Plants are predominantly
photosynthetic eukaryotes of the
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. Historically, the plant kingdom encompassed all living things that were not
animal
Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s, and included
algae and
fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
; however, all current definitions of Plantae exclude the fungi and some algae, as well as the
prokaryote
A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Conne ...
s (the
archaea and
bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
). By one definition, plants form the
clade Viridiplantae
Viridiplantae (literally "green plants") are a clade of eukaryotic organisms that comprise approximately 450,000–500,000 species and play important roles in both terrestrial and aquatic ecosystems. They are made up of the green algae, which ar ...
(Latin name for "green plants") which is sister of the
Glaucophyta
The glaucophytes, also known as glaucocystophytes or glaucocystids, are a small group of unicellular algae found in freshwater and moist terrestrial environments, less common today than they were during the Proterozoic. The stated number of speci ...
, and consists of the
green algae and
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 ...
(land plants). The latter includes the
flowering plant
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 ...
s,
conifers and other
gymnosperm
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
s,
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s and
their allies,
hornworts,
liverworts, and
mosses.
Most plants are
multicellular
A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism.
All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially un ...
organisms. Green plants obtain most of their energy from
sunlight via
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 i ...
by primary
chloroplasts that are derived from
endosymbiosis
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 ...
with
cyanobacteria. Their chloroplasts contain
chlorophylls a and b, which gives them their green color. Some plants 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 ha ...
or
mycotrophic and have lost the ability to produce normal amounts of chlorophyll or to photosynthesize, but still have flowers, fruits, and seeds. Plants are characterized by
sexual reproduction
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
and
alternation of generations, although
asexual reproduction is also common.
There are about 320,000 known
species
In biology, a species is the basic unit of 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 appropriate s ...
of plants, of which the great majority, some 260,000–290,000,
produce seeds.
Green plants provide a substantial proportion of the world's molecular oxygen,
and are the basis of most of Earth's ecosystems. Plants that produce
grain,
fruit
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
, and
vegetable
Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems, ...
s also form basic human foods and have been
domesticated
Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
for millennia. Plants have many
cultural and other uses, as ornaments,
building materials
Building material is material used for construction. Many naturally occurring substances, such as clay, rocks, sand, wood, and even twigs and leaves, have been used to construct buildings. Apart from naturally occurring materials, many man- ...
,
writing material
Writing material refers to the materials that provide the surfaces on which humans use writing instruments to inscribe writings. The same materials can also be used for symbolic or representational drawings. Building material on which writings or ...
and, in great variety, they have been the
source of medicines and
psychoactive drug
A psychoactive drug, psychopharmaceutical, psychoactive agent or psychotropic drug is a chemical substance, that changes functions of the nervous system, and results in alterations in perception, mood, consciousness, cognition or behavior. ...
s. The scientific study of plants is known as
botany
Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
, 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 i ...
.
Definition
All living things were traditionally placed into one of two groups, plants and animals. This classification may date from
Aristotle
Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ph ...
(384 BC – 322 BC), who made the distinction between plants, which generally do not move, and animals, which often are mobile to catch their food. Much later, when
Linnaeus
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 ...
(1707–1778) created the basis of the modern system of
scientific classification
Taxonomy is the practice and science of categorization or classification.
A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
, these two groups became the
kingdoms
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 ...
Vegetabilia (later Metaphyta or Plantae) and
Animalia (also called Metazoa). Since then, it has become clear that the plant kingdom as originally defined included several unrelated groups, and the
fungi
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
and several groups of
algae were removed to new kingdoms. However, these organisms are still sometimes considered plants, particularly in informal contexts.
The term "plant" generally implies the possession of the following traits: multicellularity, possession of cell walls containing
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 w ...
, and the ability to carry out photosynthesis with primary chloroplasts.
[– Definition from the Merriam-Webster Online Dictionary">]
Current definitions of Plantae
When the name Plantae or plant is applied to a specific group of organisms or
taxon
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 n ...
, it usually refers to one of four concepts. From least to most inclusive, these four groupings are:
Another way of looking at the relationships between the different groups that have been called "plants" is through a
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 ...
, which shows their evolutionary relationships. These are not yet completely settled, but .
Those which have been called "plants" are in bold (some minor groups have been omitted).
The way in which the groups of green algae are combined and named varies considerably between authors.
Algae
Algae consist of several groups of organisms which produce food by photosynthesis and thus have traditionally been included in the plant kingdom. The
seaweeds range from large multicellular algae to single-celled organisms and are classified into three groups, the
green algae,
red algae and
brown algae
Brown algae (singular: alga), comprising the class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and p ...
. There is good evidence that the brown algae evolved independently from the others, from non-photosynthetic ancestors that formed endosymbiotic relationships with red algae rather than from cyanobacteria, and they are no longer classified as plants as defined here.
