 5.5-6.5
On heavy soils pH should not be allowed to drop below 5.0.
Land Preparation
Winter ploughing, immediately after harvest kills weeds, conserves moisture, controls pests and
promotes early decomposition of weeds and residues.
Summer ploughing and harrowing will create a fine tilth for planting.
Cultivar selection
There are two major groups of cultivars:
Hybrids whose seeds cannot be recycled and
Open-pollinated varieties, whose seed can be recycled for at least 3 seasons before buying fresh seed,
Some of the advantages of OPVs
Cheaper seed
Seed can be recycled for ( 2-3 season )
Recycling hybrid seed result in loss of hybrid vigour, hence the need to buy seed every season.

Long season variety: Matures in 140+ days, e.g. SR52, SC701

Intermediate season variety: 130-140 days, e.g. SC627, ZS255
Short season variety: 120 -130 days, e.g. SC407, ZS257, SC403, SC401
Long season and medium season varieties are recommended for high rainfall areas while short season
varieties are for drier regions.
Seed rates
On average 25kg/ha of seed maize is recommended.
Planting time:
Dry planting can be done 2-3 weeks prior to the first effective rain or alternatively plant with first
effective rains.
Long season varieties can be planted with irrigation (where it is available) in mid October before the
rain.
Late planting of maize after mid-December reduces yields.
Rotation can be done with legumes.
Plant population and spacing

220
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

45 000 plants/ha ( 90 cm x 25 cm) for high rainfall areas of NRI and NRII and
37 000 plants/ha ( 90 cm x 30 cm) for low rainfall areas of NRIII and IV
22000 plant/ha (1.5m x 30 cm) on tied ridges for dry areas of NRV.
Varieties with short stature can be planted at higher plant populations.
The general Planting depth is 50 mm, but variations can occur depending on soil type and planting
method.
Fertilizer application rates and time:
Table showing general Fertilizer Recommendation according to soil fertility status
Soil Fertility Status
Good Medium Poor
Fertilizer Nutrients Kg/ha of fertilizer nutrient required

N Up to 100 100-160 160-200

P2O5 30-50 50-70 70-90
K2O 20-30 30-50 50-70
Fertilizer in bags 3-4 compound D 4-7 compound D 7-10 compound D

Table 1: General recommended fertilizer rates according to Natural Region

Fertilizer Natural II Natural III Natural IV
Compound/maize fertilizer (kg/ha) 300 to 350 200 to 300 150-200
(6-7 bags) (4-6 bags) (3-4 bags)

Ammonium nitrate (AN) top dressing (kg/ha) 250 to 300 150 to 200 100-150
(5-6 bags) (3-4 bags) (2-3 bags)

Generally, the whole of P & K requirements and at least 1/3 N should be applied at planting by either
broadcasting or banding in planting rows.
The top dressing fertilizer (AN) is recommended at 4-6 weeks after planting or at maize knee height if
applied once.
If topdressing is split because of high rainfall, the first 1/3 can be applied at 4-6 weeks and the last 1/3
at 8 weeks after maize planting.
Liming
It is recommended that fields be limed once in three years.
General recommendation is 600-kg/ha limes.
If the soil is strongly acidic comparatively large applications of lime are required, ½ to 2/3 of the
recommended lime should be applied in the first season and the remainder in the following season.
It is important that the lime should be incorporated through out the plough Zone (30 cm). It is
recommended also that it be applied after winter ploughing to give it time to react with soils before
planting.
NB manure & ash have liming effects on soils
Other soil nutrient replenishing methods to supplement inorganic fertilizers:
Use of cattle manure, cured in pits or heaped.
Rotation mainly with grain legumes such as groundnut, Bambara groundnut (Nyimo /indhlubu) and
soyabean.
Maize intercropping with legumes such as cowpea (Nyemba / indumba)
Green manuring using legumes such as velvet bean and sun hemp
Use of agro forestry species such as e.g. Sesbania sesban for improving fallows before planting maize.
Use of termitaria (termite mound) soils
Use of compost and ashes in fields to be planted to maize
Weed management
221
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

