Photosynthesis Efficiency and Radiation Balance, Water Use Efficiency, Canopy Transmissivity,

Canopy Management, Plant Geometry and Crop Yield in Agroforestry

• Species Interactions
• Competition
– Plants utilize the same resource, the growth of one or both is reduced
– Monocultures – intense competition (genetically similar – same resource
requirements)
– Threshold – increasing density, production constant
– Exceed threshold – add different species
• Complementarity
– Niche differentiation – plants utilize different growing space or have different
resource requirements
– Species mixtures = more complete/efficient resource use = increased productivity
– Theory:  Species diversity =  productivity
* Empirical evidence lacking (contradictory)
– Key: a few species from different life forms
• Facilitation
• One species directly benefits another growing in a mixture
• Nutrient cycling efficiency
• Soil structure – water & nutrient retention
• Soil moisture status – hydraulic lift
• Reduction of loss to pests, pathogens, weeds
• Balance: maintain facilitative functions and reduce competition
• Water use efficiency
• The water utilized by crop is evaluated in terms of Water Use Efficiency
• This water use efficiency can be classified into:
• i. Crop water use efficiency (CWUE) - It is the ratio of crop yield (Y) to the amount of water
used by the crop for evapotranspiration (ET)
• CWUE =Y/ET
• Expressed as kg/mm/ha
 ii. Field water use efficiency (FWUE) - It is the ratio of crop yield (Y) to the total amount of
water used in the field (WR)
 FWUE= Y/WR
 Expressed as kg/mm/ha

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  iii. Physiological water use efficiency (PWUE)- The physiological WUE is calculated in
terms of the amount of CO2 fixed per unit of water transpired
 PWUE = Rate of photosynthesis/Rate of transpiration

WUE = NPP/E
Where, NPP = Net primary productivity, E = evapotranspiration
 It will vary in space (edaphoclimatic conditions, planting density) and time (seasonality, age)
 It is a very good indicator of the production of biomass
 Tree increases water productivity in two distinct ways:
1. Increasing the quantity of water used, transpiration (Tw)
2. Increasing biomass per unit of water consumed (Ew)
Biomass = (Ew) x (Tw)
• Management Implication
 Tree water can be optimized:
 To decreased dry season demand
 Minimize below ground competition with crop and
 The diversion of precious water from other uses
 Preferentially planting deciduous tree species
 Avoiding fast growing, evergreen species viz. Pines and Eucalyptus, particularly in water
scarce areas
 Practicing root & shoot pruning to decrease below ground competition
 Avoiding rotational woodlots with frequent harvesting

Photosynthetic efficiency
 Photosynthesis consists essentially of carbon "fixation" in the green tissues of plants, in the
presence of sunlight
CO2 + 2H2O -------------> (CH2O) + H2O + O2
 Plant productivity, i.e., the amount of growth that can be attained by a plant within a given
period of time, is a function of the net rate of photosynthesis (PN), which is the difference
between gross photosynthesis (PG) and respiration (R)
PN = PG – R
 Respiration involves the oxidation (or breakdown) of complex substances such as sugars and
fats.
Factors affecting rate of photosynthesis

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Environmental factors
 Light
 Availability of CO2
 Temperature
 Soil-water
 Nutrient supply
Genetic factors
 Leaf age
 leaf angle
 leaf orientation
 chlorophyll content of leaf
• Solar Radiation
 Chlorophyll Synthesis
 Stomata regulation- open in light and closed in darkness except CAM
 Main constituent of photosynthesis
 Affect the growth and development- quality, intensity (quantity) and duration of light
influences photosynthesis
 Form and quality of tree- diameter growth observed in day time and height growth in night
time due to elongation of cell
 Species stratification- top canopy occupy by LD
 Size and structure of leaves
• Intensity of light
 Light between the wavelength of 400-700nm is PAR
 At low to moderate level of light intensity, the rate of photosynthesis increases
proportionally with increasing light intensity
 When light intensity is above a certain level, the rate of photosynthesis does not increase
with further increase in light intensity because of:
 Carbon dioxide concentration becomes limiting and
 Destruction of chlorophyll occurs
 Extremely high intensity of light favors more root growth than shoot growth
 Generally, Light passing through the tree canopy experiences change in intensity because of
reflectance, absorbance, and transmittance
•
• The Effect of Light Intensity on Photosynthetic Rate

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•

• Quality of Light
 It refers to wavelength of light spectrum i.e. colour of light
 Maximum photosynthesis-in red and blue light and minimum-in green light
 Plant grown in blue light will be short in size but show normal growth
 Infra red- elongation of cell occurs
 Violet and UV rays- plant became dwarf
• Duration of Light