The Viridiplantae, the green plants – green algae and land plants – form a
clade, a group consisting of all the descendants of a common ancestor. With a few exceptions, the green plants have the following features in common; primary
chloroplasts derived from cyanobacteria containing
chlorophylls ''a'' and ''b'', cell walls containing
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 w ...
, and food stores in the form of
starch contained within the plastids. They undergo closed
mitosis without
centrioles, and typically have
mitochondria with flat cristae. The
chloroplasts of green plants are surrounded by two membranes, suggesting they originated directly from endosymbiotic
cyanobacteria.
Two additional groups, the
Rhodophyta (red algae) and
Glaucophyta
The glaucophytes, also known as glaucocystophytes or glaucocystids, are a small group of unicellular algae found in freshwater and moist terrestrial environments, less common today than they were during the Proterozoic. The stated number of speci ...
(glaucophyte algae), also have primary chloroplasts that appear to be derived directly from endosymbiotic
cyanobacteria, although they differ from Viridiplantae in the pigments which are used in photosynthesis and so are different in colour. These groups also differ from green plants in that the storage polysaccharide is
floridean starch
Floridean starch is a type of a storage glucan found in glaucophytes and in red algae (or rhodophytes), in which it is usually the primary sink for fixed carbon from photosynthesis. It is found in grains or granules in the cell's cytoplasm and is ...
and is stored in the cytoplasm rather than in the plastids. They appear to have had a common origin with Viridiplantae and the three groups form the clade
Archaeplastida
The Archaeplastida (or kingdom Plantae '' sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor gr ...
, whose name implies that their chloroplasts were derived from a single ancient endosymbiotic event. This is the broadest modern definition of the term 'plant'.
In contrast, most other algae (e.g.
brown algae/diatoms,
haptophyte
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for '' Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
s,
dinoflagellates, and
euglenid
Euglenids (euglenoids, or euglenophytes, formally Euglenida/Euglenoida, ICZN, or Euglenophyceae, ICBN) are one of the best-known groups of flagellates, which are excavate eukaryotes of the phylum Euglenophyta and their cell structure is typical o ...
s) not only have different pigments but also have chloroplasts with three or four surrounding membranes. They are not close relatives of the Archaeplastida, presumably having acquired chloroplasts separately from ingested or symbiotic green and red algae. They are thus not included in even the broadest modern definition of the plant kingdom, although they were in the past.
The green plants or Viridiplantae were traditionally divided into the green algae (including the stoneworts) and the land plants. However, it is now known that the land plants evolved from within a group of green algae, so that the green algae by themselves are a
paraphyletic group, i.e. a group that excludes some of the descendants of a common ancestor. Paraphyletic groups are generally avoided in modern classifications, so that in recent treatments the Viridiplantae have been divided into two clades, the
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 ...
and the
Streptophyta
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 ...
(including the land plants and Charophyta).
The Chlorophyta (a name that has also been used for ''all'' green algae) are the sister group to the Charophytes, from which the land plants evolved. There are about 4,300 species,
mainly unicellular or multicellular marine organisms such as the sea lettuce, ''
Ulva
Ulva (; gd, Ulbha) is a small island in the Inner Hebrides of Scotland, off the west coast of Mull. It is separated from Mull by a narrow strait, and connected to the neighbouring island of Gometra by a bridge. Much of the island is formed f ...
''.
The other group within the Viridiplantae are the mainly freshwater or terrestrial Streptophyta, which consists of the land plants together with the Charophyta, itself consisting of several groups of green algae such as the
desmid
Desmidiales, commonly called desmids (''Gr.'' ''desmos'', bond or chain), are an order in the Charophyta, a division of green algae in which the land plants (Embryophyta) emerged. Or in other words, Desmid, (order Desmidiales), order of single- ...
s and
stoneworts
Charales is an order of freshwater green algae in the division Charophyta, class Charophyceae, commonly known as stoneworts. Depending on the treatment of the genus '' Nitellopsis'', living (extant) species are placed into either one family (Ch ...
. Streptophyte algae are either unicellular or form multicellular filaments, branched or unbranched.
The genus ''
Spirogyra
''Spirogyra'' (common names include water silk, mermaid's tresses, and blanket weed) is a genus of filamentous charophyte green algae of the order Zygnematales, named for the helical or spiral arrangement of the chloroplasts that is character ...
'' is a filamentous streptophyte alga familiar to many, as it is often used in teaching and is one of the organisms responsible for the algal "scum" on ponds. The freshwater stoneworts strongly resemble land plants and are believed to be their closest relatives. Growing immersed in fresh water, they consist of a central stalk with whorls of branchlets.