A maize crop should be maintained weed-free for at least 2 to 6 weeks after establishment.
There are basically three types of weed control practices used in Zimbabwe
Hand weeding, using a hoe
Mechanical (using ploughs and cultivators) in combination with hand hoeing
Use of herbicides. Atrazine is the most common herbicide recommended for use in maize. It can be
combined with mechanical weeding by applying a rate reduced by 50% in the crop rows only and
using the cultivator or plough in between the maize rows. Atrazine controls most broad leaf weeds,
but depending on the weed species, other herbicides may be recommended.
Pests and diseases
Table 2: Major maize diseases and insect pests and their cultural or chemical control
Diseases Cultural and/or chemical control
Leaf blight Breeding for resistance and use of rotations
Leaf streak virus 40% Dimethoate at 50ml in 455 litres of water
Boil smut Standard seed dressing eradicates the seed borne infection
Bacterial stalk rot Do not irrigate in hot, sunny conditions
Maize rust Breeding for resistance
Grey leaf spot Breeding for resistance

Chemical control:
Pests Active ingredient Trade name
Aphids
Demeton-S-Metyhyl Metasystox 25 EC
Dimethoate Dimethoate 40 EC, Rogor CE
Thiometon Ekatin 25 EC
Carbaryl
Armyworm Malathion Carbaryl 85 WP
Malathion Dust, Malathion 50 EC
Benfuracarb Oncol 20 EC
Carbofuran Curater 10 G, Carbofuran
Leaf hopper 10 G
Imidacloprid Gaucho 70 WP
Dimethoate Rogor CE, Dimethoate 40 EC

Stalk-borer Carbaryl
Carbaryl 5 Dust, Carbaryl 85 WP
Trichlofin Dipterex 2.5 Gran
Endosulfan Thiodan 1%

Fipronil Regent 250 SE, Regent 3 G

Imidacloprid Confidor 200 SL
Termites
Actellic Super
Pirimiphos methyl + Permethrin
Weevils Malathion Ingwe/Nhoro
Pirimiphos methyl Shumba 2 Dust

Harvesting, drying and storage

Harvesting indices (signs of maturity)
- Black layer at the top of mature kernel ,
- Grain moisture at onset of maturity occurs within the range of 25-350C,
222
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

- Number of days to reach maturity of the variety being grown ,

- Drying of maize cobs ,
- Kernels become hard to bite and glossing in color,
- Husks and leaves become brown and papery.
Harvesting methods
- Harvestingis the operations of gathering the useful part or parts of the plant.
- It is a voluntary intervention by man, carried out at the time when all the nutrients have been
developed and the edible parts have reached maturity.
- Maize crops are first cut either as whole or partially (cobs) and then threshold and cleaned to
separate the grain from the cobs and straw.
Maize can either be dried artificially or naturally.
In natural drying, the cob is left on the plant and on small plots the sheath could be opened to speed
up drying.
Acceptable moisture for effective storage is 12.5%.
Marketing
Marketing of maize grain is currently controlled by Legislation with the Grain Marketing Board being
the sole trader.
Innovative utilization at farm level i.e. for stock feed
Maize storage
- Maize for marketing should be stored shelled in a closed stored with the use of pesticides.
- The spread of the grain borer in the region makes the use of good storage essential.
- Delayed harvesting of maize lead to higher losses because of attacks by termites , rodents and
domestic animals.
- Crops can be infected by pests.
- Storage facilities should serve the following purposes
× provide protection –ground & water rain
-insect pests
-excessive heat.
Maize storage facilities
- Drying storage cribs – used normally when storing maize on the cobs, and can also be used to store
shelled grain in bags.
- Mud or cement – plastered buckets.
- Brick bins -
- Ferro cement bins
Basic components of a storage structure:

surface water.) and to provide a barrier against pests. In Zimbabwe a foundation consisting of a few
widely spaced stones with a platform of logs is very common. (advantages and disadvantages).
Foundation should always raise the floor of the structure to prevent surface water from entering the
store- a minimum of 20 cm above the ground is often recommended.

izontal pressure from the stored commodity; to protect

the commodity against temperature fluctuations; to protect the commodity against moisture; to keep
pests out.
entry of
pests.
Thatch is a very old and very suitable type of roof.

resist lateral pressure of the stored produce.

