Botany

Botany, also called plant science or phytology, is the

branch of natural science and biology studying plants,
especially their anatomy, taxonomy, and ecology.[1] A
botanist, plant scientist or phytologist is a scientist who
specialises in this field. Nowadays, botanists (in the strict
sense) study approximately 410,000 species of land
plants, including some 391,000 species of vascular plants
(of which approximately 369,000 are flowering plants)[2]
and approximately 20,000 bryophytes.[3]

Botany originated in prehistory as herbalism with the

efforts of early humans to identify – and later cultivate –
plants that were edible, poisonous, and possibly The fruit of Myristica fragrans, a species native
medicinal, making it one of the first endeavours of to Indonesia, is the source of two valuable
human investigation. Medieval physic gardens, often spices, the red aril (mace) enclosing the dark
attached to monasteries, contained plants possibly having brown nutmeg.
medicinal benefit. They were forerunners of the first
botanical gardens attached to universities, founded from
the 1540s onwards. One of the earliest was the Padua botanical garden. These gardens facilitated the
academic study of plants. Efforts to catalogue and describe their collections were the beginnings of plant
taxonomy and led in 1753 to the binomial system of nomenclature of Carl Linnaeus that remains in use to
this day for the naming of all biological species.

In the 19th and 20th centuries, new techniques were developed for the study of plants, including methods
of optical microscopy and live cell imaging, electron microscopy, analysis of chromosome number, plant
chemistry and the structure and function of enzymes and other proteins. In the last two decades of the
20th century, botanists exploited the techniques of molecular genetic analysis, including genomics and
proteomics and DNA sequences to classify plants more accurately.

Modern botany is a broad subject with contributions and insights from most other areas of science and
technology. Research topics include the study of plant structure, growth and differentiation, reproduction,
biochemistry and primary metabolism, chemical products, development, diseases, evolutionary
relationships, systematics, and plant taxonomy. Dominant themes in 21st-century plant science are
molecular genetics and epigenetics, which study the mechanisms and control of gene expression during
differentiation of plant cells and tissues. Botanical research has diverse applications in providing staple
foods, materials such as timber, oil, rubber, fibre and drugs, in modern horticulture, agriculture and
forestry, plant propagation, breeding and genetic modification, in the synthesis of chemicals and raw
materials for construction and energy production, in environmental management, and the maintenance of
biodiversity.
Etymology
The term "botany" comes from the Ancient Greek word botanē (βοτάνη) meaning "pasture", "herbs"
"grass", or "fodder";[4] Botanē is in turn derived from boskein (Greek: βόσκειν), "to feed" or "to
graze".[5][6][7] Traditionally, botany has also included the study of fungi and algae by mycologists and
phycologists respectively, with the study of these three groups of organisms remaining within the sphere
of interest of the International Botanical Congress.

History

Early botany
Botany originated as herbalism, the study and use of plants for
their possible medicinal properties.[8] The early recorded history
of botany includes many ancient writings and plant classifications.
Examples of early botanical works have been found in ancient
texts from India dating back to before 1100 BCE,[9][10] Ancient
Egypt,[11] in archaic Avestan writings, and in works from China
purportedly from before 221 BCE.[9][12]

Modern botany traces its roots back to Ancient Greece specifically

to Theophrastus (c. 371–287 BCE), 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".[13]
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.[13][14]
An engraving of the cells of cork,
from Robert Hooke's Micrographia,
Another work from Ancient Greece that made an early impact on 1665
botany is De materia medica, a five-volume encyclopedia about
preliminary herbal medicine written in the middle of the
first century by Greek physician and pharmacologist
Pedanius Dioscorides. De materia medica was widely read
for more than 1,500 years.[15] Important contributions from
the medieval Muslim world include Ibn Wahshiyya's
Nabatean Agriculture, Abū Ḥanīfa Dīnawarī's (828–896)
the Book of Plants, and Ibn Bassal's The Classification of Leonhart Otto Brunfels Hieronymus
Soils. In the early 13th century, Abu al-Abbas al-Nabati, Fuchs Bock
and Ibn al-Baitar (d. 1248) wrote on botany in a systematic
and scientific manner.[16][17][18]
In the mid-16th century, botanical gardens were founded in a number of Italian universities. The Padua
botanical garden in 1545 is usually considered to be the first which is still in its original location. These
gardens continued the practical value of earlier "physic gardens", often associated with monasteries, in
which plants were cultivated for suspected medicinal uses. They supported the growth of botany as an
academic subject. Lectures were given about the plants grown in the gardens. Botanical gardens came
much later to northern Europe; the first in England was the University of Oxford Botanic Garden in
1621.[19]

German physician Leonhart Fuchs (1501–1566) was one of "the three German fathers of botany", along
with theologian Otto Brunfels (1489–1534) and physician Hieronymus Bock (1498–1554) (also called
Hieronymus Tragus).[20][21] Fuchs and Brunfels broke away from the tradition of copying earlier works
to make original observations of their own. Bock created his own system of plant classification.

Physician Valerius Cordus (1515–1544) authored a botanically and pharmacologically important herbal
Historia Plantarum in 1544 and a pharmacopoeia of lasting importance, the Dispensatorium in 1546.[22]
Naturalist Conrad von Gesner (1516–1565) and herbalist John Gerard (1545–c. 1611) published herbals
covering the supposed medicinal uses of plants. Naturalist Ulisse Aldrovandi (1522–1605) was
considered the father of natural history, which included the study of plants. In 1665, using an early
microscope, Polymath Robert Hooke discovered cells (a term he coined) in cork, and a short time later in
living plant tissue.[23]

Early modern botany

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.[24] By the 18th century, new plants for study were
The Linnaean Garden of Linnaeus' arriving in Europe in increasing numbers from newly discovered
residence in Uppsala, Sweden, was countries and the European colonies worldwide. In 1753, Carl
planted according to his Systema Linnaeus published his Species Plantarum, a hierarchical
sexuale. classification of plant species that remains the reference point for
modern botanical nomenclature. This established a standardised
binomial or two-part naming scheme where the first name represented the genus and the second identified
the species within the genus.[25] For the purposes of identification, Linnaeus's Systema Sexuale classified
plants into 24 groups according to the number of their male sexual organs. The 24th group, Cryptogamia,
included all plants with concealed reproductive parts, mosses, liverworts, ferns, algae and fungi.[26]

Increasing knowledge of plant anatomy, morphology and life cycles led to the realisation that there were
more natural affinities between plants than the artificial sexual system of Linnaeus. Adanson (1763), de
Jussieu (1789), and Candolle (1819) all proposed various alternative natural systems of classification that
grouped plants using a wider range of shared characters and were widely followed. The Candollean
system reflected his ideas of the progression of morphological complexity and the later Bentham &
Hooker system, which was influential until the mid-19th century, was influenced by Candolle's approach.
Darwin's publication of the Origin of Species in 1859 and his concept of common descent required
modifications to the Candollean system to reflect evolutionary relationships as distinct from mere
morphological similarity.[27]

In the 19th century botany was a socially acceptable hobby for upper-class women. These women would
collect and paint flowers and plants from around the world with scientific accuracy. The paintings were
used to record many species that could not be transported or maintained in other environments. Marianne
North illustrated over 900 species in extreme detail with watercolor and oil paintings.[28] Her work and
many other women's botany work was the beginning of popularizing botany to a wider audience.