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 Photoperiod is the relative day and night period to which plant is exposed
 Longer the photoperiod, more photosynthesis as a result more growth
 Affects the germination, leaf fall and flowering (Phenology) and breaking the dormancy
 Photoperiod varies with species. e.g. short days may cause dormant stage in poplar but not
in juniperus
• The effect of CO2 concentration on the rate of photosynthesis
 In C3 plants, rate of photosynthesis increases with increasing CO2 concentration (upto
2000μΙ/I) when other factors are not limiting
 Reason
 High availability of substrate for the carboxylation reaction
 Reduced photorespiration
 In C4 plant, rate of photosynthesis increases with increasing CO2 but attain
saturation at much lower conc. (360 μΙ/I) than the C3 plants

• Temperature
 The rate drops if the temperature is too high because in photosynthesis the light reactions
use ATP synthase, while Rubisco is found in the "dark reactions“
 A high temperature will cause them to denature, which brings down the rate (affinity of
enzyme rubisco for CO2 is reduced)
 The optimal temperature that gives the highest rate of photosynthesis is 25 °C in C3 but in
35 °C C4 plant

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  Low temperature also inactivates enzymes
• Soil water
 Rate of photosynthesis decline under condition of water stress because:
 Stomatal closure and the resultant decrease in CO2 supply
 Reduced leaf water potential
 Low leaf water potential reduces leaf expansion which causes significant reduction
in photosynthetic surface area
• Nutrient supply
 Among various nutrients, nitrogen has a direct relationship with Photosynthesis
 Nitrogen is a basic constituent of chlorophyll and all enzyme involved in carbon reaction
 Rubisco alone accounts for more than half of the total leaf nitrogen
 N and P are probably most often limiting protein building element for growth process
• Nutrients
Mitscherlich’s law
 Assimilation activities of given leaf areas are DECISIVELY influenced by the supply of
nutrients which depends on soil quality
Hartig
 Soil with plenty supply of nutrients can produce TWICE the amount of timber than poor soils
for the same amount of foliage

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Significance of Radiation
 It provides the necessary energy for all the phenomena
concerning biomass production
 Photo synthetically Active Radiations (PAR) are the real source
of energy for photosynthesis process
 Plants are the efficient biological converters of solar energy into biomass
 Radiation defines the yield of crop in particular region
 It also provides the energy for the physical processes taking
place in plants, soil and atmosphere

Canopy Transmissivity
 It is defined as the ratio of the transmitted radiation to the total
radiation incident upon the medium
 It enable the prediction of energy transmission through the canopy to lower layers of
biomass or to ground level
 The fraction of incoming PAR absorbed by the canopy is the opposite of canopy
transmittance
 It can be calculated from the ratio of down welling PAR below the canopy to down welling
PAR above the canopy
 Down welling PAR above the canopy does not tend to vary spatially during clear conditions

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  However, down welling PAR below the canopy varies significantly both in space and time
due to canopy structure and openness
 The fraction of incoming PAR is closely related to the canopy gap fraction

Purpose of Canopy Management

 To optimize the plant model in such a way that is suitable and compatible with the
associated arable crops in any agroforestry system
Prerequisite of Canopy Management
1. Full understanding of the crown architecture and growth
2. Nature of trees
3. Nature of the companion crops
4. Objective of agroforestry system
Basic Principles of Canopy Management
 To make best use of the land
 To make maximum use of available solar radiations for increased productivity
 Convenience in carrying out the cultural practices
Options for Canopy Management
 The option for managing tree canopy are:
 Pruning
 Coppicing
 Pollarding

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  Lopping
 Partial crown removal
 Crown lifting
Pruning
 Selective removal of parts of a plant such as branches, buds, or roots
 Powerful approach to regulate above ground competition in agroforestry system
 Can increase the light transmitting in agroforestry system and increase the crop production
Objectives of Pruning in Agroforestry
 Increase crop production
 To improve the wood quality (no nodular wood)
 Fodder and mulch production
i. Fixed lift pruning
ii. Variable-lift pruning
iii. Selective pruning
iv. Tip pruning
v. Basal branch pruning/ side pruning
When to Prune?
Basic Considerations for time of Pruning:
1. Dead, damaged, diseased, and dying wood: Anytime of year, and should be removed
2. Spring-flowering trees: Should be pruned after they bloom for the season since the flower is
growing on wood that was produced during the previous years growing season
3. Summer-flowering trees: Should be pruned in late winter (after threat of frost) and/or
before new spring growth occurs, their flowers are borne of wood produced that same year
4. Trees without flowers: Should be pruned when they are dormant, typically in mid-late
winter or early spring, before new growth had developed, this group includes most
deciduous trees, needled evergreens, and many of the broadleaf evergreens
Height of pruning
 Nature of tree i.e photosynthetic efficiency of the tree
 Nature of the companion crop
 How long the acceptable log length is in the area
 Availability of the specialized equipment which allow cheap and safe high pruning
How Much Should Be Pruned?
 The amount of live tissue that should be removed depends on the tree size, species, and
age, as well as the pruning objectives

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  The optimal pruning intensity should involve a trade-off between maximising gains both in
crop production and wood quality , and minimising impacts on tree growth.