223
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

ws and compartment shutters): to seal the store or compartments

and still provide access to the produce, provide aeration or entry of light.
Types of storage
a) Bulk storage:
Very common type in Zimbabwe and the structures in which the grains are stored are known by
different local names (hozi, dura etc).
b) Bag storage bags to be raised from the ground by placing them on a platform
- should be stacked in such a way that the stack is stable and there is good air circulation
- Rodents can easily hide in stacks, so it is advisable to have the bags on raised platform as well
having them a distance from the walls. This also prevents moisture uptake though the walls and the
floor and allows cleaning and better inspection of the store.
THE IMPORTANCE OF GOOD STORAGE TECHNIQUES/ THE RATIONALE FOR
STORING GRAIN.
- Food security reasons: ( for future use).
a) To keep grain to use as planting seed in subsequent years.
b) For food and feed.
─ Allows supply of food from one harvest to the other
─ The stored grain will cushion the farmer in bad seasons. It protects consumers
─ against bad crop years.
In the smallholder farming sector, a large proportion of the grain is retained.
On average an adult requires about 155kg of grain per year.
- Grain can be stored and only sold when good prices are being offered.
Preparing Maize for marketing
- GMB buys the maize soon after harvesting.
- The farmer can find milling buyers at a Marketing Board or Co-operative Depot.
- Maize is marketed through the GMB at a price gazette by the government.
- Also due to trade liberalization, maize processing companies like millers and breweries provide an
alternative for maize growths.
Moisture Content determination
1. Subjective methods of determining MC in grain
Most farmers use these methods to get a rough idea of the MC.
– dry grain is harder
– dry grain is harder
–wet grain gives a dull sound
–wet grain gives a dull sound

A more accurate method is the salt test: (but method is also subjective).
A jar is half-filled with grain and a tablespoon of common salt is added to it. The jar is then shaken for
a few minutes to bring the mixture to equilibrium. If the salt sticks to the walls of the jar, then the
grain had a moisture content of more than 15% (15% is the equilibrium grain moisture with an air
relative humidity of 70%) If the salt still looks dry (does not stick) the moisture was less than 15%.
- Farmers can market their grain through the open market like Mbare Musika at concessionary prices.

CHAPTER 30

SOYABEAN PRODUCTION

Soils and climate

 The soyabean is suited to soils with high clay content. The optimum pH is 5.5

224
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 .Soyabean requires reliable rainfall particularly from flowering to pod maturity.

 It is a good crop to grow in rotation with maize, cotton and wheat.
Fertilisation

 Soyabeans grow well on residual fertiliser.\

 General recommendation is 200 - 300 kg/ha of either a Compound fertiliser (D or L), Gypsum or
Single Super Phosphate before planting.
 Soyabeans respond well to manure application.
 Soyabeans do not need much nitrogen either in the basal fertilizer or as top dressing they are able
to obtain their nitrogen requirements from the soil air.
 It is therefore essential to apply Bradyrhizobium inoculant to the seed at planting.
 Do not leave the inoculant in a hot place, but store it in a cool, dark place.
 When ready to plant, the inoculum is mixed with a little water and sugar, and applied to the seed
immediately prior to planting.
 One packet of inoculant is required for each 100 kg of seed. However, a higher rate is preferable
on sandy soils or in first year soyabeans fields.
Cultivars
There are two basic types of soyabean cultivars: Determinate and indeterminate

 Determinate cultivars grow vegetatively for about six weeks and then begin flowering, having put
on 10 to 12 leaves. Once flowering begins, no further new leaves are produced on the main stem.
 Indeterminate cultivars, on the other hand, grow vegetatively for about six weeks, then begin
flowering when the main stem has about 10 leaves, but at the same time as flowering, the stem
continues to grow for another three weeks or so, producing another five to seven leaves.
 Determinate cultivars are better suited to warm fast growing environments where irrigation is
available, like the lowveld, whilst on the middleveld and highveld, both types are suitable. Under
drought conditions, indeterminate varieties may have some advantage over determinates.
 Some of the cultivars available include Soprano and Solitaire, which are indeterminate, and Storm
and Soma, which are determinate.
Planting and crop management

 The seed rate is about 100 kg per ha.

