Botany, also called , plant biology or phytology, is the
science of
plant life and a branch of
biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
. A botanist, plant scientist or phytologist is a
scientist who specialises in this field. The term "botany" comes from the
Ancient Greek word (''botanē'') meaning "
pasture
Pasture (from the Latin ''pastus'', past participle of ''pascere'', "to feed") is land used for grazing. Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, cattle, sheep, or swin ...
", "
herbs" "
grass
Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns and ...
", or "
fodder
Fodder (), also called provender (), is any agricultural foodstuff used specifically to feed domesticated livestock, such as cattle, rabbits, sheep, horses, chickens and pigs. "Fodder" refers particularly to food given to the animals (includin ...
"; is in turn derived from (), "to feed" or "to
graze". Traditionally, botany has also included the study of
fungi and
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
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
International Botanical Congress (IBC) is an international meeting of botanists in all scientific fields, authorized by the International Association of Botanical and Mycological Societies (IABMS) and held every six years, with the location rotati ...
. Nowadays, botanists (in the strict sense) study approximately 410,000
species
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
of
land plants
The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as siste ...
of which some 391,000 species are
vascular plants (including approximately 369,000 species of
flowering plants), and approximately 20,000 are
bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in s ...
s.
Botany originated in prehistory as
herbalism with the efforts of early humans to identify – and later cultivate – edible, medicinal and poisonous plants, making it one of the oldest branches of science. Medieval
physic gardens, often attached to
monasteries
A monastery is a building or complex of buildings comprising the domestic quarters and workplaces of monastics, monks or nuns, whether living in communities or alone (hermits). A monastery generally includes a place reserved for prayer which m ...
, contained plants of medical importance. 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
Plant taxonomy is the science that finds, identifies, describes, classifies, and names plants. It is one of the main branches of taxonomy (the science that finds, describes, classifies, and names living things).
Plant taxonomy is closely allied ...
, 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
Live-cell imaging is the study of living cells using time-lapse microscopy. It is used by scientists to obtain a better understanding of biological function through the study of cellular dynamics. Live-cell imaging was pioneered in the first d ...
,
electron microscopy
An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a hi ...
, analysis of
chromosome number
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectivel ...
,
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
Genomics is an interdisciplinary field of biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, three-dim ...
and
proteomics
Proteomics is the large-scale study of proteins. Proteins are vital parts of living organisms, with many functions such as the formation of structural fibers of muscle tissue, enzymatic digestion of food, or synthesis and replication of DNA. I ...
and
DNA sequences to classify plants more accurately.
Modern botany is a broad, multidisciplinary 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
Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
and
primary metabolism
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
,
chemical products
Products are the species formed from chemical reactions. During a chemical reaction, reactants are transformed into products after passing through a high energy transition state. This process results in the consumption of the reactants. It can be a ...
,
development
Development or developing may refer to:
Arts
*Development hell, when a project is stuck in development
*Filmmaking, development phase, including finance and budgeting
*Development (music), the process thematic material is reshaped
*Photographi ...
,
diseases,
evolutionary relationships
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
,
systematics, and
plant taxonomy
Plant taxonomy is the science that finds, identifies, describes, classifies, and names plants. It is one of the main branches of taxonomy (the science that finds, describes, classifies, and names living things).
Plant taxonomy is closely allied ...
. Dominant themes in 21st century plant science are
molecular genetics
Molecular genetics is a sub-field of biology that addresses how differences in the structures or expression of DNA molecules manifests as variation among organisms. Molecular genetics often applies an "investigative approach" to determine the ...
and
epigenetics
In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are "o ...
, 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
A staple food, food staple, or simply a staple, is a food that is eaten often and in such quantities that it constitutes a dominant portion of a standard diet for a given person or group of people, supplying a large fraction of energy needs and ...
, materials such as
timber,
oil
An oil is any nonpolar chemical substance that is composed primarily of hydrocarbons and is hydrophobic (does not mix with water) & lipophilic (mixes with other oils). Oils are usually flammable and surface active. Most oils are unsaturated ...
, rubber,
fibre
Fiber or fibre (from la, fibra, links=no) is a natural or artificial substance that is significantly longer than it is wide. Fibers are often used in the manufacture of other materials. The strongest engineering materials often incorporate ...
and drugs, in modern
horticulture,
agriculture
Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
and
forestry,
plant propagation,
breeding
Breeding is sexual reproduction that produces offspring, usually animals or plants. It can only occur between a male and a female animal or plant.
Breeding may refer to:
* Animal husbandry, through selected specimens such as dogs, horses, and rab ...
and
genetic modification
Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including ...
, in the synthesis of chemicals and raw materials for construction and energy production, in
environmental management
Environmental resource management is the management of the interaction and impact of human societies on the environment. It is not, as the phrase might suggest, the management of the environment itself. Environmental resources management aims ...
, and the maintenance of
biodiversity
Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
.
History
Early botany
Botany originated as
herbalism, the study and use of plants for their medicinal properties. 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,
Ancient Egypt,
[Manniche, Lisa; An Ancient Egyptian Herbal; American University in Cairo Press; Cairo; 2006; ] in archaic
Avestan
Avestan (), or historically Zend, is an umbrella term for two Old Iranian languages: Old Avestan (spoken in the 2nd millennium BCE) and Younger Avestan (spoken in the 1st millennium BCE). They are known only from their conjoined use as the scri ...
writings, and in works from China purportedly from before 221 BCE.
Modern botany traces its roots back to
Ancient Greece
Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of classical antiquity ( AD 600), that comprised a loose collection of cult ...
specifically to
Theophrastus (–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". His major works, ''
Enquiry into Plants
Theophrastus's ''Enquiry into Plants'' or ''Historia Plantarum'' ( grc-gre, Περὶ φυτῶν ἱστορία, ''Peri phyton historia'') was, along with his mentor Aristotle's ''History of Animals'', Pliny the Elder's '' Natural History'' a ...
'' and ''On the Causes of Plants'', constitute the most important contributions to botanical science until the
Middle Ages, almost seventeen centuries later.
Another work from Ancient Greece that made an early impact on botany is ''
De materia medica
(Latin name for the Greek work , , both meaning "On Medical Material") is a pharmacopoeia of medicinal plants and the medicines that can be obtained from them. The five-volume work was written between 50 and 70 CE by Pedanius Dioscorides, a ...
'', a five-volume encyclopedia about
herbal medicine written in the middle of the first century by Greek physician and pharmacologist
Pedanius Dioscorides
Pedanius Dioscorides ( grc-gre, Πεδάνιος Διοσκουρίδης, ; 40–90 AD), “the father of pharmacognosy”, was a Greek physician, pharmacologist, botanist, and author of '' De materia medica'' (, On Medical Material) —a 5-vo ...
. ''De materia medica'' was widely read for more than 1,500 years. Important contributions from the
medieval Muslim world include
Ibn Wahshiyya
( ar, ابن وحشية), died , was a Nabataean (Aramaic-speaking, rural Iraqi) agriculturalist, toxicologist, and alchemist born in Qussīn, near Kufa in Iraq. He is the author of the ''Nabataean Agriculture'' (), an influential Arabic work o ...
's ''
Nabatean Agriculture'',
Abū Ḥanīfa Dīnawarī
Abū Ḥanīfa Aḥmad ibn Dāwūd Dīnawarī ( fa, ابوحنيفه دينوری; died 895) was a Persian Islamic Golden Age polymath, astronomer, agriculturist, botanist, metallurgist, geographer, mathematician, and historian.
Life
Dinaw ...
's (828–896) the ''Book of Plants'', and
Ibn Bassal
Ibn Bassal ( ar, ابن بصال) was an 11th-century Andalusian Arab botanist and agronomist in Toledo and Seville, Spain who wrote about horticulture and arboriculture. He is best known for his book on agronomy, the ''Dīwān al-filāha'' (An ...
's ''The Classification of Soils''. In the early 13th century,
Abu al-Abbas al-Nabati
Ahmad bin Muhammad bin Mufarraj bin Ani al-Khalil, better known as Abu al-Abbas al-Nabati, Ibn al-Rumiya or al-Ashshab, ( ar, أبو العباس النباتي, ''Abu’l-ʿAbbās al-Nabātī'') (c. 1200) was an Andalusian scientist, botan ...
, and
Ibn al-Baitar
Diyāʾ al-Dīn Abū Muḥammad ʿAbd Allāh ibn Aḥmad al-Mālaqī, commonly known as Ibn al-Bayṭār () (1197–1248 AD) was an Andalusian Arab physician, botanist, pharmacist and scientist. His main contribution was to systematically record ...
(d. 1248) wrote on botany in a systematic and scientific manner.
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 medical use. They supported the growth of botany as an academic subject. Lectures were given about the plants grown in the gardens and their medical uses demonstrated. Botanical gardens came much later to northern Europe; the first in England was the
University of Oxford Botanic Garden
The University of Oxford Botanic Garden is the oldest botanic garden in Great Britain and one of the oldest scientific gardens in the world. The garden was founded in 1621 as a physic garden growing plants for medicinal research. Today it conta ...
in 1621. Throughout this period, botany remained firmly subordinate to medicine.
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
Hieronymus Bock ( Latinised Hieronymus Tragus; c. 1498 – 21 February 1554) was a German botanist, physician, and Lutheran minister who began the transition from medieval botany to the modern scientific worldview by arranging plants by their re ...
(1498–1554) (also called Hieronymus Tragus). 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
Valerius Cordus (18 February 1515 – 25 September 1544) was a German physician, botanist and pharmacologist who authored the first pharmacopoeia North of the Alps and one of the most celebrated herbals in history. He is also widely credited ...
(1515–1544) authored a botanically and pharmacologically important herbal ''Historia Plantarum'' in 1544 and a
pharmacopoeia of lasting importance, the ''Dispensatorium'' in 1546. Naturalist
Conrad von Gesner
Conrad Gessner (; la, Conradus Gesnerus 26 March 1516 – 13 December 1565) was a Swiss physician, naturalist, bibliographer, and philologist. Born into a poor family in Zürich, Switzerland, his father and teachers quickly realised his tale ...