How Coppicing?
 The cut should be at angle
 The bark should not be damaged
 Young coppice should be kept weed free
 Coppicing should be done in the dormant season
 When it is done to produce mulch, fodder, or small wood, the sprouts can be left to grow
and be pruned when needed
 When the intention is to produce a new tree, all sprouts are removed except one that is
going straight up, this will form new tree

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Coppicing Frequency
 Depend upon:
 species
 environment and
 amount of foliage left
 Depending upon the tree species used and the crop with which it is grown:
 coppicing can be done 4 to 5 times in a year ( sometimes even more)
Pollarding
 Removal of all the branches and top of the tree at a height 1m or more above the ground
level so that only the trunk is left
 The tree is then allowed to regrow after the initial
cutting, but once begun, pollarding requires regular
maintenance by pruning
 Pollarding of older trees may result death of the tree,
especially
 If there are no branches below the cut, or
 The tree is of an inappropriate species

Purposes
• Applied in the alley and other agroforestry system to harvest wood, fodder, mulch and other
biomass and to reduce shade for the adjacent crops

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 • The main reason for this type of practice, rather than coppicing, is in wood pastures and
grazing areas where growth from the ground upwards was less practicable, due to the
required area for grazing which would have been reduced by thickets of low tree growth
• Pollarding above head height protects valuable timber or poles from being damaged by
browsing animals
When it is done
 For timber or pole, cut the tree at higher point, leaving a long stem to grow
 To produce smaller wood, fodder, mulch and so on , cut the tree lower making it easy to
reach
How to pollard?
 On a shrub, this might be a one stem cut to a meter high, a mass of stem will grow from the
top
 With a tree, it is more typical to leave a trunk supporting three to five branches and this
branches are cut to a desired length i.e. 1 to 2 inches from the main stem
• Conti…
How to Maintain a pollard?
 Once a tree and shrub is pollarded, continue the annual cycle of cutting
 Branch should be pruned just above the previous pollard cuts
When to pollard ?
 The best time for pollarding many trees and shrubs is in late winter or early spring
 The least favourable time for pollarding is the autumn, as decay fungi may enter the pruning
cuts
Age of pollarding
 Start pollarding while the tree is young i.e. 2 to 3 years.
Lopping
• Lopping is the harvesting of the branch in a more haphazard way
• The main reason for lopping is to collect the fodder
• Branches can be cut from any part of the tree but not too close from the trunk, however, do
not take too many branches at a time
• The lopping can be done at any time when fodder are required

 PAR are the real source of energy for photosynthesis process, plants are the efficient
biological converters of solar energy into biomass and it defines the yield of crop in
particular region

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  Basic principles of canopy management is to make best use of the land, to make maximum
use of available solar radiations for increased productivity and convenience in carrying out
the cultural practices
 Plant geometry refers to the shape of the space available for individual plants and it
influences crop yield through its influence on light interception, rooting pattern and
moisture extraction pattern

Plant Geometry and Crop Yield

• It is the pattern of distribution of plant over the ground or the shape of the area available to
the individual plant, in a crop field
• Plant geometry refers to the shape of the space available for individual plants
• It influences crop yield through its influence on light interception, rooting pattern and
moisture extraction pattern
• Plant geometry is altered by changing inter and intrarow spacing
• In agro-forestry experiments row-to-row distance should be in east-west direction and tree-
to-tree distance in north-south direction.
• This will give tree rectangularity of 1 or more than 1, which is essential for proper utilization
of light in agro-forestry
Planting patterns of perennial vegetations like trees/shrubs

i. Square planting
ii. Rectangular planting
iii. Triangular planting
iv. Miscellaneous planting

 Skip row planting

 Paired row planting

Management options in competitive tree-crop interface

 The magnitude of interactive effects between trees and other components of agroforestry systems
depends on the characteristics of the species, their planting density, and spatial arrangement and
management of the trees. Manipulating densities and arrangements is probably the most powerful
method for capitalizing on beneficial effects of trees while reducing negative ones. Whenever a tree
species with all the desired characteristics is not available (which is most likely to be the case), tree
crowns and roots can be manipulated through management operations, mainly by pruning and
thinning. Other common management operations such as fertilization, application of mulch and
manure, cut-and-carry fodder systems, and confinement or rotation of the animals can also be

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 employed. The different manipulations can be grouped as growth-enhancing or growth-reducing
according to their effect on the targeted component (Table below).
Management options to achieve

[Link] growth [Link] growth

 Microclimate amelioration  Prunning
 Fertilization  Pollarding
 Application of mulch/manure  Trenching
 Irrigation  Excessive shading
 Soil tillage  Herbicides
 Adapted species  Grazing/browsing
 Supplemental feeding