 The desirable plant population is around 350 000 plants per ha. The minimum plant population is
200 000 plants per ha, while the maximum is 500 000 plants/ha.
 The row spacing may be from 25 - 90 cm.
 Do not plant seed deeper than 5 cm.
 Covered the seed such that the soil forms a slight mound over the row, as this makes it easier for
the seedling to emerge.
 Seed should be planted 25-50 mm deep, depending on soil texture. If soil crusting occurs before
emergence, wetting the soil with irrigation or breaking the crust with a ‘millipede implement’ will
improve emergence.
 Planting is ideal around mid-December.
 A fungicide seed dressing of Thiram 80 WP (85 g/50 kg seed) or Captan 50 WP (125 g/50 kg
seed) will help ensure good emergence.
 Soyabeans are particularly sensitive to weed competition during the first six weeks of the season.
 Soyabeans are very susceptible to drought during the pod-filling stage.
Soyabean Planting Guide

225
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Required average number of seeds

Seeding rate (kg/ha)
per metre of row
Target Required
Harvest Planting
Thousand seed weight
Population Population Row width (m)
(g)
per ha per ha
0.45 0.60 0.75 0.90 250 220 190

200000 248000 11 15 19 22 62 54 47

240000 297000 13 18 22 27 74 65 57

280000 347000 16 21 26 31 87 76 66

320000 396000 18 24 30 36 100 87 75

360000 446000 20 27 33 40 111 98 85

400000 495000 22 30 37 45 124 109 94

440000 545000 25 33 41 49 136 120 104

Soyabean Management Guide

Establishment Late vegetative, early Seed filling

flowering period

Lime acid soils (Ideal pH=5.3). Control any new weed growth. Ensure good control of Rust and
Fertilise as necessary. Start scouting for Semi-loopers Semi-loopers.
Best variety. and Rust. Apply two fungicide sprays
Use certified seed. Apply first fungicide spray for during this period.
Inoculate with Rhizobium. Rust at first flower. Irrigate when there is a dry spell
Do not plant too deep. longer than 5 days.
Population: 300,000 plants/ha
Row width 45 - 75 cm. Harvest early to avoid shattering
Ensure early weed control. losses

Pests
Semi-looper caterpillars

 Often a problem during the flowering and seed-fill period.

 They eat the leaves and sometimes the pods.
226
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 Normally controlled by a naturally occurring virus disease, which kills the caterpillars.
Caterpillars that have died from the disease are black and hang from the leaves. These may be
collected, crushed, mixed with water and sprayed around the field to help control other
caterpillars.
 If the virus is not killing the semi-loopers then an insecticide spray may be required.
Diseases
Frog Eye Leaf Spot (Cercospora sojina)
Most cultivars available today are resistant to this disease.

Rust (Phakopsora pachyrhizi).

o The Rust disease may be recognised as numerous small grey to russet coloured tufts on the
underside of leaves. They may appear similar to red spider mites.
o In advanced stages the leaves become distinctly yellowy-brown and a light brown cloud of
spores are seen in and above the canopy when the plants are shaken.
o It can be controlled using a fungicide spray.
o A number of chemicals are available, such as Punch® Xtra (350 to 500 mL/ha) and Shavit®
(500 ml/ha). Two or even three, fungicide sprays are at 3 week intervals beginning at first
flower. Ensure good leaf cover when spraying.
Red Leaf Blotch (Pyrenochaeta glycines).
o Severe infections can reduce seed yields by 30% - 50%. D.
o Solitaire and Soprano are moderately resistant.
o Chemical control is not available.
Harvesting
a) Hand harvesting.
o Suitable for small areas.
o Advantages of hand harvesting are that:
Losses can be reduced to a minimum
Soyabeans of a high quality are produced
The beans normally have a high viability.

 Therefore, hand harvesting is suitable for seed production.

b) Mowing or cutting by hand and shelling.

 A variation is to use a mower to cut the plant material, and a mechanical winnower for the
final cleaning.
 This method should enable an output of ± 150 kg (3 bags)/labour/day. This method enables
harvesting to commence before the pods split, but allows sufficient moisture to be lost, thereby
preventing mould developing in the established cocks or stacks.
c) Swather plus combine.

 This method involves the use of a swather to cut and wind-row the crop before it is combined.
A pick-up attachment is required to be fitted to the combine table.
d) Combine harvesting.