(1516–1565) and herbalist
John Gerard
John Gerard (also John Gerarde, c. 1545–1612) was an English herbalist with a large garden in Holborn, now part of London. His 1,484-page illustrated ''Herball, or Generall Historie of Plantes'', first published in 1597, became a popular gard ...
(1545–) published herbals covering the medicinal uses of plants. Naturalist
Ulisse Aldrovandi
Ulisse Aldrovandi (11 September 1522 – 4 May 1605) was an Italian naturalist, the moving force behind Bologna's botanical garden, one of the first in Europe. Carl Linnaeus and the comte de Buffon reckoned him the father of natural history ...
(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
Cell most often refers to:
* Cell (biology), the functional basic unit of life
Cell may also refer to:
Locations
* Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
, a term he coined, in
cork
Cork or CORK may refer to:
Materials
* Cork (material), an impermeable buoyant plant product
** Cork (plug), a cylindrical or conical object used to seal a container
***Wine cork
Places Ireland
* Cork (city)
** Metropolitan Cork, also known as G ...
, and a short time later in living plant tissue.
Early modern botany
During the 18th century, systems of
plant identification
In biology, determination is the process of matching a specimen of an organism to a known taxon, for example identifying a plant. The term is also used in cellular biology, where it means the act of the differentiation of stem cells becoming fix ...
were developed comparable to
dichotomous keys
In phylogenetics, a single-access key (also called dichotomous key, sequential key, analytical key, or pathway key) is an identification key where the sequence and structure of identification steps is fixed by the author of the key. At each point i ...
, where unidentified plants are placed into
taxonomic groups (e.g. family, genus and species) by making a series of choices between pairs of
characters
Character or Characters may refer to:
Arts, entertainment, and media Literature
* ''Character'' (novel), a 1936 Dutch novel by Ferdinand Bordewijk
* ''Characters'' (Theophrastus), a classical Greek set of character sketches attributed to The ...
. 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 biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
in synoptic keys. By the 18th century, new plants for study were arriving in Europe in increasing numbers from newly discovered countries and the European colonies worldwide. In 1753,
Carl von Linné
Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
(Carl Linnaeus) published his
Species Plantarum, a hierarchical 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
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
within the genus. 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.
Increasing knowledge of
plant anatomy
Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plant ...
,
morphology
Morphology, from the Greek and meaning "study of shape", may refer to:
Disciplines
*Morphology (archaeology), study of the shapes or forms of artifacts
*Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
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
De Jussieu, the name of a French family which came into prominence towards the close of the sixteenth century, and was known for a century and a half for the botanists it produced. The following are its more eminent members:
*Antoine de Jussieu ( ...
(1789), and
Candolle
Augustin Pyramus (or Pyrame) de Candolle (, , ; 4 February 17789 September 1841) was a Swiss botanist. René Louiche Desfontaines launched de Candolle's botanical career by recommending him at a herbarium. Within a couple of years de 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
A taxonomic system, the Bentham & Hooker system for seed plants, was published in Bentham and Hooker's ''Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita'' in three volumes between 1862 and 1883.
George Bentham (1 ...
, 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.
Botany was greatly stimulated by the appearance of the first "modern" textbook,
Matthias Schleiden
Matthias Jakob Schleiden (; 5 April 1804 – 23 June 1881) was a German botanist and co-founder of cell theory, along with Theodor Schwann and Rudolf Virchow.
Career
Matthias Jakob Schleiden was born in Hamburg. on 5 April 1804. His father was ...
's ', published in English in 1849 as ''Principles of Scientific Botany''. Schleiden was a microscopist and an early plant anatomist who co-founded the
cell theory
In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
with
Theodor Schwann and
Rudolf Virchow and was among the first to grasp the significance of the
cell nucleus
The cell nucleus (pl. nuclei; from Latin or , meaning ''kernel'' or ''seed'') is a membrane-bound organelle found in eukaryotic cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, h ...
that had been described by
Robert Brown in 1831.
In 1855,
Adolf Fick
Adolf Eugen Fick (3 September 1829 – 21 August 1901) was a German-born physician and physiologist.
Early life and education
Fick began his work in the formal study of mathematics and physics before realising an aptitude for medicine. He ...
formulated
Fick's laws that enabled the calculation of the rates of
molecular diffusion
Molecular diffusion, often simply called diffusion, is the thermal motion of all (liquid or gas) particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size (mass) o ...
in biological systems.
Late modern botany
Building upon the gene-chromosome theory of heredity that originated with
Gregor Mendel (1822–1884),
August Weismann
August Friedrich Leopold Weismann FRS (For), HonFRSE, LLD (17 January 18345 November 1914) was a German evolutionary biologist. Fellow German Ernst Mayr ranked him as the second most notable evolutionary theorist of the 19th century, after Cha ...
(1834–1914) proved that inheritance only takes place through
gametes. No other cells can pass on inherited characters. The work of
Katherine Esau
Katherine Esau (3 April 1898 – 4 June 1997) was a German-American botanist who received the National Medal of Science for her work on plant anatomy.
Personal life and education
Esau was born on 3 April 1898 in Yekaterinoslav, Russian Empire ...
(1898–1997) on plant anatomy is still a major foundation of modern botany. Her books ''Plant Anatomy'' and ''Anatomy of Seed Plants'' have been key plant structural biology texts for more than half a century.
The discipline of
plant ecology
Plant ecology is a subdiscipline of ecology which studies the distribution and abundance of plants, the effects of environmental factors upon the abundance of plants, and the interactions among and between plants and other organisms. Examples ...
was pioneered in the late 19th century by botanists such as
Eugenius Warming, who produced the hypothesis that plants form
communities
A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, tow ...
, 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
Ecological succession is the process of change in the species structure of an ecological community over time. The time scale can be decades (for example, after a wildfire) or more or less.
Bacteria allows for the cycling of nutrients such as ca ...
was developed by
Henry Chandler Cowles,
Arthur Tansley
Sir Arthur George Tansley FLS, FRS (15 August 1871 – 25 November 1955) was an English botanist
Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phy ...
and
Frederic Clements
Frederic Edward Clements (September 16, 1874 – July 26, 1945) was an American plant ecologist and pioneer in the study of plant ecology and vegetation succession.
Biography
Born in Lincoln, Nebraska, he studied botany at the University of Neb ...
. Clements is credited with the idea of
climax vegetation
In scientific ecology, climax community or climatic climax community is a historic term for a community of plants, animals, and fungi which, through the process of ecological succession in the development of vegetation in an area over time, ha ...
as the most complex vegetation that an environment can support and Tansley introduced the concept of
ecosystems to biology. Building on the extensive earlier work of
Alphonse de Candolle
Alphonse Louis Pierre Pyramus (or Pyrame) de Candolle (28 October 18064 April 1893) was a French-Swiss botanist, the son of the Swiss botanist Augustin Pyramus de Candolle.
Biography
De Candolle, son of Augustin Pyramus de Candolle, first devot ...
,
Nikolai Vavilov (1887–1943) produced accounts of the
biogeography
Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
,
centres of origin, and evolutionary history of economic plants.
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
Molecular diffusion, often simply called diffusion, is the thermal motion of all (liquid or gas) particles at temperatures above absolute zero. The rate of this movement is a function of temperature, viscosity of the fluid and the size (mass) o ...
of water vapour and carbon dioxide through
stomatal
In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
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. Innovations in
statistical analysis by
Ronald Fisher
Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who ...
,
Frank Yates and others at
Rothamsted Experimental Station
Rothamsted Research, previously known as the Rothamsted Experimental Station and then the Institute of Arable Crops Research, is one of the oldest agricultural research institutions in the world, having been founded in 1843. It is located at Har ...
facilitated rational experimental design and data analysis in botanical research. The discovery and identification of the
auxin
Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essenti ...
plant hormones
Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, p ...
by
Kenneth V. Thimann
Kenneth Vivian Thimann (August 5, 1904 – January 15, 1997) was an English-American plant physiologist and microbiologist known for his studies of plant hormones, which were widely influential in agriculture and horticulture. He isolated and de ...
in 1948 enabled regulation of plant growth by externally applied chemicals.
Frederick Campion Steward
Frederick Campion "Camp" Steward FRS (16 June 1904 – 13 September 1993) was a British botanist and plant physiologist.
Early life and education
He was born in Pimlico, London, but brought up in Yorkshire. He was educated at Heckmondwike G ...
pioneered techniques of
micropropagation and
plant tissue culture controlled by plant hormones. The synthetic auxin
2,4-dichlorophenoxyacetic acid or 2,4-D was one of the first commercial synthetic herbicides.
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
Genomics is an interdisciplinary field of biology focusing on the structure, function, evolution, mapping, and editing of genomes. A genome is an organism's complete set of DNA, including all of its genes as well as its hierarchical, three-dim ...
,
proteomics
Proteomics is the large-scale study of proteins. Proteins are vital parts of living organisms, with many functions such as the formation of structural fibers of muscle tissue, enzymatic digestion of food, or synthesis and replication of DNA. I ...
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. The concept originally stated by
Gottlieb Haberlandt
Gottlieb Haberlandt (28 November 1854 – 30 January 1945) was an Austrian botanist. He was the son of European 'soybean' pioneer Professor Friedrich J. Haberlandt. His son Ludwig Haberlandt was an early reproductive physiologist now given credi ...
in 1902 that all plant cells are
totipotent Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta.
According to Latin terms, Pluripotentia means the ability for many thin ...
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
bioreactors
A bioreactor refers to any manufactured device or system that supports a biologically active environment. In one case, a bioreactor is a vessel in which a chemical process is carried out which involves organisms or biochemically active substances ...
to synthesise
pesticides
Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and la ...
,
antibiotics
An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
or other
pharmaceuticals
A medication (also called medicament, medicine, pharmaceutical drug, medicinal drug or simply drug) is a drug used to diagnose, cure, treat, or prevent disease. Drug therapy (pharmacotherapy) is an important part of the medical field and r ...
, as well as the practical application of
genetically modified crops designed for traits such as improved yield.