Fungi
Linnaeus' original classification placed the fungi within the Plantae, since they were unquestionably neither animals or minerals and these were the only other alternatives. With 19th century developments in
microbiology,
Ernst Haeckel introduced the new kingdom Protista in addition to Plantae and Animalia, but whether fungi were best placed in the Plantae or should be reclassified as
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 exc ...
s remained controversial. In 1969,
Robert Whittaker proposed the creation of the kingdom Fungi. Molecular evidence has since shown that the
most recent common ancestor
In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
(concestor), of the Fungi was probably more similar to that of the Animalia than to that of Plantae or any other kingdom.
Whittaker's original reclassification was based on the fundamental difference in nutrition between the Fungi and the Plantae. Unlike plants, which generally gain carbon through photosynthesis, and so are called
autotroph
An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Wo ...
s, fungi do not possess chloroplasts and generally obtain carbon by breaking down and absorbing surrounding materials, and so are called
heterotroph
A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic
saprotrophs. In addition, the substructure of multicellular fungi is different from that of plants, taking the form of many chitinous microscopic strands called
hyphae, which may be further subdivided into cells or may form a
syncytium
A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleu ...
containing many
eukaryotic
Eukaryotes () are organisms whose Cell (biology), cells have a cell nucleus, nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the ...
nuclei. Fruiting bodies, of which
mushroom
A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans.
The standard for the name "mushroom" is ...
s are the most familiar example, are the reproductive structures of fungi, and are unlike any structures produced by plants.
Diversity
The table below shows some species count estimates of different green plant (Viridiplantae) divisions. About 85–90% of all plants are flowering plants. Several projects are currently attempting to collect all plant species in online databases, e.g. the
World Flora Online
World Flora Online is an Internet-based compendium of the world's plant species.
Description
The World Flora Online (WFO) is an open-access database, launched In October 2012 as a follow-up project to The Plant List, with the aim of publishing a ...
and
World Plants both list about 391,000 species.
The naming of plants is governed by the
International Code of Nomenclature for algae, fungi, and plants and
International Code of Nomenclature for Cultivated Plants
The ''International Code of Nomenclature for Cultivated Plants'' (ICNCP), is a guide to the rules and regulations for naming cultigens, plants whose origin or selection is primarily due to intentional human activity. It is also known as Cultivat ...
(see
cultivated plant taxonomy
Cultivated plant taxonomy is the study of the theory and practice of the science that identifies, describes, classifies, and names cultigens—those plants whose origin or selection is primarily due to intentional human activity. Cultivated ...
).
Evolution
The evolution of plants has resulted in increasing
levels of complexity, from the earliest
algal mat
Algal mats are one of many types of microbial mat that forms on the surface of water or rocks. They are typically composed of blue-green cyanobacteria and sediments. Formation occurs when alternating layers of blue-green bacteria and sediments ar ...
s, through
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,
lycopod
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 ...
s,
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s to the complex
gymnosperm
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
s and
angiosperms of today. Plants in all of these groups continue to thrive, especially in the environments in which they evolved.
An algal scum formed on the land , but it was not until the
Ordovician Period
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.
Th ...
, around , that land plants appeared. However, new evidence from the study of carbon isotope ratios in Precambrian rocks has suggested that complex photosynthetic plants developed on the earth over 1000 m.y.a. For more than a century it has been assumed that the ancestors of land plants evolved in aquatic environments and then adapted to a life on land, an idea usually credited to botanist
Frederick Orpen Bower
Frederick Orpen Bower FRSE FRS (4 November 1855 – 11 April 1948) was an English botanist. He was elected a Fellow of the Royal Society in 1891. He was awarded the Gold Medal of the Linnean Society in 1909 and the Darwin Medal of the Ro ...
in his 1908 book ''The Origin of a Land Flora''. A recent alternative view, supported by genetic evidence, is that they evolved from terrestrial single-celled algae, and that even the common ancestor of red and green algae, and the unicellular freshwater algae
glaucophyte
The glaucophytes, also known as glaucocystophytes or glaucocystids, are a small group of unicellular algae found in freshwater and moist terrestrial environments, less common today than they were during the Proterozoic. The stated number of speci ...
s, originated in a terrestrial environment in freshwater biofilms or microbial mats. Primitive land plants began to diversify in the late
Silurian Period
The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
, around , and the results of their diversification are displayed in remarkable detail in an early
Devonian fossil assemblage from the
Rhynie chert
The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is locate ...
. This chert preserved early plants in cellular detail, petrified in volcanic springs. By the middle of the Devonian Period most of the features recognised in plants today are present, including roots, leaves and secondary wood, and by late Devonian times seeds had evolved.