 Large areas are usually reaped by combine harvester and losses are inevitable.

227
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

 The degree of loss depends on the efficiency of the machine and operator, evenness of the
land, the height of the pods off the ground, lodging, the moisture content of the beans, and
weed control.

CHAPTER 31

CONSERVATION FARMING:PRINCIPLES AND PRACTICES

 Explain the importance of conservation farming

 Outline the components and practices of conservation farming
 Describe different conservation farming practices.

Conservation agriculture (CA) can be defined by a statement given by the Food and Agricultural
Organization of the United Nations as

“a concept for resource-saving agricultural crop production that strives to achieve acceptable profits
together with high and sustained production levels while concurrently conserving the environment”
(FAO 2007).

“an approach to managing agro-ecosystems for improved and sustained productivity, increased profits
and food security while preserving and enhancing the resource base and the environment”

CROP PRODUCTION IN ZIMBABWE

PRINCIPAL CONSTRAINTS TO EXPANDED CROP PRODUCTION

AVAILABILITY OF CREDIT FOR COMMERCIAL PRODUCERS

ey supply situation in all sectors of the economy has tightened. Many potential
horticultural projects have been suspended as there are limited funds available for short, medium and
long term financing.
evelopment Fund, it fails to match the nature
of investments undertaken in horticultural production.

they expect the situation to ease only very slowly as the effects of drought on agricultural borrowings
are felt.

rates of up to
350% per annum
edit on firm export orders
will partly address the constraints, but only of export growers.

AVAILABILITY OF CREDIT FOR SMALL SCALE/COMMUNAL PRODUCERS

financial systems that currently prevail tend to exclude the small scale farmers, particularly the
communal farmers

only institution lending to small scale communal farmers

INADEQUATE TRANSPORT FACILITIES

228
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

markets

export production

hain from the packshed to the oversees market if premium prices

are to be paid

- insufficient capacity
- irregular take offs and cancelled flights relatively high cost of freight
- insufficient direct flights to destinations
- inadequately equipped cargo handling facilities at the airport

- lack of suitably designed and configured rolling stock

- inadequate cold storage facilities at Beira
- insufficient refrigerated trucks

- limited availability of lorries

- relatively high cost of imported lorries and the limited availability of spare parts
- insufficient competition which results in high freight rates
- lack of specialised transport required for perishable horticultural products
3.4 LOCAL RESEARCH AND SERVICES
Most of research in Zimbabwe is handled by AREX and educational centres. There are four main
centres namely Horticultural Research Centre (Marondera), Nyanga Experimental Station, and
Subtropical Horticultural Research Station at Chiredzi and Coffee Research Centre at Chipinge.

sector
current shortage of trained horticultural staff
within
AREX severely affected quality of services and horticultural research in the country.

Africa and
Europe
There is need for an urgent address of the current problems in the research areas.

sible for
issuing phytosanitary certificates to exporters

- The present site is congested creating possibilities of contamination. The problem will be overcome
if the unit relocates to more spacious sites e.g. Henderson
- The centre relies mainly on physical testing and no chemical testing is conducted
- Plant inspectors rely too much on honest declaration by individuals. Routine monitoring is required.
The unit is understaffed and under funded
LABORATORY SERVICES FOR PATHOGEN IDENTIFICATION, SOIL AND LEAF
ANALYSIS

e.g. ZFC,
University of Zimbabwe and parastatals TRB and SIRDC also offer testing services
fail to access these facilities

229
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

EXTENSION SERVICES AND TRAINING

training programme encompasses on a wide range of agricultural enterprises and the extensionists may
lack basic information on commercial horticultural expertise

programmes
PACKAGING MATERIALS
owever, most of the special packaging is still
being imported, thus creating shortages

PROCESSING FACILITIES
make the export grade or is
specifically aimed at the local or export markets in a processed form (e.g. canned, freeze dried, frozen
and bottled)

horticultural sector remains sidelined from these facilities

facilities would allow the small/communal sector

MARKET INFORMATION
nd the Europe Africa Caribbean Pacific Liason Committee
(COLEACP) provide information to Government bodies and exporters in African, Carribean and
Pacific
(ACP) countries on a regular basis
farmers in communal areas, intending to
venture into export.

Umlimi wa lamhla
and Murimi waNhasi television programmes.