Modern morphology recognises a continuum between the major morphological categories of root, stem (caulome), leaf (phyllome) and
trichome. Furthermore, it emphasises structural dynamics. Modern systematics aims to reflect and discover
phylogenetic relationships between plants. 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
The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disco ...
to publish in 1998 a
phylogeny of flowering plants, answering many of the questions about relationships among
angiosperm
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
families and species. 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.
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
Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
with the chemical energy they need to exist. Plants,
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
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 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. 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. Plants are crucial to the future of human society as they provide food, oxygen, medicine, and products for people, as well as creating and preserving soil.
Historically, all living things were classified as either animals or plants and botany covered the study of all organisms not considered animals. Botanists examine both the internal functions and processes within plant
organelle
In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' the ...
s, 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
Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
and
morphology
Morphology, from the Greek and meaning "study of shape", may refer to:
Disciplines
*Morphology (archaeology), study of the shapes or forms of artifacts
*Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
), or function (
physiology) of plant life.
The strictest definition of "plant" includes only the "land plants" or
embryophytes, which include
seed plants
A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
(gymnosperms, including the
pines
A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garde ...
, and
flowering plants) and the free-sporing
cryptogams
A cryptogam (scientific name Cryptogamae) is a plant (in the wide sense of the word) or a plant-like organism that reproduces by spores, without flowers or seeds. The name ''Cryptogamae'' () means "hidden reproduction", referring to the fact ...
including
ferns,
clubmosses
Lycopodiopsida is a class of vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching ...
,
liverworts,
hornwort
Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s 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
A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.
Life cycle
The sporophyte develops from the zygote p ...
within its tissues for at least part of its life, even in the seed plants, where the gametophyte itself is nurtured by its parent sporophyte. Other groups of organisms that were previously studied by botanists include bacteria (now studied in
bacteriology Bacteriology is the branch and specialty of biology that studies the morphology, ecology, genetics and biochemistry of bacteria as well as many other aspects related to them. This subdivision of microbiology involves the identification, classificat ...
), fungi (
mycology) – including
lichen-forming fungi (
lichenology
Lichenology is the branch of mycology that studies the lichens, symbiotic organisms made up of an intimate symbiotic association of a microscopic alga (or a cyanobacterium) with a filamentous fungus.
Study of lichens draws knowledge from several ...
), non-
chlorophyte
Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to a ...
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
(
phycology), and viruses (
virology). However, attention is still given to these groups by botanists, and fungi (including lichens) and photosynthetic
protist
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exclu ...
s are usually covered in introductory botany courses.
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
An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship.
(The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
relationship with an early eukaryote, ultimately becoming the
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s 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.
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
In organizational studies, resource management is the efficient and effective development of an organization's resources when they are needed. Such resources may include the financial resources, inventory, human skills, production resources, or i ...
,
conservation,
human food security,
biologically invasive organisms,
carbon sequestration
Carbon sequestration is the process of storing carbon in a carbon pool. Carbon dioxide () is naturally captured from the atmosphere through biological, chemical, and physical processes. These changes can be accelerated through changes in lan ...
,
climate change, and
sustainability.
Human nutrition
Virtually all staple foods come either directly from
primary production by plants, or indirectly from animals that eat them. Plants and other photosynthetic organisms are at the base of most
food chain
A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), det ...
s 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
The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it i ...
. The modern forms of the major
staple food
A staple food, food staple, or simply a staple, is a food that is eaten often and in such quantities that it constitutes a dominant portion of a standard Diet (nutrition), diet for a given person or group of people, supplying a large fraction of ...
s, such as
hemp,
teff, maize, rice, wheat and other cereal grasses,
pulses
In medicine, a pulse represents the tactile arterial palpation of the cardiac cycle (heartbeat) by trained fingertips. The pulse may be palpated in any place that allows an artery to be compressed near the surface of the body, such as at the nec ...
, bananas and plantains, as well as
hemp,
flax
Flax, also known as common flax or linseed, is a flowering plant, ''Linum usitatissimum'', in the family Linaceae. It is cultivated as a food and fiber crop in regions of the world with temperate climates. Textiles made from flax are known in ...
and
cotton grown for their fibres, are the outcome of prehistoric selection over thousands of years from among
wild ancestral plants with the most desirable characteristics.
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
Food security speaks to the availability of food in a country (or geography) and the ability of individuals within that country (geography) to access, afford, and source adequate foodstuffs. According to the United Nations' Committee on World F ...
for future generations. Botanists also study weeds, which are a considerable problem in agriculture, and the biology and control of
plant pathogens
Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oomy ...
in agriculture and natural
ecosystems.
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. Some of the earliest plant-people relationships arose between the
indigenous people
Indigenous peoples are culturally distinct ethnic groups whose members are directly descended from the earliest known inhabitants of a particular geographic region and, to some extent, maintain the language and culture of those original people ...
of Canada in identifying edible plants from inedible plants.
This relationship the indigenous people had with plants was recorded by ethnobotanists.
Plant biochemistry
Plant biochemistry is the study of the chemical processes used by plants. Some of these processes are used in their
primary metabolism
Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
like the photosynthetic
Calvin cycle
The Calvin cycle, light-independent reactions, bio synthetic phase, dark reactions, or photosynthetic carbon reduction (PCR) cycle of photosynthesis is a series of chemical reactions that convert carbon dioxide and hydrogen-carrier compounds into ...
and
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 that allows a plant to photosynthesize during the day, but only exchange gases at night. I ...
. Others make specialised materials like the
cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
and
lignin used to build their bodies, and
secondary products like
resins and
aroma compounds
An aroma compound, also known as an odorant, aroma, fragrance or flavoring, is a chemical compound that has a smell or odor. For an individual chemical or class of chemical compounds to impart a smell or fragrance, it must be sufficiently vol ...
.
Plants and various other groups of photosynthetic eukaryotes collectively known as "
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
" have unique organelles known as
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s. Chloroplasts are thought to be descended from
cyanobacteria that formed
endosymbiotic
An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship.
(The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
relationships with ancient plant and algal ancestors. Chloroplasts and cyanobacteria contain the blue-green pigment
chlorophyll ''a''. Chlorophyll ''a'' (as well as its plant and green algal-specific cousin
chlorophyll ''b'') absorbs light in the blue-violet and orange/red parts of the
spectrum
A spectrum (plural ''spectra'' or ''spectrums'') is a condition that is not limited to a specific set of values but can vary, without gaps, across a continuum. The word was first used scientifically in optics to describe the rainbow of colors ...
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
Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
, a process that generates
molecular oxygen
There are several known allotropes of oxygen. The most familiar is molecular oxygen (O2), present at significant levels in Earth's atmosphere and also known as dioxygen or triplet oxygen. Another is the highly reactive ozone (O3). Others are ...
(O
2) as a by-product.
The light energy captured by
chlorophyll ''a'' is initially in the form of electrons (and later a
proton gradient
An electrochemical gradient is a gradient of electrochemical potential, usually for an ion that can move across a membrane. The gradient consists of two parts, the chemical gradient, or difference in solute concentration across a membrane, and th ...
) that's used to make molecules of
ATP and
NADPH
Nicotinamide adenine dinucleotide phosphate, abbreviated NADP or, in older notation, TPN (triphosphopyridine nucleotide), is a cofactor used in anabolic reactions, such as the Calvin cycle and lipid and nucleic acid syntheses, which require NAD ...
which temporarily store and transport energy. Their energy is used in the
light-independent reactions
The Calvin cycle, light-independent reactions, bio synthetic phase, dark reactions, or photosynthetic carbon reduction (PCR) cycle of photosynthesis is a series of chemical reactions that convert carbon dioxide and hydrogen-carrier compounds into ...
of the Calvin cycle by the enzyme
rubisco to produce molecules of the 3-carbon sugar
glyceraldehyde 3-phosphate
Glyceraldehyde 3-phosphate, also known as triose phosphate or 3-phosphoglyceraldehyde and abbreviated as G3P, GA3P, GADP, GAP, TP, GALP or PGAL, is a metabolite that occurs as an intermediate in several central pathways of all organisms.Nelson, D ...
(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. Starch is the characteristic energy store of most land plants and algae, while
inulin
Inulins are a group of naturally occurring polysaccharides produced by many types of plants, industrially most often extracted from chicory. The inulins belong to a class of dietary fibers known as fructans. Inulin is used by some plants as a me ...
, a polymer of
fructose
Fructose, or fruit sugar, is a ketonic simple sugar found in many plants, where it is often bonded to glucose to form the disaccharide sucrose. It is one of the three dietary monosaccharides, along with glucose and galactose, that are absorbe ...
is used for the same purpose in the sunflower family
Asteraceae
The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae w ...
. 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
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s, including synthesising all their
fatty acids, and most
amino acids
Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
. 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
Cutin is one of two waxy polymers that are the main components of the plant cuticle, which covers all aerial surfaces of plants. It is an insoluble substance with waterproof quality. Cutin also harbors cuticular waxes, which assist in cuticle stru ...
which is found in the
plant cuticle that protects land plants from drying out.
Plants synthesise a number of unique
polymers like the
polysaccharide molecules
cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
,
pectin and
xyloglucan
Xyloglucan is a hemicellulose that occurs in the primary cell wall of all vascular plants; however, all enzymes responsible for xyloglucan metabolism are found in Charophyceae algae.LEV Del Bem and M Vincentz (2010) Evolution of xyloglucan-related ...
from which the land plant cell wall is constructed.
Vascular land plants make
lignin, a polymer used to strengthen the
secondary cell walls of xylem
tracheid
A tracheid is a long and tapered lignified cell in the xylem of vascular plants. It is a type of conductive cell called a tracheary element. Angiosperms use another type of tracheary element, called vessel elements, to transport water through the ...
s and
vessels
Vessel(s) or The Vessel may refer to:
Biology
* Blood vessel, a part of the circulatory system and function to transport blood throughout the body
*Lymphatic vessel, a thin walled, valved structure that carries lymph
*Vessel element, a narrow wa ...
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
270px, SEM image of pollen grains
Sporopollenin is one of the most chemically inert biological polymers. It is a major component of the tough outer (exine) walls of plant spores and pollen grains. It is chemically very stable and is usually well ...