 The goals of management practices should be to increase the production of the desired
products and to decrease growth and, hence, competition of undesired components. In many cases,
one cultural treatment will accomplish both goals simultaneously, e.g., in the case of pruning trees in
alley cropping and applying the biomass to the soil. While the removal of parts or the entire crown
will obviously reduce the tree's competitive ability, it will automatically increase the growth of the
associated intercrop by providing green manure and by allowing more light to penetrate to the
crop.' Below-ground competition may also be reduced as a result of pruning-induced root die-back
(Cannell, 1983).- These observations also apply to pruning or pollarding operations on trees grown
for shade or as live supports, such as legumes of the genera Erythrina, Inga or Gliricidia. Species such
as Erythrina berteroana, which have large thick leaves and high rates of biomass production when
grown as a shade tree, will require more intensive pruning than trees with a less dense canopy such
as Gliricidia sepium

• The term ”architecture” in reference to biological objects usually denotes the spatial
configuration of some complex assemblage of subunits, with the implication that the overall
configuration has some functional significance
• In its simplest form, the term root architecture refers to the spatial configuration of a
plant’s root system
• This system can be extremely complex and is dependent upon multiple factors such as the
species of the plant itself, the composition of the soil and the availability of nutrients
• Root architecture plays the important role of providing a secure supply of nutrients and
water as well as anchorage and support
The main terms used to classify the architecture of a root system are:
• Branch magnitude: the number of links (exterior or interior)
• Topology: the pattern of branching, including:
1. Herringbone: alternate lateral branching off a parent root
2. Dichotomous: opposite, forked branches
3. Radial: whorl(s) of branches around a root

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 4. Link length: the distance between branches
• Root angle: the radial angle of a lateral root’s base around the parent root’s circumference,
the angle of a lateral root from its parent root, and the angle an entire system spreads
• Link radius: the diameter of a root.
• The configuration of root systems is important to support the plant, compete with other
plants and for uptake of nutrients from the soil.
• Roots grow to specific conditions, which, if changed, can impede a plants growth. For
example, a root system that has developed in dry soil may not be as efficient in flooded soil,
however, plants are still able to adapt to changes in the environment, such as seasonal
changes.
Mechanisms of root interactions and perspectives of their management
Root distribution and root interactions at the root systems level
• A fundamental hypothesis of agroforestry is that different plant life forms such as trees and
herbaceous crops or pastures occupy to some extent different soil strata with their root
systems when grown in association, leading to a degree of complementarity in their use of
soil resources
• For this reason, rooting depth and the vertical distribution of root systems are of particular
interest for agroforestry
• Regarding rooting depth, it is important to distinguish between the total depth explored by a
root system and the distribution of the roots in the soil profile
• In certain cases, associated plant species form vertically stratified root systems, i.e. their
root systems occupy more or less separate soil layers.
• This situation is desirable in many agroforestry situations because it is likely to reduce
overall root competition and to lead to a more complete utilization of soil resources.
• The ‘safety net‘ of tree roots below the root systems of associated crops refers to a situation
of vertically stratified root systems, in which the tree roots absorb nutrients which have not
been taken up by the shallower-rooted crops and have therefore been leached out of the
topsoil.
a. Coarse roots
Function: tree support, storage of carbohydrates and nutrients
b. Fine roots (< 2-5 mm diameter)
Function: water and nutrient absorption, synthesis of growth regulation compounds. Most
fine roots occur in the upper 30 cm (12 inches) of soil. Fine root growth can occur any time of year as

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long as the soil is not frozen. Typically peaks in fine root growth correspond with water and nutrient
availability.
• -quantity variable
• -regional variations
c. Lateral Roots-extend laterally from the tree
d. Specialized root- pneumatophores or knee, buttress roots
• Plants accumulate and redistribute macro and micronutrients throughout their life cycle.
• Ultimately the plant is seeking to optimize the way it responds to environmental cues and to
integrate these with the genetically determined elements of development in order to
maximize its reproductive potential and ensure the production and survival of the next
generation.
• For annual plants under optimal growth conditions, the process of resource allocation
ensures that ultimately nearly all the nitrogen resources produced during the photosynthetic
period will be remobilized from the vegetative tissues into the developing seeds.
• In contrast, perennial plants commit a more limited proportion of resources to the
reproductive phase of their life cycle as it is advantageous to retain some photosynthetic
capacity for as long as the environmental conditions allow.
• Most species of higher plants have qualitatively similar resource requirements for growth
and reproduction. They differ, however, in the way they use resources to carry out three
essential functions reproduction, defence against herbivores, and growth.
• Variation in resource allocation occurs through differences in the chemical composition of
structures, the relative mass of different structures or organs, and the relative numbers of
different structures a plant produces.

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