IMPORTANCE OF CONSERVATION FARMING

Specifically, conservation agriculture (CA) increases the productivity of:

Land - Conservation agriculture improves soil structure and protects the soil against erosion and
nutrient losses by maintaining a permanent soil cover and minimizing soil disturbance. Furthermore,
CA practices enhance soil organic matter (SOM) levels and nutrient availability by utilizing the
previous crop residues or growing green manure/ cover crops (GMCC's) and keeping these residues as
a surface mulch rather than burning. Thus, arable land under CA is more productive for much longer
periods of time.

Labor - Because land under no-till is not cleared before planting and involves less weeding and pest
problems following the establishment of permanent soil cover/crop rotations, farmers in Ghana
reported a 22% savings in labor associated with maize production. Similar reductions in labor
requirements have been reported with no-till rice-wheat systems in South Asia and various CA
technologies in South America. Much of the reduced labor comes from the absence of tillage
operations under CA, which use up valuable labor days during the planting season.

Water - Conservation agriculture requires significantly less water use due to increased infiltration and
enhanced water holding capacity from crop residues left on the soil surface. Mulches also protect the

230
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

soil surface from extreme temperatures and greatly reduce surface evaporation, which is particularly
important in tropical and sub-tropical climates. In Sub-Saharan Africa, as with other dryland regions,
the benefits of conservation agriculture are most salient during drought years, when the risk of total
crop failure is significantly reduced due to enhanced water use efficiency.

Nutrients - Soil nutrient supplies and cycling are enhanced by the biochemical decomposition of
organic crop residues at the soil surface that are also vital for feeding the soil microbes. While much
of the nitrogen needs of primary food crops can be achieved by planting nitrogen-fixing legume
species, other plant essential nutrients often must be supplemented by additional chemical and/or
organic fertilizer inputs. In general, soil fertility is built up over time under conservation agriculture,
and fewer fertilizer amendments are required to achieve optimal yields over time.

Soil biota - Insect pests and other disease causing organisms are held in check by an abundant and
diverse community of beneficial soil organisms, including predatory wasps, spiders, nematodes,
springtails, mites and beneficial bacteria and fungi, among other species. Furthermore, the burrowing
activity of earthworms and other fauna create tiny channels or pores in the soil that facilitate the
exchange of water and gases and loosen the soil for enhanced root penetration.

Economic benefits - Farmers using CA technologies typically report higher yields (up to 45-48%
higher) with fewer water, fertilizer and labor inputs, thereby resulting in higher overall farm profits. In
Paraguay, net farm income of no-till (NT) farming on large-scale commercial farms increased from
$2,3467 to $32,608 more than farms using conventional tillage over a 10 year period. The economic
benefits of NT and other conservation agriculture technologies, more than any other factor, has lead to
widespread adoption among both large- and small-scale farmers throughout the world.

Environmental benefits - Conservation agriculture represents an environmentally-friendly set of

technologies. Because it uses resources more efficiently than conventional agriculture, these resources
become available for other uses, including conserving them for future generations. The significant
reduction in fossil fuel use under no-till agriculture results in fewer greenhouse gases being emitted
into the atmosphere and cleaner air in general. Reduced applications of agrochemicals under CA also
significantly lessens pollution levels in air, soil and water.

Equity considerations -Conservation agriculture also has the benefit of being accessible to many
small-scale farmers who need to obtain the highest possible yields with limited land area and inputs.
Perhaps the biggest obstacle thus far for the technology spreading to more small-scale farmers
worldwide has been limited access in certain areas to certain specialized equipment and machinery,
such as no-till planters.

Active role for farmers -As with any new agricultural technology, CA methods are most effective
when used with skillful management and careful consideration of the many agroecolgical factors
affecting production on any given farm or field. Rather than being a fixed technology to be adopted in
blueprint-like fashion, CA should be seen as a set of sound agricultural principles and practices that
can be applied either individually or together, based on resource availability and other factors. For this
reason, farmers are encouraged to experiment with the methods and to evaluate the results for
themselves- not just to "adopt" CA technologies. Selecting among different cover crop species, for
example, needs to be determined in relation to particular agroecological conditions of the farm,
231
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

including soil type, climate, topography as well as seed availability and what the primary function of
the GMCC will be. Similarly, planting distances, irrigation requirements and the use of agrochemicals
to control weeds and pests among other considerations, must be decided based on what the farmer
needs as well as the availability of these and other resources.