Late Devonian plants had thereby reached a degree of sophistication that allowed them to form forests of tall trees. Evolutionary innovation continued in the Carboniferous and later geological periods and is ongoing today. Most plant groups were relatively unscathed by the
Permo-Triassic extinction event, although the structures of communities changed. This may have set the scene for the evolution of flowering plants in the Triassic (~), which exploded in the Cretaceous and Tertiary. The latest major group of plants to evolve were the grasses, which became important in the mid Tertiary, from around . The grasses, as well as many other groups, evolved new mechanisms of metabolism to survive the low and warm, dry conditions of the tropics over the last .
A 1997 proposed
phylogenetic tree of Plantae, after Kenrick and Crane, is as follows, with modification to the Pteridophyta from Smith ''et al.'' The
Prasinophyceae
The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapma ...
are a
paraphyletic assemblage of early diverging green algal lineages, but are treated as a group outside the Chlorophyta:
later authors have not followed this suggestion.
A newer proposed classification follows Leliaert et al. 2011
and modified with Silar 2016
for the green algae clades and Novíkov & Barabaš-Krasni 2015 for the land plants clade. Notice that the Prasinophyceae are here placed inside the Chlorophyta.
Later, a phylogeny based on genomes and transcriptomes from 1,153 plant species was proposed.
The placing of algal groups is supported by phylogenies based on genomes from the Mesostigmatophyceae and Chlorokybophyceae that have since been sequenced.
The classification of Bryophyta is supported both by Puttick ''et al.'' 2018,
and by phylogenies involving the hornwort genomes that have also since been sequenced.
Embryophytes
The plants that are likely most familiar to us are the
multicellular
A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism.
All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially un ...
land plants, called
embryophyte
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 si ...
s. Embryophytes include the
vascular plant
Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s, such as ferns, conifers and flowering plants. They also include the ''
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'', of which
mosses and
liverworts are the most common.
All of these plants have
eukaryotic
Eukaryotes () are organisms whose Cell (biology), cells have a cell nucleus, nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the ...
cells with
cell walls composed of
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 w ...
, and most obtain their energy through
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 i ...
, using
light
Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 te ...
, water and
carbon dioxide
Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
to synthesize food. About three hundred plant species do not photosynthesize but are
parasite
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 ...
s on other species of photosynthetic plants. Embryophytes are distinguished from
green algae, which represent a mode of photosynthetic life similar to the kind modern plants are believed to have evolved from, by having specialized reproductive organs protected by non-reproductive tissues.
Bryophytes first appeared during the early
Paleozoic
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 ' ...
. They mainly live in habitats where moisture is available for significant periods, although some species, such as
''Targionia'', are desiccation-tolerant. Most species of bryophytes remain small throughout their life-cycle. This involves an alternation between two generations: a
haploid stage, called the
gametophyte
A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...
, and a
diploid stage, called the
sporophyte. In bryophytes, the sporophyte is always unbranched and remains nutritionally dependent on its parent gametophyte. The embryophytes have the ability to secrete a
cuticle on their outer surface, a waxy layer that confers resistance to desiccation. In the
mosses and
hornworts a cuticle is usually only produced on the sporophyte.
Stomata are absent from liverworts, but occur on the sporangia of mosses and hornworts, allowing gas exchange.
Vascular plants first appeared during the
Silurian period, and by the
Devonian had diversified and spread into many different terrestrial environments. They developed a number of adaptations that allowed them to spread into increasingly more arid places, notably 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
Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is c ...
, that transport water and food throughout the organism. Root systems capable of obtaining soil water and nutrients also evolved during the Devonian. In modern vascular plants, the sporophyte is typically large, branched, nutritionally independent and long-lived, but there is increasing evidence that Paleozoic gametophytes were just as complex as the sporophytes. The gametophytes of all vascular plant groups evolved to become reduced in size and prominence in the life cycle.
In seed plants, the
microgametophyte
A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sex ...
is reduced from a multicellular free-living organism to a few cells in a pollen grain and the miniaturised
megagametophyte remains inside the megasporangium, attached to and dependent on the parent plant. A megasporangium enclosed in a protective layer called an integument is known as an
ovule
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the '' integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the ...
. After fertilisation by means of sperm produced by
pollen grains, an embryo sporophyte develops inside the ovule. The integument becomes a seed coat, and the ovule develops into a seed. Seed plants can survive and reproduce in extremely arid conditions, because they are not dependent on free water for the movement of sperm, or the development of free living gametophytes.
The first seed plants,
pteridosperms (seed ferns), now extinct, appeared in the Devonian and diversified through the Carboniferous. They were the ancestors of modern
gymnosperm
The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
s, of which four surviving groups are widespread today, particularly the
conifers, which are dominant
tree
In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s in several
biome
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 ...
s. The name gymnosperm comes from the
Greek
Greek may refer to:
Greece
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group.