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
The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya.
The ...
period.
The concentration of carbon dioxide in the atmosphere today is much lower than it was when plants emerged onto land during the
Ordovician
The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya.
The ...
and
Silurian periods. Many
monocots
Monocotyledons (), commonly referred to as monocots, ( Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
like
maize and the
pineapple and some
dicots like the
Asteraceae
The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae w ...
have since independently evolved pathways like
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 that allows a plant to photosynthesize during the day, but only exchange gases at night. I ...
and the
carbon fixation pathway for photosynthesis which avoid the losses resulting from
photorespiration in the more common
carbon fixation pathway. These biochemical strategies are unique to land plants.
Medicine and materials
Phytochemistry
Phytochemistry is the study of phytochemicals, which are chemicals derived from plants. Phytochemists strive to describe the structures of the large number of secondary metabolites found in plants, the functions of these compounds in human and ...
is a branch of plant biochemistry primarily concerned with the chemical substances produced by plants during
secondary metabolism
Secondary metabolism (also called specialized metabolism) is a term for pathways and small molecule products of metabolism that are involved in ecological interactions, but are not absolutely required for the survival of the organism. These molecul ...
. Some of these compounds are toxins such as the
alkaloid
Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of similar s ...
coniine
Coniine is a poisonous chemical compound, an alkaloid present in and isolable from poison hemlock (''Conium maculatum''), where its presence has been a source of significant economic, medical, and historico-cultural interest; coniine is also produ ...
from
hemlock. Others, such as the
essential oils
peppermint oil Peppermint extract is an herbal extract of peppermint (''Mentha × piperita'') made from the essential oils of peppermint leaves. Peppermint is a hybrid of water mint and spearmint and was indigenous to Europe and the Middle East before it became ...
and lemon oil are useful for their aroma, as flavourings and spices (e.g.,
capsaicin
Capsaicin (8-methyl-''N''-vanillyl-6-nonenamide) ( or ) is an active component of chili peppers, which are plants belonging to the genus '' Capsicum''. It is a chemical irritant for mammals, including humans, and produces a sensation of burni ...
), and in medicine as pharmaceuticals as in
opium from
opium poppies. Many
medicinal and
recreational drugs, such as
tetrahydrocannabinol
Tetrahydrocannabinol (THC) is the principal psychoactive constituent of cannabis and one of at least 113 total cannabinoids identified on the plant. Although the chemical formula for THC (C21H30O2) describes multiple isomers, the term ''THC'' ...
(active ingredient in
cannabis
''Cannabis'' () is a genus of flowering plants in the family Cannabaceae. The number of species within the genus is disputed. Three species may be recognized: ''Cannabis sativa'', '' C. indica'', and '' C. ruderalis''. Alternativel ...
),
caffeine,
morphine
Morphine is a strong opiate that is found naturally in opium, a dark brown resin in poppies (''Papaver somniferum''). It is mainly used as a pain medication, and is also commonly used recreationally, or to make other illicit opioids. There ...
and
nicotine
Nicotine is a naturally produced alkaloid in the nightshade family of plants (most predominantly in tobacco and ''Duboisia hopwoodii'') and is widely used recreationally as a stimulant and anxiolytic. As a pharmaceutical drug, it is used for ...
come directly from plants. Others are simple
derivatives of botanical natural products. For example, the pain killer
aspirin
Aspirin, also known as acetylsalicylic acid (ASA), is a nonsteroidal anti-inflammatory drug (NSAID) used to reduce pain, fever, and/or inflammation, and as an antithrombotic. Specific inflammatory conditions which aspirin is used to treat in ...
is the acetyl
ester of
salicylic acid
Salicylic acid is an organic compound with the formula HOC6H4CO2H. A colorless, bitter-tasting solid, it is a precursor to and a metabolite of aspirin (acetylsalicylic acid). It is a plant hormone, and has been listed by the EPA Toxic Substa ...
, originally isolated from the
bark
Bark may refer to:
* Bark (botany), an outer layer of a woody plant such as a tree or stick
* Bark (sound), a vocalization of some animals (which is commonly the dog)
Places
* Bark, Germany
* Bark, Warmian-Masurian Voivodeship, Poland
Arts, e ...
of
willow trees, and a wide range of
opiate painkillers like
heroin
Heroin, also known as diacetylmorphine and diamorphine among other names, is a potent opioid mainly used as a recreational drug for its euphoric effects. Medical grade diamorphine is used as a pure hydrochloride salt. Various white and brown ...
are obtained by chemical modification of
morphine
Morphine is a strong opiate that is found naturally in opium, a dark brown resin in poppies (''Papaver somniferum''). It is mainly used as a pain medication, and is also commonly used recreationally, or to make other illicit opioids. There ...
obtained from the
opium poppy. Popular
stimulants come from plants, such as
caffeine from coffee, tea and chocolate, and
nicotine
Nicotine is a naturally produced alkaloid in the nightshade family of plants (most predominantly in tobacco and ''Duboisia hopwoodii'') and is widely used recreationally as a stimulant and anxiolytic. As a pharmaceutical drug, it is used for ...
from tobacco. Most alcoholic beverages come from
fermentation of
carbohydrate-rich plant products such as
barley (beer), rice (
sake) and grapes (wine).
Native Americans have used various plants as ways of treating illness or disease for thousands of years. 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.
Plants can synthesise useful coloured dyes and pigments such as the
anthocyanin
Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical com ...
s 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
Gamboge ( , ) is a partially transparent deep saffron to mustard yellow pigment.Other forms and spellings are: cambodia, cambogium, camboge, cambugium, gambaugium, gambogia, gambozia, gamboidea, gambogium, gumbouge, gambouge, gamboge, gambooge, g ...
and
rose madder.
Sugar,
starch, cotton,
linen,
hemp, some types of
rope, wood and
particle board
Particle board, also known as chipboard or low-density fiberboard, is an engineered wood product manufactured from wood chips and a synthetic resin or other suitable binder, which is pressed and extruded. Particle board is often confused with o ...
s,
papyrus and paper,
vegetable oils,
wax
Waxes are a diverse class of organic compounds that are lipophilic, malleable solids near ambient temperatures. They include higher alkanes and lipids, typically with melting points above about 40 °C (104 °F), melting to give low ...
, and
natural rubber
Rubber, also called India rubber, latex, Amazonian rubber, ''caucho'', or ''caoutchouc'', as initially produced, consists of polymers of the organic compound isoprene, with minor impurities of other organic compounds. Thailand, Malaysia, and ...
are examples of commercially important materials made from plant tissues or their secondary products.
Charcoal
Charcoal is a lightweight black carbon residue produced by strongly heating wood (or other animal and plant materials) in minimal oxygen to remove all water and volatile constituents. In the traditional version of this pyrolysis process, cal ...
, a pure form of carbon made by
pyrolysis of wood, has a long
history as a metal-
smelting
Smelting is a process of applying heat to ore, to extract a base metal. It is a form of extractive metallurgy. It is used to extract many metals from their ores, including silver, iron, copper, and other base metals. Smelting uses heat and a c ...
fuel, as a filter material and
adsorbent
Adsorption is the adhesion of atoms, ions or molecules from a gas, liquid or dissolved solid to a surface. This process creates a film of the ''adsorbate'' on the surface of the ''adsorbent''. This process differs from absorption, in which a f ...
and as an artist's material and is one of the three ingredients of
gunpowder.
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
, the world's most abundant organic polymer, can be converted into energy, fuels, materials and chemical feedstock.
Products made from cellulose include
rayon and
cellophane
Cellophane is a thin, transparent sheet made of regenerated cellulose. Its low permeability to air, oils, greases, bacteria, and liquid water makes it useful for food packaging. Cellophane is highly permeable to water vapour, but may be coat ...
,
wallpaper paste,
biobutanol
220px, Butanol, a C-4 hydrocarbon is a promising bio-derived fuel, which shares many properties with gasoline.
Butanol may be used as a fuel in an internal combustion engine. It is more similar to gasoline than it is to ethanol. A C4-hydrocarbo ...
and
gun cotton
Nitrocellulose (also known as cellulose nitrate, flash paper, flash cotton, guncotton, pyroxylin and flash string, depending on form) is a highly flammable compound formed by nitrating cellulose through exposure to a mixture of nitric acid a ...
.
Sugarcane
Sugarcane or sugar cane is a species of (often hybrid) tall, perennial grass (in the genus '' Saccharum'', tribe Andropogoneae) that is used for sugar production. The plants are 2–6 m (6–20 ft) tall with stout, jointed, fibrous stalks ...
,
rapeseed
Rapeseed (''Brassica napus ''subsp.'' napus''), also known as rape, or oilseed rape, is a bright-yellow flowering member of the family Brassicaceae (mustard or cabbage family), cultivated mainly for its oil-rich seed, which naturally contains ...
and
soy
The soybean, soy bean, or soya bean (''Glycine max'') is a species of legume native to East Asia, widely grown for its edible bean, which has numerous uses.
Traditional unfermented food uses of soybeans include soy milk, from which tofu and ...
are some of the plants with a highly fermentable sugar or oil content that are used as sources of
biofuel
Biofuel is a fuel that is produced over a short time span from biomass, rather than by the very slow natural processes involved in the formation of fossil fuels, such as oil. According to the United States Energy Information Administration (EIA ...
s, important alternatives to
fossil fuels, such as
biodiesel
Biodiesel is a form of diesel fuel derived from plants or animals and consisting of long-chain fatty acid esters. It is typically made by chemically reacting lipids such as animal fat (tallow), soybean oil, or some other vegetable oil wit ...
. Sweetgrass was used by Native Americans to ward off bugs like
mosquitoes.
These bug repelling properties of sweetgrass were later found by the
American Chemical Society
The American Chemical Society (ACS) is a scientific society based in the United States that supports scientific inquiry in the field of chemistry. Founded in 1876 at New York University, the ACS currently has more than 155,000 members at all ...
in the molecules
phytol and
coumarin
Coumarin () or 2''H''-chromen-2-one is an aromatic organic chemical compound with formula . Its molecule can be described as a benzene molecule with two adjacent hydrogen atoms replaced by a lactone-like chain , forming a second six-membered ...