Botany was greatly stimulated by the appearance of the first "modern" textbook, Matthias Schleiden's
Grundzüge der Wissenschaftlichen Botanik, published in English in 1849 as Principles of Scientific
Botany.[29] Schleiden was a microscopist and an early plant anatomist who co-founded the cell theory
with Theodor Schwann and Rudolf Virchow and was among the first to grasp the significance of the cell
nucleus that had been described by Robert Brown in 1831.[30] In 1855, Adolf Fick formulated Fick's laws
that enabled the calculation of the rates of molecular diffusion in biological systems.[31]

Late modern botany

Building upon the gene-chromosome theory of heredity that
originated with Gregor Mendel (1822–1884), August Weismann
(1834–1914) proved that inheritance only takes place through
gametes. No other cells can pass on inherited characters.[32] The
work of Katherine Esau (1898–1997) on plant anatomy is still a
major foundation of modern botany. Her books Plant Anatomy and Echeveria glauca in a Connecticut
Anatomy of Seed Plants have been key plant structural biology greenhouse. Botany uses Latin
texts for more than half a century.[33][34] names for identification; here, the
specific name glauca means blue.
The discipline of plant ecology was
pioneered in the late 19th century
by botanists such as Eugenius Warming, who produced the hypothesis that
plants form communities, and his mentor and successor Christen C.
Raunkiær whose system for describing plant life forms is still in use today.
The concept that the composition of plant communities such as temperate
broadleaf forest changes by a process of ecological succession was
developed by Henry Chandler Cowles, Arthur Tansley and Frederic
Clements. Clements is credited with the idea of climax vegetation as the
most complex vegetation that an environment can support and Tansley
introduced the concept of ecosystems to biology.[35][36][37] Building on
the extensive earlier work of Alphonse de Candolle, Nikolai Vavilov
Class of alpine botany in
(1887–1943) produced accounts of the biogeography, centres of origin,
Switzerland, 1936
and evolutionary history of economic plants.[38]

Particularly since the mid-1960s there have been advances in

understanding of the physics of plant physiological processes such as transpiration (the transport of water
within plant tissues), the temperature dependence of rates of water evaporation from the leaf surface and
the molecular diffusion of water vapour and carbon dioxide through stomatal apertures. These
developments, coupled with new methods for measuring the size of stomatal apertures, and the rate of
photosynthesis have enabled precise description of the rates of gas exchange between plants and the
atmosphere.[39][40] Innovations in statistical analysis by Ronald Fisher,[41] Frank Yates and others at
Rothamsted Experimental Station facilitated rational experimental design and data analysis in botanical
research.[42] The discovery and identification of the auxin plant hormones by Kenneth V. Thimann in
1948 enabled regulation of plant growth by externally applied chemicals. Frederick Campion Steward
pioneered techniques of micropropagation and plant tissue culture controlled by plant hormones.[43] The
synthetic auxin 2,4-dichlorophenoxyacetic acid or 2,4-D was one of the first commercial synthetic
herbicides.[44]

20th century developments in plant biochemistry have been driven by

modern techniques of organic chemical analysis, such as spectroscopy,
chromatography and electrophoresis. With the rise of the related
molecular-scale biological approaches of molecular biology, genomics,
proteomics and metabolomics, the relationship between the plant genome
and most aspects of the biochemistry, physiology, morphology and
behaviour of plants can be subjected to detailed experimental analysis.[45]
The concept originally stated by Gottlieb Haberlandt in 1902[46] that all
plant cells are totipotent and can be grown in vitro ultimately enabled the
use of genetic engineering experimentally to knock out a gene or genes
responsible for a specific trait, or to add genes such as GFP that report
when a gene of interest is being expressed. These technologies enable the
biotechnological use of whole plants or plant cell cultures grown in
Micropropagation of
bioreactors to synthesise pesticides, antibiotics or other pharmaceuticals,
transgenic plants
as well as the practical application of genetically modified crops designed
for traits such as improved yield.[47]

Modern morphology recognises a continuum between the major morphological categories of root, stem
(caulome), leaf (phyllome) and trichome.[48] Furthermore, it emphasises structural dynamics.[49] Modern
systematics aims to reflect and discover phylogenetic relationships between plants.[50][51][52][53] Modern
Molecular phylogenetics largely ignores morphological characters, relying on DNA sequences as data.
Molecular analysis of DNA sequences from most families of flowering plants enabled the Angiosperm
Phylogeny Group to publish in 1998 a phylogeny of flowering plants, answering many of the questions
about relationships among angiosperm families and species.[54] The theoretical possibility of a practical
method for identification of plant species and commercial varieties by DNA barcoding is the subject of
active current research.[55][56]

Branches of botany
Botany is divided along several axes.

Some subfields of botany relate to particular groups of organisms. Divisions related to the broader
historical sense of botany include bacteriology, mycology (or fungology) and phycology - the study of
bacteria, fungi and algae respectively - with lichenology as a subfield of mycology. The narrower sense of
botany in the sense of the study of embryophytes (land plants) is disambiguated as phytology. Bryology is
the study of mosses (and in the broader sense also liverworts and hornworts). Pteridology (or filicology)
is the study of ferns and allied plants. A number of other taxa of ranks varying from family to subgenus
have terms for their study, including agrostology (or graminology) for the study of grasses,
synantherology for the study of composites, and batology for the study of brambles.

Study can also be divided by guild rather than clade or grade. Dendrology is the study of woody plants.