PRINCIPLES OF CONSERVATION FARMING

The FAO has determined that CA has three key principles that producers (farmers) can proceed
through in the process of CA. These three principles outline what conservationists and producers
believe can be done to conserve what we use for a longer period of time.

The first key principle in CA (Conservation Agriculture) is practicing minimum soil disturbance
which is essential to maintaining minerals within the soil, stopping erosion, and preventing water loss
from occurring within the soil. In the past agriculture has looked at soil tillage as a main process in the
introduction of new crops to an area. It was believed that tilling the soil would increase fertility within
the soil through mineralization that takes place in the soil. Also tilling of soil can cause severe erosion
and crusting which leads to a decrease in soil fertility. Today tillage is seen as destroying organic
matter that can be found within the soil cover. No-till farming has caught on as a process that can save
soil organic levels for a longer period and still allow the soil to be productive for longer periods (FAO
2007). Additionally, the process of tilling can increase time and labor for producing that crop.
Minimum soil disturbance also reduce destruction of soil micro and macro-organism habitats that is
common in conventional ploughing practices.

When no-till practices are followed, the producer sees a reduction in production cost for a certain crop.
Tillage of the ground requires more money in order to fuel tractors or to provide feed for the animals
pulling the plough. The producer sees a reduction in labor because he or she does not have to be in the
fields as long as a conventional farmer.

The second key principle in CA is permanent soil cover with live or dead plant material (e.g crop
residues). The principle of managing the top soil to create a permanent organic soil cover can allow
for growth of organisms within the soil structure. This growth will break down the mulch that is left
on the soil surface. The breaking down of this mulch will produce a high organic matter level which
will act as a fertilizer for the soil surface. If CA practices were used done for many years and enough
organic matter was being built up at the surface, then a layer of mulch would start to form. This layer
helps prevent soil erosion from taking place and ruining the soil's profile or layout. The presence of
mulching also reduce the velocity of runoff and the impact of rain drops thus reducing soil erosion and
runoff.

According to the article “The role of conservation agriculture and sustainable agriculture”, the layer of
mulch that is built up over time will become like a buffer zone between soil and mulch and this will
help reduce wind and water erosion. With this comes the protection of the soil's surface when rain falls
on the ground. Land that is not protected by a layer of mulch is left open to the elements (Hobbs et al.
2007). This type of ground cover also helps keep the temperature and moisture levels of the soil at a
higher level rather than if it was tilled every year (FAO 2007).

232
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

The third principle is crop diversification, (e.g crop rotations, cover crops or intercrops with legumes)
According to an article published in the Physiological Transactions of the Royal Society called “The
role of conservation agriculture and sustainable agriculture,” crop rotation can be used best as a
disease control against other preferred crops (Hobbs et al. 2007). This process will not allow pests
such as insects and weeds to be set into a rotation with specific crops. Rotational crops will act as a
natural insecticide and herbicide against specific crops. Not allowing insects or weeds to establish a
pattern will help to eliminate problems with yield reduction and infestations within fields (FAO 2007).
Crop rotation can also help build up soil infrastructure. Establishing crops in a rotation allows for an
extensive buildup of rooting zones which will allow for better water infiltration (Hobbs et al. 2007).

Organic molecules in the soil break down into phosphates, nitrates and other beneficial elements
which are thus better absorbed by plants. Plowing increases the amount of oxygen in the soil and
increases the aerobic processes, hastening the breakdown of organic material. Thus more nutrients are
available for the next crop but, at the same time, the soil is depleted more quickly of its nutrient
reserves.

CONSERVATION FARMING PRACTICES

The process of no-till is one that follows the first principle of CA, causing minimal mechanical soil
disturbance. No-till also brings other benefits to the producer . According to the FAO, tillage is one of
the most “energy consuming” processes that can be used: It requires a lot of labor, time, and fuel to
till. Producers can save 30% to 40% of time and labor by practicing the no-till process. (FAO 3020)

Besides conserving the soil, there are other examples of how CA is used. According to an article
in Science called “Farming and the Fate of Wild Nature” there are two more kinds of CA . The
practice of wildlife-friendly farming and land sparing are ideas for producers who are looking to
practice better conservation towards biodiversity (Green, et al. 2005).