*Greek language, a branch of the Indo-European language family.
**Proto-Greek language, the assumed last common ancestor ...
, a composite of ( ) and ( ), as the ovules and subsequent seeds are not enclosed in a protective structure (carpels or fruit), but are borne naked, typically on cone scales.
Fossils
Plant
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s include roots, wood, leaves, seeds, fruit,
pollen,
spores,
phytolith
Phytoliths (from Greek, "plant stone") are rigid, microscopic structures made of silica, found in some plant tissues and persisting after the decay of the plant. These plants take up silica from the soil, whereupon it is deposited within different ...
s, and
amber
Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In ...
(the fossilized resin produced by some plants). Fossil land plants are recorded in terrestrial, lacustrine, fluvial and nearshore marine sediments.
Pollen,
spores
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, ...
and algae (
dinoflagellates and
acritarch
Acritarchs are organic microfossils, known from approximately 1800 million years ago to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity refle ...
s) are used for dating sedimentary rock sequences. The remains of fossil plants are not as common as fossil animals, although plant fossils are locally abundant in many regions worldwide.
The earliest fossils clearly assignable to Kingdom Plantae are fossil green algae from the
Cambrian. These fossils resemble
calcified
Calcification is the accumulation of calcium salts in a body tissue. It normally occurs in the formation of bone, but calcium can be deposited abnormally in soft tissue,Miller, J. D. Cardiovascular calcification: Orbicular origins. ''Nature Ma ...
multicellular
A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism.
All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially un ...
members of the
Dasycladales
Dasycladales is an order of large unicellular green algae in the class Ulvophyceae. It contains two families, the Dasycladaceae and the Polyphysaceae.
These single celled algae are from 2 mm to 200 mm long. They live on substrates ...
. Earlier
Precambrian fossils are known that resemble single-cell green algae, but definitive identity with that group of algae is uncertain.
The earliest fossils attributed to green algae date from the
Precambrian (ca. 1200 mya). The resistant outer walls of
prasinophyte
The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapma ...
cysts (known as phycomata) are well preserved in fossil deposits of the
Paleozoic
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 ' ...
(ca. 250–540 mya). A filamentous fossil (''Proterocladus'') from middle Neoproterozoic deposits (ca. 750 mya) has been attributed to the
Cladophorales
Cladophorales are an order of green algae, in the class Ulvophyceae
The Ulvophyceae or ulvophytes are a class of green algae, distinguished mainly on the basis of ultrastructural morphology, life cycle and molecular phylogenetic data.Graham L ...
, while the oldest reliable records of the
Bryopsidales
Bryopsidales is an order of green algae, in the class Ulvophyceae.
Characteristics
The thallus is filamentous and much branched and may be packed into a mass. It is coenocytic, having multi-nucleate cells consisting of cytoplasm contained withi ...
,
Dasycladales
Dasycladales is an order of large unicellular green algae in the class Ulvophyceae. It contains two families, the Dasycladaceae and the Polyphysaceae.
These single celled algae are from 2 mm to 200 mm long. They live on substrates ...
) and
stoneworts
Charales is an order of freshwater green algae in the division Charophyta, class Charophyceae, commonly known as stoneworts. Depending on the treatment of the genus '' Nitellopsis'', living (extant) species are placed into either one family (Ch ...
are from the
Paleozoic
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 ' ...
.
The oldest known fossils of embryophytes date from 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.
T ...
, though such fossils are fragmentary. By the
Silurian, fossils of whole plants are preserved, including the simple vascular plant ''
Cooksonia
''Cooksonia'' is an extinct group of primitive land plants, treated as a genus, although probably not monophyletic. The earliest ''Cooksonia'' date from the middle of the Silurian (the Wenlock epoch); the group continued to be an important comp ...
'' in mid-Silurian and the much larger and more complex
lycophyte
The lycophytes, when broadly circumscribed, are a vascular plant (tracheophyte) subgroup of the kingdom Plantae. They are sometimes placed in a division Lycopodiophyta or Lycophyta or in a subdivision Lycopodiophytina. They are one of the oldes ...
''
Baragwanathia longifolia
''Baragwanathia'' is a genus of extinct lycopsid plants of Late Silurian to Early Devonian age (), fossils of which have been found in Australia, Canada, China and Czechia. The name derives from William Baragwanath who discovered the first spec ...
'' in late Silurian. From the early Devonian
Rhynie chert
The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is locate ...