Many divisions of biology have botanical subfields. These are commonly denoted by prefixing the word
plant (e.g. plant taxonomy, plant ecology, plant anatomy, plant morphology, plant systematics, plant
ecology), or prefixing or substituting the prefix phyto- (e.g. phytochemistry, phytogeography). The study
of fossil plants is palaeobotany. Other fields are denoted by adding or substituting the word botany (e.g.
systematic botany).

Phytosociology is a subfield of plant ecology that classifies and studies communities of plants.

The intersection of fields from the above pair of categories gives rise to fields such as bryogeography (the
study of the distribution of mosses).

Different parts of plants also give rise to their own subfields, including xylology, carpology (or
fructology) and palynology, these being the study of wood, fruit and pollen/spores respectively.

Botany also overlaps on the one hand with agriculture, horticulture and silviculture, and on the other hand
with medicine and pharmacology, giving rise to fields such as agronomy, horticultural botany,
phytopathology and phytopharmacology.

Scope and importance

The study of plants is vital because they underpin almost all animal life on
Earth by generating a large proportion of the oxygen and food that provide
humans and other organisms with aerobic respiration with the chemical
energy they need to exist. Plants, algae and cyanobacteria are the major
groups of organisms that carry out photosynthesis, a process that uses the
energy of sunlight to convert water and carbon dioxide[57] into sugars that
can be used both as a source of chemical energy and of organic molecules
that are used in the structural components of cells.[58] As a by-product of
photosynthesis, plants release oxygen into the atmosphere, a gas that is
required by nearly all living things to carry out cellular respiration. In
addition, they are influential in the global carbon and water cycles and
plant roots bind and stabilise soils, preventing soil erosion.[59] Plants are
crucial to the future of human society as they provide food, oxygen,
Botany involves the
recording and description of
biochemicals, and products for people, as well as creating and preserving
plants, such as this soil.[60]
herbarium specimen of the
lady fern Athyrium filix- Historically, all living things were classified as either animals or plants[61]
femina. and botany covered the study of all organisms not considered animals.[62]
Botanists examine both the internal functions and processes within plant
organelles, cells, tissues, whole plants, plant populations and plant communities. At each of these levels, a
botanist may be concerned with the classification (taxonomy), phylogeny and evolution, structure
(anatomy and morphology), or function (physiology) of plant life.[63]

The strictest definition of "plant" includes only the "land plants" or embryophytes, which include seed
plants (gymnosperms, including the pines, and flowering plants) and the free-sporing cryptogams
including ferns, clubmosses, liverworts, hornworts and mosses. Embryophytes are multicellular
eukaryotes descended from an ancestor that obtained its energy from sunlight by photosynthesis. They
have life cycles with alternating haploid and diploid phases. The sexual haploid phase of embryophytes,
known as the gametophyte, nurtures the developing diploid embryo sporophyte within its tissues for at
least part of its life,[64] even in the seed plants, where the gametophyte itself is nurtured by its parent
sporophyte.[65] Other groups of organisms that were previously studied by botanists include bacteria
(now studied in bacteriology), fungi (mycology) – including lichen-forming fungi (lichenology), non-
chlorophyte algae (phycology), and viruses (virology). However, attention is still given to these groups by
botanists, and fungi (including lichens) and photosynthetic protists are usually covered in introductory
botany courses.[66][67]

Palaeobotanists study ancient plants in the fossil record to provide information about the evolutionary
history of plants. Cyanobacteria, the first oxygen-releasing photosynthetic organisms on Earth, are
thought to have given rise to the ancestor of plants by entering into an endosymbiotic relationship with an
early eukaryote, ultimately becoming the chloroplasts in plant cells. The new photosynthetic plants (along
with their algal relatives) accelerated the rise in atmospheric oxygen started by the cyanobacteria,
changing the ancient oxygen-free, reducing, atmosphere to one in which free oxygen has been abundant
for more than 2 billion years.[68][69]

Among the important botanical questions of the 21st century are the role of plants as primary producers
in the global cycling of life's basic ingredients: energy, carbon, oxygen, nitrogen and water, and ways that
our plant stewardship can help address the global environmental issues of resource management,
conservation, human food security, biologically invasive organisms, carbon sequestration, climate
change, and sustainability.[70]

Human nutrition
Virtually all staple foods come either directly from primary
production by plants, or indirectly from animals that eat them.[71]
Plants and other photosynthetic organisms are at the base of most
food chains because they use the energy from the sun and nutrients
from the soil and atmosphere, converting them into a form that can
be used by animals. This is what ecologists call the first trophic
level.[72] The modern forms of the major staple foods, such as
hemp, teff, maize, rice, wheat and other cereal grasses, pulses, The food we eat comes directly or
bananas and plantains,[73] as well as hemp, flax and cotton grown indirectly from plants such as rice.
for their fibres, are the outcome of prehistoric selection over
thousands of years from among wild ancestral plants with the most
desirable characteristics.[74]
Botanists study how plants produce food and how to increase yields, for example through plant breeding,
making their work important to humanity's ability to feed the world and provide food security for future
generations.[75] Botanists also study weeds, which are a considerable problem in agriculture, and the
biology and control of plant pathogens in agriculture and natural ecosystems.[76] Ethnobotany is the study
of the relationships between plants and people. When applied to the investigation of historical plant–
people relationships ethnobotany may be referred to as archaeobotany or palaeoethnobotany.[77] Some of
the earliest plant-people relationships arose between the indigenous people of Canada in identifying
edible plants from inedible plants. This relationship the indigenous people had with plants was recorded
by ethnobotanists.[78]

Plant biochemistry
Plant biochemistry is the study of the chemical processes used by plants. Some of these processes are
used in their primary metabolism like the photosynthetic Calvin cycle and crassulacean acid
metabolism.[79] Others make specialised materials like the cellulose and lignin used to build their bodies,
and secondary products like resins and aroma compounds.

Plants and various other groups

of photosynthetic eukaryotes
collectively known as "algae"
have unique organelles known
as chloroplasts. Chloroplasts are
thought to be descended from
cyanobacteria that formed
endosymbiotic relationships
with ancient plant and algal
Paper chromatography of some spinach leaf extract shows the various
ancestors. Chloroplasts and pigments present in their chloroplasts: yellowish xanthophylls, greenish
cyanobacteria contain the blue- chlorophylls a and b.
green pigment chlorophyll a.[80]
Chlorophyll a (as well as its
plant and green algal-specific cousin chlorophyll b)[a] absorbs light in the blue-violet and orange/red parts
of the spectrum while reflecting and transmitting the green light that we see as the characteristic colour of
these organisms. The energy in the red and blue light that these pigments absorb is used by chloroplasts to
make energy-rich carbon compounds from carbon dioxide and water by oxygenic photosynthesis, a
process that generates molecular oxygen (O2) as a by-product.