Wildlife-friendly farming

Wildlife-friendly farming is a practice of setting aside land that will not be developed by the producer
(farmer). This land will be set aside so that biodiversity has a chance to establish itself in areas with
agricultural fields. At the same time, the producer is attempting to lower the amount of fertilizer and
pesticides used on the fields so that organisms and microbial activity have a chance to establish
themselves in the soil and habitat (Green, et al. 2005). But as in all systems, not all can be perfect. To
create a habitat suitable for biodiversity something has to be reduced, and as in this case for
agriculture farmers, yields can be reduced. This is where the second idea of land sparing can be looked
on as an alternative manner

Land sparing

Land sparing is another way that producer and conservationist can be on the same page. Land sparing
advocates for the land that is being used for agricultural purposes to continue to produce crops at
increased yield. With an increase in yield on all land that is in use, other land can be set aside for
conservation and production for biodiversity. Agricultural land stays in production but would have to
increase its yield potential to keep up with demand. Land that is not being put into agriculture would
be used for conserving biodiversity (Green, et al. 2005). In fact, data from the Food and Agriculture
233
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

Organization shows that between 1961 and 2012, the amount of arable land needed to produce the
same amount of food declined by 68 percent worldwide.

CHAPTER 32

CROP MANAGEMENT

 Describe harvesting indices and methods of a named cereal and legume crops
 describe post harvest technology of a named legume and cereal crops
 discuss amrketing a named legume and cereal crops
 discuss signifiance of record-keeping in the production of a named cereal and legume crop.
 Keeping records of a named cereal and legume crop

Maturity Indices and Harvest Indices

Maturity can be described as the attainment of the particular size or stage after which ripening takes
place.

On the other hand, ripening means the qualitative changes in fruits after maturity of which it become
edible.

MATURITY

It is the stage of fully development of tissue of fruit and vegetables only after which it will ripen
normally. During the

Physiological maturity

It refers to the stage in the development of the harvestable part when maximum growth and maturation
has occurred. It is usually associated with full ripening in the fruits. The Physiological mature stage is
followed by senescence.

Commercial maturity

It is the state of plant organ required by a market. It commonly bears little relation to Physiological
maturity and may occur at any stage during development stage.

Harvest Maturity

It may be defined in terms of Physiological maturity and horticultural maturity, it is a stage, which
will allow fruits / vegetables at its peak condition when it reaches to the consumers and develop
acceptable flavour or appearance and having adequate shelf life.

Various Maturity Indices are:

234
Prepared by Shamuyarira, Mvurwi High School.2020
 Comprehensive Crop Science( 6049) Hand out

i) Number of days from fruit set,

ii) Visual indicators,
iii) Size of fruits,
iv) Shape of fruit,
v) Colour of fruit,
vi) Appearance (External)
vii) Texture
viii) Lenticel number
ix) Specific gravity
x) Starch Content
xi) Soluble solids
xii) Sugar acid ratio
xiii) Oil content

These indices are used to determination of fruits, Bhatnagar and Suramanyam ( 1971) reported that in
Mango varieties – Alphanso and Pairi it took 110 to 125 days after fruit set for surface colour to
change from dark green to olive green and flesh colour from white to pale yellow. Haden mangos are
ready for picking at 1.02 specific gravity. The general practice mango observed in this process is
known as TAPKA. The stage of maturity of Banana is determined by experience and judged largely
by the visual appearances of the hanging branch and particularly by angularity of the individual
banana fingers.

Harvest Indices:

The quality of fruits and vegetable cannot be improved but it can be presented when harvesting is
done at proper stage of maturity. Immature fruits when harvested will give poor quality and erratic
ripening.

In some cases, if the produce is to shipped to distant markets, or stored, to wait for a better price, it
should be picked in the mature but unripe stage. Here lies the difficulty, because unlike the ripening
stage, the boundary between pre maturation and maturation stage is hard to detect. No prominent
changes in firmness or colour are evident often harvest indices becomes allitrary and subjective.

235
Prepared by Shamuyarira, Mvurwi High School.2020