.
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
flora
Flora is all the plant life present in a particular region or time, generally the naturally occurring ( indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or ''skin flora''.
...
s, their
biodiversity
Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
, genetic diversity and
fitness, the
adaptation
In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
of plants to their environment, and their competitive or
mutualistic interactions with other species. Some ecologists even rely on
empirical data
Empirical evidence for a proposition is evidence, i.e. what supports or counters this proposition, that is constituted by or accessible to sense experience or experimental procedure. Empirical evidence is of central importance to the sciences a ...
from indigenous people that is gathered by ethnobotanists.
This information can relay a great deal of information on how the land once was thousands of years ago and how it has changed over that time.
The goals of plant ecology are to understand the causes of their distribution patterns, productivity, environmental impact, evolution, and responses to environmental change.
Plants depend on certain
edaphic
Edaphology (from Greek , ''edaphos'', "ground",, ''-logia'') is concerned with the influence of soils on living beings, particularly plants.
It is one of two main divisions of soil science, the other being pedology. Edaphology includes the study ...
(soil) and climatic factors in their environment but can modify these factors too. For example, they can change their environment's
albedo
Albedo (; ) is the measure of the diffuse reflection of sunlight, solar radiation out of the total solar radiation and measured on a scale from 0, corresponding to a black body that absorbs all incident radiation, to 1, corresponding to a body ...
, increase
runoff
Runoff, run-off or RUNOFF may refer to:
* RUNOFF, the first computer text-formatting program
* Runoff or run-off, another name for bleed, printing that lies beyond the edges to which a printed sheet is trimmed
* Runoff or run-off, a stock marke ...
interception, stabilise mineral soils and develop their organic content, and affect local temperature. Plants compete with other organisms in their
ecosystem for resources. They interact with their neighbours at a variety of
spatial scale
Spatial scale is a specific application of the term scale for describing or categorizing (e.g. into orders of magnitude) the size of a space (hence ''spatial''), or the extent of it at which a phenomenon or process occurs.
For instance, in phy ...
s in groups, populations and
communities
A community is a social unit (a group of living things) with commonality such as place, norms, religion, values, customs, or identity. Communities may share a sense of place situated in a given geographical area (e.g. a country, village, tow ...
that collectively constitute vegetation. Regions with characteristic
vegetation types
Vegetation classification is the process of classifying and mapping the vegetation over an area of the earth's surface. Vegetation classification is often performed by state based agencies as part of land use, resource and environmental management. ...
and dominant plants as well as similar
abiotic
In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them unde ...
and
biotic factors,
climate
Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
, and
geography make up
biomes
A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
like
tundra or
tropical rainforest
Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equatori ...
.
Herbivore
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthp ...
s eat plants, but plants can
defend themselves and some species are
parasitic
Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
or even
carnivorous
A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other so ...
. Other organisms form
mutually beneficial relationships with plants. For example,
mycorrhizal fungi and
rhizobia
Rhizobia are diazotrophic bacteria that fix nitrogen after becoming established inside the root nodules of legumes (Fabaceae). To express genes for nitrogen fixation, rhizobia require a plant host; they cannot independently fix nitrogen. In ...
provide plants with nutrients in exchange for food,
ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,0 ...
s are recruited by
ant plants to provide protection,
honey bee
A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmop ...
s,
bat
Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s and other animals
pollinate
Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
flowers and
humans
Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
and
other animals act as
dispersal vectors to spread
spore
In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s 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
Phenology is the study of periodic events in biological life cycles and how these are influenced by seasonal and interannual variations in climate, as well as habitat factors (such as elevation).
Examples include the date of emergence of leaves ...
can be a useful
proxy
Proxy may refer to:
* Proxy or agent (law), a substitute authorized to act for another entity or a document which authorizes the agent so to act
* Proxy (climate), a measured variable used to infer the value of a variable of interest in climate re ...
for temperature in
historical climatology
Historical climatology is the study of historical changes in climate and their effect on civilization from the emergence of homininis to the present day. This differs from paleoclimatology which encompasses climate change over the entire history ...
, and the biological
impact of climate change
The effects of climate change impact the physical environment, ecosystems and human societies. The environmental effects of climate change are broad and far-reaching. They affect the water cycle, oceans, sea and land ice (glaciers), sea ...
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. Estimates of atmospheric concentrations since the
Palaeozoic
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ''z ...
have been obtained from
stomatal
In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
densities and the leaf shapes and sizes of ancient
land plants
The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as siste ...
.
Ozone depletion can expose plants to higher levels of
ultraviolet radiation-B (UV-B), resulting in lower growth rates. Moreover, information from studies of
community ecology
In ecology, a community is a group or association (ecology), association of Population ecology, populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, ...
, plant
systematics, and
taxonomy is essential to understanding
vegetation change,
habitat destruction and
species extinction
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
.
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
A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Tra ...
" were discovered by
Barbara McClintock
Barbara McClintock (June 16, 1902 – September 2, 1992) was an American scientist and cytogenetics, cytogeneticist who was awarded the 1983 Nobel Prize in Physiology or Medicine. McClintock received her PhD in botany from Cornell University in ...
while she was studying maize. 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
Peppermint (''Mentha'' × ''piperita'') is a hybrid species of mint, a cross between watermint and spearmint. Indigenous to Europe and the Middle East, the plant is now widely spread and cultivated in many regions of the world.Euro+Med Plantb ...
, ''Mentha'' × ''piperita'', a
sterile hybrid between ''
Mentha aquatica
''Mentha aquatica'' (water mint; syn. ''Mentha hirsuta'' Huds.Euro+Med Plantbase Project''Mentha aquatica'') is a perennial flowering plant in the mint family Lamiaceae. It grows in moist places and is native to much of Europe, northwest Africa ...
'' and spearmint, ''
Mentha spicata
Spearmint, also known as garden mint, common mint, lamb mint and mackerel mint, is a species of mint, ''Mentha spicata'' (, native to Europe and southern temperate Asia, extending from Ireland in the west to southern China in the east. It is nat ...
''. The many cultivated varieties of wheat are the result of multiple inter- and intra-
specific
Specific may refer to:
* Specificity (disambiguation)
* Specific, a cure or therapy for a specific illness
Law
* Specific deterrence, focussed on an individual
* Specific finding, intermediate verdict used by a jury in determining the final ...
crosses between wild species and their hybrids.
Angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
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. This is one of several methods used by plants to promote
outcrossing. 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
sporophyte
A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.
Life cycle
The sporophyte develops from the zygote p ...
s and
dioicous when referring to
bryophyte
The Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited in s ...
gametophytes.
Unlike in higher animals, where
parthenogenesis is rare,
asexual reproduction
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the f ...
may occur in plants by several different mechanisms. The formation of stem
tuber
Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
s in potato is one example. Particularly in
arctic
The Arctic ( or ) is a polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenland), Finland, Iceland, N ...
or
alpine
Alpine may refer to any mountainous region. It may also refer to:
Places Europe
* Alps, a European mountain range
** Alpine states, which overlap with the European range
Australia
* Alpine, New South Wales, a Northern Village
* Alpine National P ...
habitats, where opportunities for fertilisation of flowers
by animals are rare, plantlets or
bulbs
In botany, a bulb is structurally a short stem with fleshy leaves or leaf basesBell, A.D. 1997. ''Plant form: an illustrated guide to flowering plant morphology''. Oxford University Press, Oxford, U.K. that function as food storage organs durin ...
, 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
In botany, apomixis is asexual reproduction without fertilization. Its etymology is Greek for "away from" + "mixing". This definition notably does not mention meiosis. Thus "normal asexual reproduction" of plants, such as propagation from cuttin ...
that occur in plants. Apomixis can also happen in a
seed, producing a seed that contains an embryo genetically identical to the parent.
Most sexually reproducing organisms are diploid, with paired chromosomes, but doubling of their
chromosome number
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectivel ...
may occur due to errors in
cytokinesis
Cytokinesis () is the part of the cell division process during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division in mitosis and meios ...
. This can occur early in development to produce an
autopolyploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
or partly autopolyploid organism, or during normal processes of cellular differentiation to produce some cell types that are polyploid (
endopolyploidy
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
), or during gamete formation. An
allopolyploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
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
The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring ...
from the parent species but live within the same geographical area, may be sufficiently successful to form a new
species
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. Some otherwise sterile plant polyploids can still reproduce
vegetatively
Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is any form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or speci ...
or by seed apomixis, forming clonal populations of identical individuals.
Durum wheat is a fertile
tetraploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
allopolyploid, while
bread wheat
Common wheat (''Triticum aestivum''), also known as bread wheat, is a cultivated wheat species. About 95% of wheat produced worldwide is common wheat; it is the most widely grown of all crops and the cereal with the highest monetary yield.
Ta ...
is a fertile
hexaploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei ( eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set conta ...
. The commercial banana is an example of a sterile, seedless
triploid
Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
hybrid.
Common dandelion
''Taraxacum officinale'', the dandelion or common dandelion, is a flowering herbaceous perennial plant of the dandelion genus in the family Asteraceae (syn. Compositae). The common dandelion is well known for its yellow flower heads that turn in ...
is a triploid that produces viable seeds by apomictic seed.
As in other eukaryotes, the inheritance of
endosymbiotic
An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship.
(The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
organelles like
mitochondria and
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s in plants is non-
Mendelian
Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularize ...
. Chloroplasts are inherited through the male parent in gymnosperms but often through the female parent in flowering plants.
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
''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land.
A winter a ...
'', a weedy species in the mustard family (
Brassicaceae
Brassicaceae () or (the older) Cruciferae () is a medium-sized and economically important family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous plants, while some are shrubs. The leav ...