The light energy captured by chlorophyll a is initially in the form of electrons (and later a proton
gradient) that's used to make molecules of ATP and NADPH which temporarily store and transport
energy. Their energy is used in the light-independent reactions of the Calvin cycle by the enzyme rubisco
to produce molecules of the 3-carbon sugar glyceraldehyde 3-phosphate (G3P). Glyceraldehyde 3-
phosphate is the first product of photosynthesis and the raw material from which glucose and almost all
other organic molecules of biological origin are synthesised. Some of the glucose is converted to starch
which is stored in the chloroplast.[84] Starch is the characteristic energy store of most land plants and
algae, while inulin, a polymer of fructose is used for the same purpose in the sunflower family
Asteraceae. Some of the glucose is converted to sucrose (common table sugar) for export to the rest of the
plant.
Unlike in animals (which lack chloroplasts), plants and their eukaryote relatives have delegated many
biochemical roles to their chloroplasts, including synthesising all their fatty acids,[85][86] and most amino
acids.[87] The fatty acids that chloroplasts make are used for many things, such as providing material to
build cell membranes out of and making the polymer cutin which is found in the plant cuticle that
protects land plants from drying out.[88]

Plants synthesise a number of unique polymers like the polysaccharide molecules cellulose, pectin and
xyloglucan[89] from which the land plant cell wall is constructed.[90] Vascular land plants make lignin, a
polymer used to strengthen the secondary cell walls of xylem tracheids and vessels to keep them from
collapsing when a plant sucks water through them under water stress. Lignin is also used in other cell
types like sclerenchyma fibres that provide structural support for a plant and is a major constituent of
wood. Sporopollenin is a chemically resistant polymer found in the outer cell walls of spores and pollen
of land plants responsible for the survival of early land plant spores and the pollen of seed plants in the
fossil record. It is widely regarded as a marker for the start of land plant evolution during the Ordovician
period.[91] The concentration of carbon dioxide in the atmosphere today is much lower than it was when
plants emerged onto land during the Ordovician and Silurian periods. Many monocots like maize and the
pineapple and some dicots like the Asteraceae have since independently evolved[92] pathways like
Crassulacean acid metabolism and the C4 carbon fixation pathway for photosynthesis which avoid the
losses resulting from photorespiration in the more common C3 carbon fixation pathway. These
biochemical strategies are unique to land plants.

Medicine and materials

Phytochemistry is a branch of plant biochemistry primarily concerned with the chemical substances
produced by plants during secondary metabolism.[93] Some of these compounds are toxins such as the
alkaloid coniine from hemlock. Others, such as the essential oils peppermint oil and lemon oil are useful
for their aroma, as flavourings and spices (e.g., capsaicin), and in medicine as pharmaceuticals as in
opium from opium poppies. Many medicinal and recreational drugs, such as tetrahydrocannabinol (active
ingredient in cannabis), caffeine, morphine and nicotine come directly from plants. Others are simple
derivatives of botanical natural products. For example, the pain killer aspirin is the acetyl ester of
salicylic acid, originally isolated from the bark of willow trees,[94] and a wide range of opiate painkillers
like heroin are obtained by chemical modification of morphine obtained from the opium poppy.[95]
Popular stimulants come from plants, such as caffeine from coffee, tea and chocolate, and nicotine from
tobacco. Most alcoholic beverages come from fermentation of carbohydrate-rich plant products such as
barley (beer), rice (sake) and grapes (wine).[96] Native Americans have used various plants as ways of
treating illness or disease for thousands of years.[97] This knowledge Native Americans have on plants
has been recorded by enthnobotanists and then in turn has been used by pharmaceutical companies as a
way of drug discovery.[98]

Plants can synthesise coloured dyes and pigments such as the anthocyanins responsible for the red colour
of red wine, yellow weld and blue woad used together to produce Lincoln green, indoxyl, source of the
blue dye indigo traditionally used to dye denim and the artist's pigments gamboge and rose madder.

Sugar, starch, cotton, linen, hemp, some types of rope, wood and particle boards, papyrus and paper,
vegetable oils, wax, and natural rubber are examples of commercially important materials made from
plant tissues or their secondary products. Charcoal, a pure form of carbon made by pyrolysis of wood, has
a long history as a metal-smelting fuel, as a filter material and adsorbent and as an artist's material and is
one of the three ingredients of gunpowder. Cellulose, the world's most abundant organic polymer,[99] can
be converted into energy, fuels, materials and chemical feedstock. Products made from cellulose include
rayon and cellophane, wallpaper paste, biobutanol and gun cotton. Sugarcane, rapeseed and soy are some
of the plants with a highly fermentable sugar or oil content that are used as sources of biofuels, important
alternatives to fossil fuels, such as biodiesel.[100] Sweetgrass was used by Native Americans to ward off
bugs like mosquitoes.[101] These bug repelling properties of sweetgrass were later found by the American
Chemical Society in the molecules phytol and coumarin.[101]

Plant ecology
Plant ecology is the science of the functional relationships
between plants and their habitats – the environments where they
complete their life cycles. Plant ecologists study the composition
of local and regional floras, their biodiversity, genetic diversity
and fitness, the adaptation of plants to their environment, and their
competitive or mutualistic interactions with other species.[103]
Some ecologists even rely on empirical data from indigenous
people that is gathered by ethnobotanists.[104] This information
The nodules of Medicago italica
contain the nitrogen fixing bacterium can relay a great deal of information on how the land once was
Sinorhizobium meliloti. The plant thousands of years ago and how it has changed over that time.[104]
provides the bacteria with nutrients The goals of plant ecology are to understand the causes of their
and an anaerobic environment, and distribution patterns, productivity, environmental impact,
the bacteria fix nitrogen for the evolution, and responses to environmental change.[105]
plant.[102]
Plants depend on certain edaphic (soil) and climatic factors in their
environment but can modify these factors too. For example, they
can change their environment's albedo, increase runoff interception, stabilise mineral soils and develop
their organic content, and affect local temperature. Plants compete with other organisms in their
ecosystem for resources.[106][107] They interact with their neighbours at a variety of spatial scales in
groups, populations and communities that collectively constitute vegetation. Regions with characteristic
vegetation types and dominant plants as well as similar abiotic and biotic factors, climate, and geography
make up biomes like tundra or tropical rainforest.[108]

Herbivores eat plants, but plants can defend themselves and some species are parasitic or even
carnivorous. Other organisms form mutually beneficial relationships with plants. For example,
mycorrhizal fungi and rhizobia provide plants with nutrients in exchange for food, ants are recruited by
ant plants to provide protection,[109] honey bees, bats and other animals pollinate flowers[110][111] and
humans and other animals[112] act as dispersal vectors to spread spores and seeds.