, detailed fossils of lycophytes and
rhyniophyte
The rhyniophytes are a group of extinct early vascular plants that are considered to be similar to the genus '' Rhynia'', found in the Early Devonian (around ). Sources vary in the name and rank used for this group, some treating it as the class ...
s have been found that show details of the individual cells within the plant organs and the symbiotic association of these plants with fungi of the order
Glomales. The
Devonian period
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, w ...
also saw the evolution of leaves and roots, and the first modern tree, ''
Archaeopteris''. This tree with fern-like foliage and a trunk with conifer-like wood was
heterosporous
Heterospory is the production of spores of two different sizes and sexes by the sporophytes of land plants. The smaller of these, the microspore, is male and the larger megaspore is female. Heterospory evolved during the Devonian period from is ...
producing spores of two different sizes, an early step in the evolution of seeds.
The
Coal measure
In lithostratigraphy, the coal measures are the coal-bearing part of the Upper Carboniferous System. In the United Kingdom, the Coal Measures Group consists of the Upper Coal Measures Formation, the Middle Coal Measures Formation and the Lower Coal ...
s are a major source of
Paleozoic
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 ' ...
plant fossils, with many groups of plants in existence at this time. The spoil heaps of coal mines are the best places to collect;
coal
Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen.
Coal is formed when ...
itself is the remains of fossilised plants, though structural detail of the plant fossils is rarely visible in coal. In the
Fossil Grove at
Victoria Park Victoria Park may refer to:
Places Australia
* Victoria Park Nature Reserve, a protected area in Northern Rivers region, New South Wales
* Victoria Park, Adelaide, a park and racecourse
* Victoria Park, Brisbane, a public park and former golf ...
in
Glasgow
Glasgow ( ; sco, Glesca or ; gd, Glaschu ) is the most populous city in Scotland and the fourth-most populous city in the United Kingdom, as well as being the 27th largest city by population in Europe. In 2020, it had an estimated popul ...
, Scotland, the stumps of ''
Lepidodendron
''Lepidodendron'' is an extinct genus of primitive vascular plants belonging to the family Lepidodendraceae, part of a group of Lycopodiopsida known as scale trees or arborescent lycophytes, related to quillworts and lycopsids (club mosses). Th ...
'' trees are found in their original growth positions.
The fossilized remains of
conifer and
angiosperm root
In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the su ...
s,
stems and
branch
A branch, sometimes called a ramus in botany, is a woody structural member connected to the central trunk of a tree (or sometimes a shrub). Large branches are known as boughs and small branches are known as twigs. The term ''twig'' usually ...
es may be locally abundant in lake and inshore
sedimentary rock
Sedimentary rocks are types of rock that are formed by the accumulation or deposition of mineral or organic particles at Earth's surface, followed by cementation. Sedimentation is the collective name for processes that cause these particles ...
s from the
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
and
Cenozoic eras.
Sequoia and its allies,
magnolia
''Magnolia'' is a large genus of about 210 to 340The number of species in the genus ''Magnolia'' depends on the taxonomic view that one takes up. Recent molecular and morphological research shows that former genera ''Talauma'', ''Dugandiodendro ...
,
oak
An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ''L ...
, and
palms are often found.
Petrified wood
Petrified wood, also known as petrified tree (from Ancient Greek meaning 'rock' or 'stone'; literally 'wood turned into stone'), is the name given to a special type of '' fossilized wood'', the fossilized remains of terrestrial vegetation. ' ...
is common in some parts of the world, and is most frequently found in arid or desert areas where it is more readily exposed by
erosion
Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is dis ...
. Petrified wood is often heavily
silicified
In geology, petrifaction or petrification () is the process by which organic material becomes a fossil through the replacement of the original material and the filling of the original pore spaces with minerals. Petrified wood typifies this proce ...
(the
organic material
Organic matter, organic material, or natural organic matter refers to the large source of carbon-based compounds found within natural and engineered, terrestrial, and aquatic environments. It is matter composed of organic compounds that have c ...
replaced by
silicon dioxide
Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is one ...
), and the impregnated tissue is often preserved in fine detail. Such specimens may be cut and polished using
lapidary
Lapidary (from the Latin ) is the practice of shaping stone, minerals, or gemstones into decorative items such as cabochons, engraved gems (including cameos), and faceted designs. A person who practices lapidary is known as a lapidarist. A lap ...
equipment. Fossil forests of petrified wood have been found in all continents.
Fossils of seed ferns such as ''
Glossopteris
''Glossopteris'' tymology: from Ancient Greek γλῶσσα (glôssa, " tongue ") + πτερίς (pterís, " fern ")is the largest and best-known genus of the extinct Permian order of seed ferns known as Glossopteridales (also known as Arberia ...
'' are widely distributed throughout several continents of the
Southern Hemisphere, a fact that gave support to
Alfred Wegener
Alfred Lothar Wegener (; ; 1 November 1880 – November 1930) was a German climatologist, geologist, geophysicist, meteorologist, and polar researcher.