). The
genome or hereditary information contained in the genes of this species is encoded by about 135 million
base pairs
A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
of DNA, forming one of the smallest genomes among
flowering plants. ''Arabidopsis'' was the first plant to have its genome sequenced, in 2000. The sequencing of some other relatively small genomes, of rice (''
Oryza sativa
''Oryza sativa'', commonly known as Asian rice or indica rice, is the plant species most commonly referred to in English as ''rice''. It is the type of farmed rice whose cultivars are most common globally, and was first domesticated in the Yan ...
'') and ''
Brachypodium distachyon
''Brachypodium distachyon'', commonly called purple false brome or stiff brome, is a grass species native to southern Europe, northern Africa and southwestern Asia east to India. It is related to the major cereal grain species wheat, barley, ...
'', has made them important model species for understanding the genetics, cellular and molecular biology of
cereals
A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
,
grasses and
monocots
Monocotyledons (), commonly referred to as monocots, ( Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
generally.
Model plants such as ''Arabidopsis thaliana'' are used for studying the molecular biology of
plant cells and the
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
. 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
plants. The
single celled
A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms and ...
green alga
The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
''
Chlamydomonas reinhardtii
''Chlamydomonas reinhardtii'' is a single-cell green alga about 10 micrometres in diameter that swims with two flagella. It has a cell wall made of hydroxyproline-rich glycoproteins, a large cup-shaped chloroplast, a large pyrenoid, and an eyes ...
'', while not an
embryophyte itself, contains a
green-pigmented chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
related to that of land plants, making it useful for study. A
red alga
Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
''
Cyanidioschyzon merolae
''Cyanidioschyzon merolae'' is a small (2μm), club-shaped, unicellular haploid red alga adapted to high sulfur acidic hot spring environments (pH 1.5, 45 °C). The cellular architecture of ''C. merolae'' is extremely simple, containing only ...
'' has also been used to study some basic chloroplast functions.
Spinach
Spinach (''Spinacia oleracea'') is a leafy green flowering plant native to central and western Asia. It is of the order Caryophyllales, family Amaranthaceae, subfamily Chenopodioideae. Its leaves are a common edible vegetable consumed either fr ...
,
peas,
soybeans and a moss ''
Physcomitrella patens
''Physcomitrium patens'', (synonym: ''Physcomitrella patens'' ) the spreading earthmoss, is a moss (bryophyte) used as a model organism for studies on plant evolution, development, and physiology.
Distribution and ecology
''Physcomitrella p ...
'' are commonly used to study plant cell biology.
''
Agrobacterium tumefaciens
''Agrobacterium radiobacter'' (more commonly known as ''Agrobacterium tumefaciens'') is the causal agent of crown gall disease (the formation of tumours) in over 140 species of eudicots. It is a rod-shaped, Gram-negative soil bacterium. Symptom ...
'', a soil
rhizosphere bacterium, can attach to plant cells and infect them with a
callus
A callus is an area of thickened and sometimes hardened skin that forms as a response to repeated friction, pressure, or other irritation. Since repeated contact is required, calluses are most often found on the feet and hands, but they may o ...
-inducing
Ti plasmid
A tumour inducing (Ti) plasmid is a plasmid found in pathogenic species of ''Agrobacterium'', including ''A. tumefaciens, ''A. rhizogenes'', ''A. rubi'' and ''A. vitis''.
Evolutionarily, the Ti plasmid is part of a family of plasmids carried b ...
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
The ''nif'' genes are genes encoding enzymes involved in the fixation of atmospheric nitrogen into a form of nitrogen available to living organisms. The primary enzyme encoded by the ''nif'' genes is the nitrogenase complex which is in charge of c ...
responsible for
nitrogen fixation
Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. A ...
in the root nodules of
legumes
A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock for ...
and other plant species. Today, genetic modification of the Ti plasmid is one of the main techniques for introduction of
transgene
A transgene is a gene that has been transferred naturally, or by any of a number of genetic engineering techniques, from one organism to another. The introduction of a transgene, in a process known as transgenesis, has the potential to change t ...
s to plants and the creation of
genetically modified crops.
Epigenetics
Epigenetics
In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are "o ...
is the study of heritable changes in
gene function that cannot be explained by changes in the underlying
DNA sequence but cause the organism's genes to behave (or "express themselves") differently. One example of epigenetic change is the marking of the genes by
DNA methylation
DNA methylation is a biological process by which methyl groups are added to the DNA molecule. Methylation can change the activity of a DNA segment without changing the sequence. When located in a gene promoter, DNA methylation typically acts t ...
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 division
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there a ...
s for the remainder of the cell's life. Some epigenetic changes have been shown to be
heritable, while others are reset in the germ cells.
Epigenetic changes in
eukaryotic biology serve to regulate the process of
cellular differentiation
Cellular differentiation is the process in which a stem cell alters from one type to a differentiated one. Usually, the cell changes to a more specialized type. Differentiation happens multiple times during the development of a multicellular ...
. During
morphogenesis,
totipotent Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta.
According to Latin terms, Pluripotentia means the ability for many thin ...
stem cells become the various
pluripotent Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta.
According to Latin terms, Pluripotentia means the ability for many thin ...
cell line
An immortalised cell line is a population of cells from a multicellular organism which would normally not proliferate indefinitely but, due to mutation, have evaded normal cellular senescence and instead can keep undergoing division. The cell ...
s 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
Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms.
Etymology
The term ''parenchyma'' is New Latin from the word � ...
,
xylem vessel elements,
phloem sieve tubes,
guard cell
Guard cells are specialized plant cells in the epidermis of leaves, stems and other organs that are used to control gas exchange. They are produced in pairs with a gap between them that forms a stomatal pore. The stomatal pores are largest when ...
s of the
epidermis
The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
, etc. as it continues to
divide. The process results from the epigenetic activation of some genes and inhibition of others.
Unlike animals, many plant cells, particularly those of the
parenchyma
Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms.
Etymology
The term ''parenchyma'' is New Latin from the word � ...
, 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
Chromatin remodeling is the dynamic modification of chromatin architecture to allow access of condensed genomic DNA to the regulatory Transcription (genetics), transcription machinery proteins, and thereby control gene expression. Such remodeling i ...
, 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.
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.
Plant evolution
The
chloroplast
A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s 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
An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship.
(The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within" ...
relationship between an ancestral
eukaryotic cell and a
cyanobacterial resident.
The
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
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
Charophyta () is a group of freshwater green algae, called charophytes (), sometimes treated as a division, yet also as a superdivision or an unranked clade. The terrestrial plants, the Embryophyta emerged within Charophyta, possibly from terre ...
, sister to the green algal division
Chlorophyta
Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to a ...
, is considered to contain the ancestor of true plants. The Charophyte class
Charophyceae and the land plant sub-kingdom
Embryophyta
The Embryophyta (), or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes () have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as siste ...
together form the
monophyletic group or clade
Streptophytina
The Phragmoplastophyta (Lecointre & Guyader 2006) are a proposed sister clade of the Klebsormidiaceae in the Streptophyte/Charophyte clade. The Phragmoplastophyta consist of the Charophycaea and another unnamed clade which contains the Coleocha ...
.
Nonvascular land plants are
embryophytes that lack the vascular tissues
xylem
Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
and
phloem. They include
mosses,
liverworts and
hornwort
Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s.
Pteridophytic vascular plants with true xylem and phloem that reproduced by spores germinating into free-living 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
Lycopodiopsida is a class of vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching s ...
,
sphenophylls and
progymnosperm
The progymnosperms are an extinct group of woody, spore-bearing plants that is presumed to have evolved from the trimerophytes, and eventually gave rise to the gymnosperms, ancestral to acrogymnosperms and angiosperms (flowering plants). They h ...
s, had independently evolved "megaspory" – their spores were of two distinct sizes, larger
megaspore
Megaspores, also called macrospores, are a type of spore that is present in heterosporous plants. These plants have two spore types, megaspores and microspores. Generally speaking, the megaspore, or large spore, germinates into a female gametophy ...
s 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 (
integument
In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, shell, germ or rind.
Etymology
The term is derived from ''integumentum'', which is Latin for "a covering". In a transferred, or ...
s). The young sporophyte develops within the seed, which on
germination
Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
splits to release it. The earliest known seed plants date from the latest Devonian
Famennian
The Famennian is the latter of two faunal stages in the Late Devonian Epoch. The most recent estimate for its duration estimates that it lasted from around 371.1 million years ago to 359.3 million years ago. An earlier 2012 estimate, still used ...
stage. Following the evolution of the seed habit,
seed plants
A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
diversified, giving rise to a number of now-extinct groups, including
seed ferns, as well as the modern gymnosperms and angiosperms.
Gymnosperms produce "naked seeds" not fully enclosed in an ovary; modern representatives include
conifers
Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All exta ...
,
cycad
Cycads are seed plants that typically have a stout and woody ( ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or ...
s, ''
Ginkgo'', and
Gnetales.
Angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
produce seeds enclosed in a structure such as a
carpel
Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils' ...
or an
ovary
The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
. Ongoing research on the molecular phylogenetics of living plants appears to show that the angiosperms are a
sister clade to the gymnosperms.
Plant physiology
Plant
physiology encompasses all the internal chemical and physical activities of plants associated with life. Chemicals obtained from the air, soil and water form the basis of all
plant metabolism
Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants. Closely related fields include plant morphology (structure of plants), plant ecology (interactions with the environment), phytochemistry ( ...
. The energy of sunlight, captured by oxygenic photosynthesis and released by
cellular respiration
Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
, 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.
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.
Molecules are moved within plants by transport processes that operate at a variety of
spatial scale
Spatial scale is a specific application of the term scale for describing or categorizing (e.g. into orders of magnitude) the size of a space (hence ''spatial''), or the extent of it at which a phenomenon or process occurs.
For instance, in phy ...
s. 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
In plants, the transpiration stream is the uninterrupted stream of water and solutes which is taken up by the roots and transported via the xylem to the leaves where it evaporates into the air/apoplast-interface of the substomatal cavity. It is dr ...
.
Diffusion,
osmosis, and
active transport
In cellular biology, ''active transport'' is the movement of molecules or ions across a cell membrane from a region of lower concentration to a region of higher concentration—against the concentration gradient. Active transport requires cellul ...
and
mass flow are all different ways transport can occur. Examples of
elements that plants need to transport are
nitrogen,
phosphorus,
potassium,
calcium
Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar ...