Plants, climate and environmental change

Plant responses to climate and other environmental changes can inform our understanding of how these
changes affect ecosystem function and productivity. For example, plant phenology can be a useful proxy
for temperature in historical climatology, and the biological impact of climate change and global
warming. Palynology, the analysis of fossil pollen deposits in sediments from thousands or millions of
years ago allows the reconstruction of past climates.[113] Estimates of atmospheric CO2 concentrations
since the Palaeozoic have been obtained from stomatal densities and the leaf shapes and sizes of ancient
land plants.[114] Ozone depletion can expose plants to higher levels of ultraviolet radiation-B (UV-B),
resulting in lower growth rates.[115] Moreover, information from studies of community ecology, plant
systematics, and taxonomy is essential to understanding vegetation change, habitat destruction and
species extinction.[116]

Genetics
Inheritance in plants follows the same fundamental principles
of genetics as in other multicellular organisms. Gregor Mendel
discovered the genetic laws of inheritance by studying
inherited traits such as shape in Pisum sativum (peas). What
Mendel learned from studying plants has had far-reaching
benefits outside of botany. Similarly, "jumping genes" were
discovered by Barbara McClintock while she was studying
maize.[117] Nevertheless, there are some distinctive genetic
differences between plants and other organisms.

Species boundaries in plants may be weaker than in animals,

and cross species hybrids are often possible. A familiar
example is peppermint, Mentha × piperita, a sterile hybrid
A Punnett square depicting a
between Mentha aquatica and spearmint, Mentha spicata.[118]
cross between two pea plants
The many cultivated varieties of wheat are the result of
heterozygous for purple (B) and
multiple inter- and intra-specific crosses between wild species
white (b) blossoms
and their hybrids.[119] Angiosperms with monoecious flowers
often have self-incompatibility mechanisms that operate
between the pollen and stigma so that the pollen either fails to reach the stigma or fails to germinate and
produce male gametes.[120] This is one of several methods used by plants to promote outcrossing.[121] In
many land plants the male and female gametes are produced by separate individuals. These species are
said to be dioecious when referring to vascular plant sporophytes and dioicous when referring to
bryophyte gametophytes.[122]

Charles Darwin in his 1878 book The Effects of Cross and Self-Fertilization in the Vegetable
Kingdom[123] at the start of chapter XII noted "The first and most important of the conclusions which
may be drawn from the observations given in this volume, is that generally cross-fertilisation is beneficial
and self-fertilisation often injurious, at least with the plants on which I experimented." An important
adaptive benefit of outcrossing is that it allows the masking of deleterious mutations in the genome of
progeny. This beneficial effect is also known as hybrid vigor or heterosis. Once outcrossing is
established, subsequent switching to inbreeding becomes disadvantageous since it allows expression of
the previously masked deleterious recessive mutations, commonly referred to as inbreeding depression.

Unlike in higher animals, where parthenogenesis is rare, asexual reproduction may occur in plants by
several different mechanisms. The formation of stem tubers in potato is one example. Particularly in
arctic or alpine habitats, where opportunities for fertilisation of flowers by animals are rare, plantlets or
bulbs, may develop instead of flowers, replacing sexual reproduction with asexual reproduction and
giving rise to clonal populations genetically identical to the parent. This is one of several types of
apomixis that occur in plants. Apomixis can also happen in a seed, producing a seed that contains an
embryo genetically identical to the parent.[124]

Most sexually reproducing organisms are diploid, with paired chromosomes, but doubling of their
chromosome number may occur due to errors in cytokinesis. This can occur early in development to
produce an autopolyploid or partly autopolyploid organism, or during normal processes of cellular
differentiation to produce some cell types that are polyploid (endopolyploidy), or during gamete
formation. An allopolyploid plant may result from a hybridisation event between two different species.
Both autopolyploid and allopolyploid plants can often reproduce normally, but may be unable to cross-
breed successfully with the parent population because there is a mismatch in chromosome numbers.
These plants that are reproductively isolated from the parent species but live within the same
geographical area, may be sufficiently successful to form a new species.[125] Some otherwise sterile plant
polyploids can still reproduce vegetatively or by seed apomixis, forming clonal populations of identical
individuals.[125] Durum wheat is a fertile tetraploid allopolyploid, while bread wheat is a fertile
hexaploid. The commercial banana is an example of a sterile, seedless triploid hybrid. Common
dandelion is a triploid that produces viable seeds by apomictic seed.

As in other eukaryotes, the inheritance of endosymbiotic organelles like mitochondria and chloroplasts in
plants is non-Mendelian. Chloroplasts are inherited through the male parent in gymnosperms but often
through the female parent in flowering plants.[126]

Molecular genetics
A considerable amount of new knowledge about plant function
comes from studies of the molecular genetics of model plants such
as the Thale cress, Arabidopsis thaliana, a weedy species in the
mustard family (Brassicaceae).[93] The genome or hereditary
information contained in the genes of this species is encoded by
about 135 million base pairs of DNA, forming one of the smallest
genomes among flowering plants. Arabidopsis was the first plant
to have its genome sequenced, in 2000.[127] The sequencing of
some other relatively small genomes, of rice (Oryza sativa)[128] Thale cress, Arabidopsis thaliana,
and Brachypodium distachyon,[129] has made them important the first plant to have its genome
model species for understanding the genetics, cellular and sequenced, remains the most
important model organism.
molecular biology of cereals, grasses and monocots generally.

Model plants such as Arabidopsis thaliana are used for studying

the molecular biology of plant cells and the chloroplast. Ideally, these organisms have small genomes that
are well known or completely sequenced, small stature and short generation times. Corn has been used to
study mechanisms of photosynthesis and phloem loading of sugar in C4 plants.[130] The single celled
green alga Chlamydomonas reinhardtii, while not an embryophyte itself, contains a green-pigmented
chloroplast related to that of land plants, making it useful for study.[131] A red alga Cyanidioschyzon
merolae has also been used to study some basic chloroplast functions.[132] Spinach,[133] peas,[134]
soybeans and a moss Physcomitrella patens are commonly used to study plant cell biology.[135]
Agrobacterium tumefaciens, a soil rhizosphere bacterium, can attach to plant cells and infect them with a
callus-inducing Ti plasmid by horizontal gene transfer, causing a callus infection called crown gall
disease. Schell and Van Montagu (1977) hypothesised that the Ti plasmid could be a natural vector for
introducing the Nif gene responsible for nitrogen fixation in the root nodules of legumes and other plant
species.[136] Today, genetic modification of the Ti plasmid is one of the main techniques for introduction
of transgenes to plants and the creation of genetically modified crops.