During his lifetime he was primarily known for his achievements in meteorology and ...
's early ideas regarding
Continental drift
Continental drift is the hypothesis that the Earth's continents have moved over geologic time relative to each other, thus appearing to have "drifted" across the ocean bed. The idea of continental drift has been subsumed into the science of pl ...
theory.
Structure, growth, and development
Most of the solid material in a plant is taken from the
atmosphere. Through the process of
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 i ...
, most plants use the energy in
sunlight to convert
carbon dioxide
Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
from the atmosphere, plus
water
Water (chemical formula ) is an Inorganic compound, inorganic, transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living ...
, into simple
sugars. These sugars are then used as building blocks and form the main structural component of the plant.
Chlorophyll, a green-colored,
magnesium
Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...
-containing
pigment
A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...
is essential to this process; it is generally present in plant
leaves, and often in other plant parts as well.
Parasitic plant
A parasitic plant is a plant that derives some or all of its nutritional requirements from another living plant. They make up about 1% of angiosperms and are found in almost every biome. All parasitic plants develop a specialized organ called the ...
s, on the other hand, use the resources of their host to provide the materials needed for metabolism and growth.
Plants usually rely on soil primarily for support and water (in quantitative terms), but they also obtain chemical compound, compounds of nitrogen, phosphorus, potassium, magnesium and other elemental nutrients from the soil. Epiphyte, Epiphytic and lithophyte, lithophytic plants depend on air and nearby debris for nutrients, and carnivorous plants supplement their nutrient requirements, particularly for nitrogen and phosphorus, with insect prey that they capture. For the majority of plants to grow successfully they also require oxygen in the atmosphere and around their roots (soil gas) for Cellular respiration, respiration. Plants use oxygen and glucose (which may be produced from stored
starch) to provide energy. Some plants grow as submerged aquatics, using oxygen dissolved in the surrounding water, and a few specialized vascular plants, such as mangroves and reed (''Phragmites australis''),
can grow with their roots in anoxic waters, anoxic conditions.
Factors affecting growth
The genome of a plant controls its growth. For example, selected varieties or genotypes of wheat grow rapidly, maturing within 110 days, whereas others, in the same environmental conditions, grow more slowly and mature within 155 days.
[Robbins, W.W.; Weier, T.E.; ''et al.''., ''Botany: Plant Science'', 3rd edition, Wiley International, New York, 1965.]
Growth is also determined by environmental factors, such as temperature, available
water
Water (chemical formula ) is an Inorganic compound, inorganic, transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living ...
, available
light
Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 te ...
,
carbon dioxide
Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
and available nutrients in the soil. Any change in the availability of these external conditions will be reflected in the plant's growth and the timing of its development.
Biotic factors also affect plant growth. Plants can be so crowded that no single individual produces normal growth, causing etiolation and chlorosis. Optimal plant growth can be hampered by grazing animals, suboptimal soil composition, lack of mycorrhizal fungi, and attacks by insects or plant pathology, plant diseases, including those caused by bacteria, fungi, viruses, and nematodes.
Simple plants like algae may have short life spans as individuals, but their populations are commonly seasonal. Annual plants grow and reproduce within one growing season, biennial plants grow for two growing seasons and usually reproduce in second year, and perennial plants live for many growing seasons and once mature will often reproduce annually. These designations often depend on climate and other environmental factors. Plants that are annual in alpine climate, alpine or temperate regions can be biennial or perennial in warmer climates. Among the vascular plants, perennials include both evergreens that keep their leaves the entire year, and deciduous plants that lose their leaves for some part of it. In temperate and boreal climates, they generally lose their leaves during the winter; many tropical plants lose their leaves during the dry season.
The growth rate of plants is extremely variable. Some mosses grow less than 0.001 millimeters per hour (mm/h), while most trees grow 0.025–0.250 mm/h. Some climbing species, such as kudzu, which do not need to produce thick supportive tissue, may grow up to 12.5 mm/h. Plants protect themselves from frost and dehydration stress with antifreeze proteins, Heat shock protein, heat-shock proteins and sugars (sucrose is common). LEA (Late Embryogenesis Abundant proteins, Late Embryogenesis Abundant) protein expression is induced by stresses and protects other proteins from aggregation as a result of desiccation and freezing.
Effects of freezing
When water freezes in plants, the consequences for the plant depend very much on whether the freezing occurs within cells (intracellularly) or outside cells in intercellular spaces.