,
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
Plant nutrition is the study of the chemical elements and compounds necessary for plant growth and reproduction, plant metabolism and their external supply. In its absence the plant is unable to complete a normal life cycle, or that the element i ...
come from the chemical breakdown of soil minerals.
Sucrose produced by photosynthesis is transported from the leaves to other parts of the plant in the phloem and
plant hormones
Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, p ...
are transported by a variety of processes.
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
''Mimosa pudica'' (from la, pudica "shy, bashful or shrinking"; also called sensitive plant, sleepy plant, action plant, touch-me-not, shameplant) is a creeping annual or perennial flowering plant of the pea/legume family Fabaceae. It is often ...
'', the insect traps of
Venus flytrap and
bladderwort
''Utricularia'', commonly and collectively called the bladderworts, is a genus of carnivorous plants consisting of approximately 233 species (precise counts differ based on classification opinions; a 2001 publication lists 215 species).Salmon, Br ...
s, and the pollinia of orchids.
The hypothesis that plant growth and development is coordinated by
plant hormone
Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, p ...
s or plant growth regulators first emerged in the late 19th century. Darwin experimented on the movements of plant shoots and roots towards
light and
gravity, and concluded "It is hardly an exaggeration to say that the tip of the radicle . . acts like the brain of one of the lower animals . . directing the several movements". About the same time, the role of
auxin
Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essenti ...
s (from the Greek , to grow) in control of plant growth was first outlined by the Dutch scientist
Frits Went
Friedrich August Ferdinand Christian Went ForMemRS (June 18, 1863 – July 24, 1935) was a Dutch botanist.
Went was born in Amsterdam. He was professor of botany and director of the Botanical Garden at the University of Utrecht. His eldes ...
. The first known auxin,
indole-3-acetic acid (IAA), which promotes cell growth, was only isolated from plants about 50 years later. This compound mediates the tropic responses of shoots and roots towards light and gravity. The finding in 1939 that plant
callus
A callus is an area of thickened and sometimes hardened skin that forms as a response to repeated friction, pressure, or other irritation. Since repeated contact is required, calluses are most often found on the feet and hands, but they may o ...
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.
Cytokinin
Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and lea ...
s are a class of plant hormones named for their control of cell division (especially
cytokinesis
Cytokinesis () is the part of the cell division process during which the cytoplasm of a single eukaryotic cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of nuclear division in mitosis and meios ...
). The natural cytokinin
zeatin
Zeatin is a cytokinin derived from adenine, which occurs in the form of a ''cis''- and a ''trans''- isomer and conjugates. Zeatin was discovered in immature corn kernels from the genus '' Zea''. It promotes growth of lateral buds and when sprayed ...
was discovered in corn, ''
Zea mays'', and is a derivative of the
purine adenine
Adenine () (symbol A or Ade) is a nucleobase (a purine derivative). It is one of the four nucleobases in the nucleic acid of DNA that are represented by the letters G–C–A–T. The three others are guanine, cytosine and thymine. Its derivativ ...
. Zeatin is produced in roots and transported to shoots in the xylem where it promotes cell division, bud development, and the greening of chloroplasts. The
gibberelins
Gibberellins (GAs) are plant hormones that regulate various developmental processes, including stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. GAs are one of the longest-known classes of plan ...
, such as
gibberelic acid
Gibberellic acid (also called gibberellin A3, GA, and GA3) is a hormone found in plants and fungi. Its chemical formula is C19H22O6. When purified, it is a white to pale-yellow solid.
Plants in their normal state produce large amounts of GA3. ...
are
diterpenes synthesised from
acetyl CoA
Acetyl-CoA (acetyl coenzyme A) is a molecule that participates in many biochemical reactions in protein, carbohydrate and lipid metabolism. Its main function is to deliver the acetyl group to the citric acid cycle (Krebs cycle) to be oxidized fo ...
via the
mevalonate pathway
The mevalonate pathway, also known as the isoprenoid pathway or HMG-CoA reductase pathway is an essential metabolic pathway present in eukaryotes, archaea, and some bacteria. The pathway produces two five-carbon building blocks called isopentenyl ...
. 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.
Abscisic acid
Abscisic acid (ABA) is a plant hormone. ABA functions in many plant developmental processes, including seed and bud dormancy, the control of organ size and stomatal closure. It is especially important for plants in the response to environmental s ...
(ABA) occurs in all land plants except liverworts, and is synthesised from
carotenoid
Carotenoids (), also called tetraterpenoids, are yellow, orange, and red organic pigments that are produced by plants and algae, as well as several bacteria, and fungi. Carotenoids give the characteristic color to pumpkins, carrots, parsnips, ...
s 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
Abscission () is the shedding of various parts of an organism, such as a plant dropping a leaf, fruit, flower, or seed. In zoology, abscission is the intentional shedding of a body part, such as the shedding of a claw, husk, or the autotomy of a ...
.
Ethylene
Ethylene ( IUPAC name: ethene) is a hydrocarbon which has the formula or . It is a colourless, flammable gas with a faint "sweet and musky" odour when pure. It is the simplest alkene (a hydrocarbon with carbon-carbon double bonds).
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, and it, or the synthetic growth regulator
ethephon
Ethephon is a plant growth regulator.
Mechanism of action
Upon metabolism by the plant, it is converted into ethylene, a potent regulator of plant growth and ripeness. It is also a butyrylcholinesterase inhibitor.
Uses in various crops
Etheph ...
which is rapidly metabolised to produce ethylene, are used on industrial scale to promote ripening of cotton,
pineapples and other
climacteric crops.
Another class of
phytohormone
Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
s is the
jasmonates, first isolated from the oil of ''
Jasminum grandiflorum
''Jasminum grandiflorum'', also known variously as the Spanish jasmine, Royal jasmine, Catalan jasmine, among others, is a species of jasmine native to South Asia, the Arabian peninsula, East and Northeast Africa and the Yunnan and Sichuan region ...
'' which regulates wound responses in plants by unblocking the expression of genes required in the
systemic acquired resistance Systemic acquired resistance (SAR) is a "whole-plant" resistance response that occurs following an earlier localized exposure to a pathogen. SAR is analogous to the innate immune system found in animals, and although there are many shared aspects be ...
response to pathogen attack.
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 In developmental biology, photomorphogenesis is light-mediated development, where plant growth patterns respond to the light spectrum. This is a completely separate process from photosynthesis where light is used as a source of energy. Phytochromes, ...
.
Phytochrome
Phytochromes are a class of photoreceptor in plants, bacteria and fungi used to detect light. They are sensitive to light in the red and far-red region of the visible spectrum and can be classed as either Type I, which are activated by far-re ...
s are the
photoreceptors in a plant that are sensitive to light.
Plant anatomy and morphology
Plant anatomy
Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plant ...
is the study of the structure of plant cells and tissues, whereas
plant morphology
Phytomorphology is the study of the morphology (biology), physical form and external structure of plants.Raven, P. H., R. F. Evert, & S. E. Eichhorn. ''Biology of Plants'', 7th ed., page 9. (New York: W. H. Freeman, 2005). . This is usually cons ...
is the study of their external form.
All plants are multicellular eukaryotes, their DNA stored in nuclei. The characteristic features of
plant cells that distinguish them from those of animals and fungi include a primary
cell wall
A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
composed of the polysaccharides
cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
,
hemicellulose and
pectin, larger
vacuole
A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s 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
Amyloplasts are a type of plastid, double-enveloped organelles in plant cells that are involved in various biological pathways. Amyloplasts are specifically a type of leucoplast, a subcategory for colorless, non-pigment-containing plastids. Amylopl ...
) or lipids (
elaioplasts). Uniquely,
streptophyte
Streptophyta (), informally the streptophytes (, from the Greek ''strepto'' 'twisted', for the morphology of the sperm of some members), is a clade of plants. The composition of the clade varies considerably between authors, but the definition e ...
cells and those of the green algal order
Trentepohliales
Trentepohliales is an order of Ulvophyceaen green algae.See the NCBI
The National Center for Biotechnology Information (NCBI) is part of the United States National Library of Medicine (NLM), a branch of the National Institutes of Health (NI ...
divide by construction of a
phragmoplast
image:Phragmoplast.png, 300px, Phragmoplast and cell plate formation in a plant cell during cytokinesis. Left side: Phragmoplast forms and cell plate starts to assemble in the center of the cell. Towards the right: Phragmoplast enlarges in a donut- ...
as a template for building a
cell plate
image:Phragmoplast.png, 300px, Phragmoplast and cell plate formation in a plant cell during cytokinesis. Left side: Phragmoplast forms and cell plate starts to assemble in the center of the cell. Towards the right: Phragmoplast enlarges in a donut ...
late in
cell division
Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there a ...
.
The bodies of
vascular plants including
clubmosses
Lycopodiopsida is a class of vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching ...
,
ferns and
seed plants
A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
(
gymnosperms and
angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
) generally have aerial and subterranean subsystems. The
shoot
In botany, a plant shoot consists of any plant stem together with its appendages, leaves and lateral buds, flowering stems, and flower buds. The new growth from seed germination that grows upward is a shoot where leaves will develop. In the spr ...
s consist of
stems bearing green photosynthesising
leaves
A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
and reproductive structures. The underground vascularised
roots bear
root hairs at their tips and generally lack chlorophyll. Non-vascular plants, the
liverworts,
hornworts and
mosses do not produce ground-penetrating vascular roots and most of the plant participates in photosynthesis. The
sporophyte
A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.
Life cycle
The sporophyte develops from the zygote p ...
generation is nonphotosynthetic in liverworts but may be able to contribute part of its energy needs by photosynthesis in mosses and hornworts.
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. Cells in each system are capable of creating cells of the other and producing
adventitious Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant a ...
shoots or roots.
Stolons and
tuber
Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
s are examples of shoots that can grow roots. Roots that spread out close to the surface, such as those of willows, can produce shoots and ultimately new plants. 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'', or even a single
cell
Cell most often refers to:
* Cell (biology), the functional basic unit of life
Cell may also refer to:
Locations
* Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...