Epigenetics
Epigenetics is the study of heritable changes in gene function that cannot be explained by changes in the
underlying DNA sequence[137] but cause the organism's genes to behave (or "express themselves")
differently.[138] One example of epigenetic change is the marking of the genes by DNA methylation
which determines whether they will be expressed or not. Gene expression can also be controlled by
repressor proteins that attach to silencer regions of the DNA and prevent that region of the DNA code
from being expressed. Epigenetic marks may be added or removed from the DNA during programmed
stages of development of the plant, and are responsible, for example, for the differences between anthers,
petals and normal leaves, despite the fact that they all have the same underlying genetic code. Epigenetic
changes may be temporary or may remain through successive cell divisions for the remainder of the cell's
life. Some epigenetic changes have been shown to be heritable,[139] while others are reset in the germ
cells.

Epigenetic changes in eukaryotic biology serve to regulate the process of cellular differentiation. During
morphogenesis, totipotent stem cells become the various pluripotent cell lines of the embryo, which in
turn become fully differentiated cells. A single fertilised egg cell, the zygote, gives rise to the many
different plant cell types including parenchyma, xylem vessel elements, phloem sieve tubes, guard cells
of the epidermis, etc. as it continues to divide. The process results from the epigenetic activation of some
genes and inhibition of others.[140]

Unlike animals, many plant cells, particularly those of the parenchyma, do not terminally differentiate,
remaining totipotent with the ability to give rise to a new individual plant. Exceptions include highly
lignified cells, the sclerenchyma and xylem which are dead at maturity, and the phloem sieve tubes which
lack nuclei. While plants use many of the same epigenetic mechanisms as animals, such as chromatin
remodelling, an alternative hypothesis is that plants set their gene expression patterns using positional
information from the environment and surrounding cells to determine their developmental fate.[141]

Epigenetic changes can lead to paramutations, which do not follow the Mendelian heritage rules. These
epigenetic marks are carried from one generation to the next, with one allele inducing a change on the
other.[142]

Plant evolution
The chloroplasts of plants have a number of biochemical, structural and genetic similarities to
cyanobacteria, (commonly but incorrectly known as "blue-green algae") and are thought to be derived
from an ancient endosymbiotic relationship between an ancestral eukaryotic cell and a cyanobacterial
resident.[143][144][145][146]
 The algae are a polyphyletic group and are placed in various
divisions, some more closely related to plants than others. There
are many differences between them in features such as cell wall
composition, biochemistry, pigmentation, chloroplast structure and
nutrient reserves. The algal division Charophyta, sister to the
green algal division Chlorophyta, is considered to contain the
ancestor of true plants.[147] The Charophyte class Charophyceae
and the land plant sub-kingdom Embryophyta together form the
monophyletic group or clade Streptophytina.[148]

Nonvascular land plants are embryophytes that lack the vascular

tissues xylem and phloem. They include mosses, liverworts and
Transverse section of a fossil stem hornworts. Pteridophytic vascular plants with true xylem and
of the Devonian vascular plant phloem that reproduced by spores germinating into free-living
Rhynia gwynne-vaughani
gametophytes evolved during the Silurian period and diversified
into several lineages during the late Silurian and early Devonian.
Representatives of the lycopods have survived to the present day. By the end of the Devonian period,
several groups, including the lycopods, sphenophylls and progymnosperms, had independently evolved
"megaspory" – their spores were of two distinct sizes, larger megaspores and smaller microspores. Their
reduced gametophytes developed from megaspores retained within the spore-producing organs
(megasporangia) of the sporophyte, a condition known as endospory. Seeds consist of an endosporic
megasporangium surrounded by one or two sheathing layers (integuments). The young sporophyte
develops within the seed, which on germination splits to release it. The earliest known seed plants date
from the latest Devonian Famennian stage.[149][150] Following the evolution of the seed habit, seed plants
diversified, giving rise to a number of now-extinct groups, including seed ferns, as well as the modern
gymnosperms and angiosperms.[151] Gymnosperms produce "naked seeds" not fully enclosed in an
ovary; modern representatives include conifers, cycads, Ginkgo, and Gnetales. Angiosperms produce
seeds enclosed in a structure such as a carpel or an ovary.[152][153] Ongoing research on the molecular
phylogenetics of living plants appears to show that the angiosperms are a sister clade to the
gymnosperms.[154]

Plant physiology
Plant physiology encompasses all the internal chemical and physical activities of plants associated with
life.[155] Chemicals obtained from the air, soil and water form the basis of all plant metabolism. The
energy of sunlight, captured by oxygenic photosynthesis and released by cellular respiration, is the basis
of almost all life. Photoautotrophs, including all green plants, algae and cyanobacteria gather energy
directly from sunlight by photosynthesis. Heterotrophs including all animals, all fungi, all completely
parasitic plants, and non-photosynthetic bacteria take in organic molecules produced by photoautotrophs
and respire them or use them in the construction of cells and tissues.[156] Respiration is the oxidation of
carbon compounds by breaking them down into simpler structures to release the energy they contain,
essentially the opposite of photosynthesis.[157]

Molecules are moved within plants by transport processes that operate at a variety of spatial scales.
Subcellular transport of ions, electrons and molecules such as water and enzymes occurs across cell
membranes. Minerals and water are transported from roots to other parts of the plant in the transpiration
stream. Diffusion, osmosis, and active transport and mass
flow are all different ways transport can occur.[158]
Examples of elements that plants need to transport are
nitrogen, phosphorus, potassium, calcium, magnesium,
and sulfur. In vascular plants, these elements are
extracted from the soil as soluble ions by the roots and
transported throughout the plant in the xylem. Most of
the elements required for plant nutrition come from the
chemical breakdown of soil minerals.[159] Sucrose
produced by photosynthesis is transported from the
leaves to other parts of the plant in the phloem and plant
hormones are transported by a variety of processes. Five of the key areas of study within plant
physiology