– which can dedifferentiate into a
callus
A callus is an area of thickened and sometimes hardened skin that forms as a response to repeated friction, pressure, or other irritation. Since repeated contact is required, calluses are most often found on the feet and hands, but they may o ...
(a mass of unspecialised cells) that can grow into a new plant.
In vascular plants, the xylem and phloem are the conductive tissues that transport resources between shoots and roots. Roots are often adapted to store food such as sugars or
starch, as in
sugar beet
A sugar beet is a plant whose root contains a high concentration of sucrose and which is grown commercially for sugar production. In plant breeding, it is known as the Altissima cultivar group of the common beet (''Beta vulgaris''). Together w ...
s and carrots.
Stems mainly provide support to the leaves and reproductive structures, but can store water in succulent plants such as
cacti
A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Gree ...
, food as in potato
tubers, or
reproduce vegetatively as in the
stolons of
strawberry
The garden strawberry (or simply strawberry; ''Fragaria × ananassa'') is a widely grown hybrid species of the genus ''Fragaria'', collectively known as the strawberries, which are cultivated worldwide for their fruit. The fruit is widely ap ...
plants or in the process of
layering
Layering has evolved as a common means of vegetative propagation of numerous species in natural environments. Layering is also utilized by horticulturists to propagate desirable plants.
Natural layering typically occurs when a branch touches ...
. Leaves gather sunlight and carry out
photosynthesis. Large, flat, flexible, green leaves are called foliage leaves.
Gymnosperms, such as
conifers,
cycad
Cycads are seed plants that typically have a stout and woody ( ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male or ...
s, ''
Ginkgo'', and
gnetophytes
Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three reli ...
are seed-producing plants with open seeds.
Angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
are
seed-producing plants that produce flowers and have enclosed seeds. Woody plants, such as
azalea
Azaleas are flowering shrubs in the genus ''Rhododendron'', particularly the former sections '' Tsutsusi'' (evergreen) and '' Pentanthera'' (deciduous). Azaleas bloom in the spring (April and May in the temperate Northern Hemisphere, and Octo ...
s and
oaks, undergo a secondary growth phase resulting in two additional types of tissues: wood (secondary
xylem
Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
) and bark (secondary
phloem and
cork
Cork or CORK may refer to:
Materials
* Cork (material), an impermeable buoyant plant product
** Cork (plug), a cylindrical or conical object used to seal a container
***Wine cork
Places Ireland
* Cork (city)
** Metropolitan Cork, also known as G ...
). All gymnosperms and many angiosperms are woody plants. Some plants reproduce sexually, some asexually, and some via both means.
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. Furthermore, structures can be seen as processes, that is, process combinations.
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. 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
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. 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
Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations that ...
principle of
common descent
Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
– 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 evolutionary lines and is likely to continue to do so. The dominant classification system is called
Linnaean taxonomy. It includes ranks and
binomial nomenclature
In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
. The nomenclature of botanical organisms is codified in the
(ICN) and administered by the
International Botanical Congress
International Botanical Congress (IBC) is an international meeting of botanists in all scientific fields, authorized by the International Association of Botanical and Mycological Societies (IABMS) and held every six years, with the location rotati ...
.
Kingdom
Kingdom commonly refers to:
* A monarchy ruled by a king or queen
* Kingdom (biology), a category in biological taxonomy
Kingdom may also refer to:
Arts and media Television
* ''Kingdom'' (British TV series), a 2007 British television drama s ...
Plantae belongs to
Domain Eukaryota and is broken down recursively until each species is separately classified. The order is:
Kingdom
Kingdom commonly refers to:
* A monarchy ruled by a king or queen
* Kingdom (biology), a category in biological taxonomy
Kingdom may also refer to:
Arts and media Television
* ''Kingdom'' (British TV series), a 2007 British television drama s ...
;
Phylum (or Division);
Class
Class or The Class may refer to:
Common uses not otherwise categorized
* Class (biology), a taxonomic rank
* Class (knowledge representation), a collection of individuals or objects
* Class (philosophy), an analytical concept used differently ...
;
Order
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
;
Family;
Genus (plural ''genera'');
Species
In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
. The scientific name of a plant represents its genus and its species within the genus, resulting in a single worldwide name for each organism. For example, the tiger lily is ''
Lilium columbianum
''Lilium columbianum'' is a lily native to western North America. It is also known as the Columbia lily, Columbia tiger lily, or simply tiger lily (sharing the latter common name with several other lily species in its genus).
Distribution and ha ...
''. ''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).
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. As an example, species of ''
Pereskia
''Pereskia'' is a small genus of about four species of cacti that do not look much like other types of cacti, having substantial leaves and non-succulent stems. The genus is named after Nicolas-Claude Fabri de Peiresc, a 16th-century French bota ...
'' are trees or bushes with prominent leaves. They do not obviously resemble a typical leafless
cactus
A cactus (, or less commonly, cactus) is a member of the plant family Cactaceae, a family comprising about 127 genera with some 1750 known species of the order Caryophyllales. The word ''cactus'' derives, through Latin, from the Ancient Gree ...
such as an ''
Echinocactus
''Echinocactus'' is a genus of cacti in the subfamily Cactoideae. The generic name derives from the Ancient Greek εχινος (''echinos''), meaning "spiny," and ''cactus''. It and '' Ferocactus'' are the two genera of barrel cactus. Members ...
''. However, both ''Pereskia'' and ''Echinocactus'' have spines produced from
areoles
In botany, areoles are small light- to dark-colored bumps on cacti out of which grow clusters of spines. Areoles are important diagnostic features of cacti, and identify them as a family distinct from other succulent plants. Gordon Rowley - Wh ...
(highly specialised pad-like structures) suggesting that the two genera are indeed related.
Judging relationships based on shared characters requires care, since plants may resemble one another through
convergent evolution in which characters have arisen independently. Some
euphorbia
''Euphorbia'' is a very large and diverse genus of flowering plants, commonly called spurge, in the family Euphorbiaceae. "Euphorbia" is sometimes used in ordinary English to collectively refer to all members of Euphorbiaceae (in deference to t ...
s 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
Homoplasy, in biology and phylogenetics, is the term used to describe a feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from homology, which is the term used to characterize ...
) 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
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
s: tree-like diagrams showing the pattern of evolutionary branching and descent.
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." 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.
In 1998, the
Angiosperm Phylogeny Group
The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disco ...
published a
phylogeny for flowering plants based on an analysis of DNA sequences from most families of flowering plants. As a result of this work, many questions, such as which families represent the earliest branches of
angiosperms
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
, have now been answered. Investigating how plant species are related to each other allows botanists to better understand the process of evolution in plants. 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
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
. Technological developments such as computers and
electron microscope
An electron microscope is a microscope that uses a beam of accelerated electrons as a source of illumination. As the wavelength of an electron can be up to 100,000 times shorter than that of visible light photons, electron microscopes have a h ...
s have greatly increased the level of detail studied and speed at which data can be analysed.
Symbols
A few symbols are in current use in botany. A number of others are obsolete; for example, Linnaeus used planetary symbols for woody, herbaceous and perennial plants, and Willd used (Saturn) for neuter in addition to (Mercury) for hermaphroditic. The following symbols are still used:
[Niki Simpson]
Botanical symbols: a new symbol set for new images
''Botanical Journal of the Linnean Society'', Volume 162, Issue 2, February 2010, Pages 117–129
:♀ female
:♂ male
:⚥
hermaphrodite/bisexual
:⚲ vegetative (asexual) reproduction
:◊ sex unknown
:☉ annual
:
16px (⚇)
biennial
Biennial means (an event) lasting for two years or occurring every two years. The related term biennium is used in reference to a period of two years.
In particular, it can refer to:
* Biennial plant, a plant which blooms in its second year and th ...
:
16px (♾)
perennial
A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
:☠ poisonous
:🛈 further information
:× crossbred hybrid
:+ grafted hybrid
See also
*
Branches of botany
Botany is a natural science concerned with the study of plants. The main branches of botany (also referred to as "plant science") are commonly divided into three groups: core topics, concerned with the study of the fundamental natural phenomena and ...
*
Evolution of plants
*
Glossary of botanical terms
*
Glossary of plant morphology
This page provides a glossary of plant morphology. Botanists and other biologists who study plant morphology use a number of different terms to classify and identify plant organs and parts that can be observed using no more than a handheld magnif ...
*
List of botany journals
The following is a list of notable botanical scientific journals.
General botany
The following table is a list of scientific journals publishing articles on many areas of botany.
Agronomy and horticulture
The following table is a list of ...
*
List of botanists
This is a list of botanists who have Wikipedia articles, in alphabetical order by surname. The List of botanists by author abbreviation is mostly a list of plant taxonomists because an author receives a standard abbreviation only when that aut ...
*
List of botanical gardens
A botanical garden is a place where plants, especially ferns, conifers and flowering plants, are grown and displayed for the purposes of research, conservation, and education. This distinguishes them from parks and pleasure gardens where plants, us ...
*
List of botanists by author abbreviation
*
List of domesticated plants
This is a list of plants that have been domesticated by humans. The list includes individual plant species identified by their common names as well as larger formal and informal botanical categories which include at least some domesticated individ ...
*
List of flowers
A ''list'' is any set of items in a row. List or lists may also refer to:
People
* List (surname)
Organizations
* List College, an undergraduate division of the Jewish Theological Seminary of America
* SC Germania List, German rugby union ...
*
List of systems of plant taxonomy
*
Outline of botany
The following outline is provided as an overview of and topical guide to botany:
Botany – biological discipline which involves the study of plants.
Core concepts of botany
* Bud
* Cell wall
* Chlorophyll
* Chloroplast
* Flora
* Flower
...
*
Timeline of British botany
__NOTOC__
This article lists (in chronological order) notable events in the history of botany in Britain.
pre-1801
* 1538 First British flora "Libellus de Herbaria" by William Turner's published.
"A new Herball, wherin are conteined the names of ...
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{{Authority control
Articles containing video clips