Plant hormones
Plants are not passive, but respond to external signals
such as light, touch, and injury by moving or growing
towards or away from the stimulus, as appropriate.
Tangible evidence of touch sensitivity is the almost
instantaneous collapse of leaflets of Mimosa pudica, the
insect traps of Venus flytrap and bladderworts, and the
pollinia of orchids.[161]
1 An oat coleoptile with the sun overhead.
The hypothesis that plant growth and development is Auxin (pink) is evenly distributed in its tip.
coordinated by plant hormones or plant growth 2 With the sun at an angle and only shining on
regulators first emerged in the late 19th century. Darwin one side of the shoot, auxin moves to the
experimented on the movements of plant shoots and opposite side and stimulates cell elongation
roots towards light [162] and gravity, and concluded "It is there.
hardly an exaggeration to say that the tip of the radicle . . 3 and 4 Extra growth on that side causes the

acts like the brain of one of the lower animals . . shoot to bend towards the sun.[160]

directing the several movements".[163] About the same

time, the role of auxins (from the Greek auxein, to grow) in control of plant growth was first outlined by
the Dutch scientist Frits Went.[164] The first known auxin, indole-3-acetic acid (IAA), which promotes
cell growth, was only isolated from plants about 50 years later.[165] This compound mediates the tropic
responses of shoots and roots towards light and gravity.[166] The finding in 1939 that plant callus could be
maintained in culture containing IAA, followed by the observation in 1947 that it could be induced to
form roots and shoots by controlling the concentration of growth hormones were key steps in the
development of plant biotechnology and genetic modification.[167]

Cytokinins are a class of plant hormones named for their control of cell division (especially cytokinesis).
The natural cytokinin zeatin was discovered in corn, Zea mays, and is a derivative of the purine adenine.
Zeatin is produced in roots and transported to shoots in the xylem where it promotes cell division, bud
development, and the greening of chloroplasts.[168][169] The gibberelins, such as gibberelic acid are
diterpenes synthesised from acetyl CoA via the mevalonate pathway. They are involved in the promotion
of germination and dormancy-breaking in seeds, in regulation of plant height by controlling stem
elongation and the control of flowering.[170] Abscisic acid (ABA) occurs in all land plants except
liverworts, and is synthesised from carotenoids in the chloroplasts and other plastids. It inhibits cell
division, promotes seed maturation, and dormancy, and
promotes stomatal closure. It was so named because it
was originally thought to control abscission.[171]
Ethylene is a gaseous hormone that is produced in all
higher plant tissues from methionine. It is now known to
be the hormone that stimulates or regulates fruit ripening
and abscission,[172][173] and it, or the synthetic growth
regulator ethephon which is rapidly metabolised to
0:00
produce ethylene, are used on industrial scale to promote
ripening of cotton, pineapples and other climacteric
crops. Venus's fly trap, Dionaea muscipula, showing
the touch-sensitive insect trap in action
Another class of phytohormones is the jasmonates, first
isolated from the oil of Jasminum grandiflorum[174]
which regulates wound responses in plants by unblocking the expression of genes required in the
systemic acquired resistance response to pathogen attack.[175]

In addition to being the primary energy source for plants, light functions as a signalling device, providing
information to the plant, such as how much sunlight the plant receives each day. This can result in
adaptive changes in a process known as photomorphogenesis. Phytochromes are the photoreceptors in a
plant that are sensitive to light.[176]

Plant anatomy and morphology

Plant anatomy is the study of the structure of plant cells and
tissues, whereas plant morphology is the study of their external
form.[177] All plants are multicellular eukaryotes, their DNA
stored in nuclei.[178][179] The characteristic features of plant cells
that distinguish them from those of animals and fungi include a
primary cell wall composed of the polysaccharides cellulose,
hemicellulose and pectin,[180] larger vacuoles than in animal cells
and the presence of plastids with unique photosynthetic and
biosynthetic functions as in the chloroplasts. Other plastids
contain storage products such as starch (amyloplasts) or lipids
(elaioplasts). Uniquely, streptophyte cells and those of the green
algal order Trentepohliales[181] divide by construction of a
phragmoplast as a template for building a cell plate late in cell
division.[84]
A nineteenth-century illustration
The bodies of vascular plants including clubmosses, ferns and showing the morphology of the
seed plants (gymnosperms and angiosperms) generally have aerial roots, stems, leaves and flowers of
and subterranean subsystems. The shoots consist of stems bearing the rice plant Oryza sativa
green photosynthesising leaves and reproductive structures. The
underground vascularised roots bear root hairs at their tips and
generally lack chlorophyll.[183] Non-vascular plants, the liverworts, hornworts and mosses do not produce
 ground-penetrating vascular roots and most of the plant
participates in photosynthesis.[184] The sporophyte
generation is nonphotosynthetic in liverworts but may be
able to contribute part of its energy needs by
photosynthesis in mosses and hornworts.[185]

The root system and the shoot system are interdependent –

the usually nonphotosynthetic root system depends on the
shoot system for food, and the usually photosynthetic shoot
system depends on water and minerals from the root
system.[183] Cells in each system are capable of creating
cells of the other and producing adventitious shoots or
roots.[186] Stolons and tubers are examples of shoots that
can grow roots.[187] Roots that spread out close to the
surface, such as those of willows, can produce shoots and
ultimately new plants.[188] In the event that one of the
systems is lost, the other can often regrow it. In fact it is
possible to grow an entire plant from a single leaf, as is the
case with plants in Streptocarpus sect. Saintpaulia,[189] or
A diagram of a "typical" eudicot, the
even a single cell – which can dedifferentiate into a callus
most common type of plant (three-
(a mass of unspecialised cells) that can grow into a new
fifths of all plant species).[182]
plant.[186] In vascular plants, the xylem and phloem are the
However, no plant actually looks
exactly like this.
conductive tissues that transport resources between shoots
and roots. Roots are often adapted to store food such as
sugars or starch,[183] as in sugar beets and carrots.[188]

Stems mainly provide support to the leaves and reproductive structures, but can store water in succulent
plants such as cacti, food as in potato tubers, or reproduce vegetatively as in the stolons of strawberry
plants or in the process of layering.[190] Leaves gather sunlight and carry out photosynthesis.[191] Large,
flat, flexible, green leaves are called foliage leaves.[192] Gymnosperms, such as conifers, cycads, Ginkgo,
and gnetophytes are seed-producing plants with open seeds.[193] Angiosperms are seed-producing plants
that produce flowers and have enclosed seeds.[152] Woody plants, such as azaleas and oaks, undergo a
secondary growth phase resulting in two additional types of tissues: wood (secondary xylem) and bark
(secondary phloem and cork). All gymnosperms and many angiosperms are woody plants.[194] Some
plants reproduce sexually, some asexually, and some via both means.[195]

Although reference to major morphological categories such as root, stem, leaf, and trichome are useful,
one has to keep in mind that these categories are linked through intermediate forms so that a continuum
between the categories results.[196] Furthermore, structures can be seen as processes, that is, process
combinations.[49]

Systematic botany
Systematic botany is part of systematic biology, which is concerned with the range and diversity of
organisms and their relationships, particularly as determined by their evolutionary history.[197] It
involves, or is related to, biological classification, scientific taxonomy and phylogenetics. Biological
 classification is the method by which botanists group
organisms into categories such as genera or species.
Biological classification is a form of scientific taxonomy.
Modern taxonomy is rooted in the work of Carl
Linnaeus, who grouped species according to shared
physical characteristics. These groupings have since been
revised to align better with the Darwinian principle of
common descent – grouping organisms by ancestry
rather than superficial characteristics. While scientists do
not always agree on how to classify organisms,
molecular phylogenetics, which uses DNA sequences as
data, has driven many recent revisions along
A botanist preparing a plant specimen for evolutionary lines and is likely to continue to do so. The
mounting in the herbarium dominant classification system is called Linnaean
taxonomy. It includes ranks and binomial nomenclature.
The nomenclature of botanical organisms is codified in
the International Code of Nomenclature for algae, fungi, and plants (ICN) and administered by the
International Botanical Congress.[198][199]

Kingdom Plantae belongs to Domain Eukaryota and is broken down recursively until each species is
separately classified. The order is: Kingdom; Phylum (or Division); Class; Order; Family; Genus (plural
genera); Species. The scientific name of a plant represents its genus and its species within the genus,
resulting in a single worldwide name for each organism.[199] For example, the tiger lily is Lilium
columbianum. Lilium is the genus, and columbianum the specific epithet. The combination is the name of
the species. When writing the scientific name of an organism, it is proper to capitalise the first letter in the
genus and put all of the specific epithet in lowercase. Additionally, the entire term is ordinarily italicised
(or underlined when italics are not available).[200][201][202]

The evolutionary relationships and heredity of a group of organisms is called its phylogeny. Phylogenetic
studies attempt to discover phylogenies. The basic approach is to use similarities based on shared
inheritance to determine relationships.[203] As an example, species of Pereskia are trees or bushes with
prominent leaves. They do not obviously resemble a typical leafless cactus such as an Echinocactus.
However, both Pereskia and Echinocactus have spines produced from areoles (highly specialised pad-like
structures) suggesting that the two genera are indeed related.[204][205]

Two cacti of very different appearance

Pereskia aculeata Echinocactus grusonii

Although Pereskia is a tree with leaves, it has spines and areoles like a more typical cactus, such as
Echinocactus.
Judging relationships based on shared characters requires care, since plants may resemble one another
through convergent evolution in which characters have arisen independently. Some euphorbias have
leafless, rounded bodies adapted to water conservation similar to those of globular cacti, but characters
such as the structure of their flowers make it clear that the two groups are not closely related. The
cladistic method takes a systematic approach to characters, distinguishing between those that carry no
information about shared evolutionary history – such as those evolved separately in different groups
(homoplasies) or those left over from ancestors (plesiomorphies) – and derived characters, which have
been passed down from innovations in a shared ancestor (apomorphies). Only derived characters, such as
the spine-producing areoles of cacti, provide evidence for descent from a common ancestor. The results of
cladistic analyses are expressed as cladograms: tree-like diagrams showing the pattern of evolutionary
branching and descent.[206]

From the 1990s onwards, the predominant approach to constructing phylogenies for living plants has
been molecular phylogenetics, which uses molecular characters, particularly DNA sequences, rather than
morphological characters like the presence or absence of spines and areoles. The difference is that the
genetic code itself is used to decide evolutionary relationships, instead of being used indirectly via the
characters it gives rise to. Clive Stace describes this as having "direct access to the genetic basis of
evolution."[207] As a simple example, prior to the use of genetic evidence, fungi were thought either to be
plants or to be more closely related to plants than animals. Genetic evidence suggests that the true
evolutionary relationship of multicelled organisms is as shown in the cladogram below – fungi are more
closely related to animals than to plants.[208]

⁠ plants In 1998, the Angiosperm Phylogeny Group published a phylogeny for

⁠ flowering plants based on an analysis of DNA sequences from most
⁠ ⁠ fungi families of flowering plants. As a result of this work, many questions,
⁠ ⁠ ⁠ such as which families represent the earliest branches of angiosperms,
⁠ ⁠ animals have now been answered.[54] Investigating how plant species are
⁠ related to each other allows botanists to better understand the process of
evolution in plants. [209] Despite the study of model plants and increasing use of DNA evidence, there is
ongoing work and discussion among taxonomists about how best to classify plants into various taxa.[210]
Technological developments such as computers and electron microscopes have greatly increased the level
of detail studied and speed at which data can be analysed.[211]

Symbols
A few symbols are in current use in botany. A number of others are obsolete; for example, Linnaeus used
planetary symbols ⟨ ♂ ⟩ (Mars) for biennial plants, ⟨♃⟩ (Jupiter) for herbaceous perennials and ⟨♄⟩
(Saturn) for woody perennials, based on the planets' orbital periods of 2, 12 and 30 years; and Willd used
⟨♄⟩ (Saturn) for neuter in addition to ⟨☿⟩ (Mercury) for hermaphroditic.[212] The following symbols are
still used:[213]

♀ female ⚇ biennial
♂ male ♾ perennial
⚥ hermaphrodite/bisexual ☠ poisonous
⚲ vegetative (asexual) reproduction 🛈 further information
◊ sex unknown
× crossbred hybrid
☉ annual
+ grafted hybrid
See also
Branches of botany List of botanists by author abbreviation
Evolution of plants List of domesticated plants
Glossary of botanical terms List of flowers
Glossary of plant morphology List of systems of plant taxonomy
List of botany journals Outline of botany
List of botanists Timeline of British botany
List of botanical gardens

Plants portal

Biology portal

Notes
a. Chlorophyll b is also found in some cyanobacteria. A bunch of other chlorophylls exist in
cyanobacteria and certain algal groups, but none of them are found in land plants.[81][82][83]

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External links
Media related to Botany at Wikimedia Commons

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