[Glerum, C. 1985. Frost hardiness of coniferous seedlings: principles and applications. pp. 107–123 ''in ''Duryea, M.L. (Ed.). Proceedings: Evaluating seedling quality: principles, procedures, and predictive abilities of major tests. Workshop, October 1984, Oregon State Univ., For. Res. Lab., Corvallis OR.] Intracellular freezing, which usually kills the cell
[Lyons, J.M.; Raison, J.K.; Steponkus, P.L. 1979. The plant membrane in response to low temperature: an overview. pp. 1–24 ''in'' Lyons, J.M.; Graham, D.; Raison, J.K. (Eds.). Low Temperature Stress in Crop Plants. Academic Press, New York NY.] regardless of the hardiness of the plant and its tissues, seldom occurs in nature because rates of cooling are rarely high enough to support it. Rates of cooling of several degrees Celsius per minute are typically needed to cause intracellular formation of ice.
[Mazur, P. 1977. The role of intracellular freezing in the death of cells cooled at supraoptimal rates. Cryobiology 14:251–272.] At rates of cooling of a few degrees Celsius per hour, segregation of ice occurs in intercellular spaces.
[Sakai, A.; Larcher, W. (Eds.) 1987. Frost Survival of Plants. Springer-Verlag, New York. 321 p.] This may or may not be lethal, depending on the hardiness of the tissue. At freezing temperatures, water in the intercellular spaces of plant tissue freezes first, though the water may remain unfrozen until temperatures drop below .
After the initial formation of intercellular ice, the cells shrink as water is lost to the segregated ice, and the cells undergo freeze-drying. This dehydration is now considered the fundamental cause of freezing injury.
DNA damage and repair
Plants are continuously exposed to a range of biotic and abiotic stresses. These stresses often cause DNA damage (naturally occurring), DNA damage directly, or indirectly via the generation of reactive oxygen species.
Plants are capable of a DNA damage response that is a critical mechanism for maintaining genome stability.
The DNA damage response is particularly important during seed germination, since seed quality tends to deteriorate with age in association with DNA damage accumulation.
During germination repair processes are activated to deal with this accumulated DNA damage.
In particular, single-stranded break, single- and double-stranded break, double-strand breaks in DNA can be DNA repair, repaired.
Double-strand repair in plants often produce DNA junctions with Genetic recombination, structural alterations.
The DNA checkpoint kinase ATM serine/threonine kinase, ATM has a key role in integrating progression through germination with repair responses to the DNA damages accumulated by the aged seed.
Plant cells
Plant cells are typically distinguished by their large water-filled central vacuole,
chloroplasts, and rigid
cell walls that are made up of
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 w ...
, hemicellulose, and pectin. Cell division is also characterized by the development of a phragmoplast for the construction of a cell plate in the late stages of cytokinesis. Just as in animals, plant cells differentiate and develop into multiple cell types. Totipotent meristematic cells can differentiate into vascular tissue, vascular, storage, protective (e.g. epidermis (botany), epidermal layer), or plant sexuality, reproductive tissues, with more primitive plants lacking some tissue types.
Physiology
Photosynthesis
Plants photosynthesis, photosynthesize, which means that they manufacture their own food molecules using energy obtained from
light
Light or visible light is electromagnetic radiation that can be perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nanometres (nm), corresponding to frequencies of 750–420 te ...
. The primary mechanism plants have for capturing light energy is the
pigment
A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...
chlorophyll. All green plants contain two forms of chlorophyll, chlorophyll a, chlorophyll ''a'' and chlorophyll b, chlorophyll ''b''. The latter of these pigments is not found in red or brown algae.
The simple equation of photosynthesis is as follows:
:
6CO2 + 6H2O ->[\text] C6H12O6 + 6O2
Immune system
By means of cells that behave like Nerve, nerves, plants receive and distribute within their systems information about incident light intensity and quality. Incident light that stimulates a chemical reaction in one leaf, will cause a chain reaction of signals to the entire plant via a type of cell termed a ''Vascular bundle, bundle sheath cell''. Researchers, from the Warsaw University of Life Sciences in Poland, found that plants have a specific memory for varying light conditions, which prepares their immune systems against seasonal pathogens. Plants use pattern-recognition receptors to recognize conserved microbial signatures. This recognition triggers an immune response. The first plant receptors of conserved microbial signatures were identified in rice (XA21, 1995) and in ''Arabidopsis thaliana'' (FLS2, 2000). Plants also carry immune receptors that recognize highly variable pathogen effectors. These include the NBS-LRR class of proteins.
Internal distribution
Vascular plants differ from other plants in that nutrients are transported between their different parts through specialized structures, called
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
Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is c ...
. They also have
root
In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the su ...
s for taking up water and minerals. The xylem moves water and minerals from the root to the rest of the plant, and the phloem provides the roots with sugars and other nutrient produced by the leaves.
Genomics
Plants have some of the largest genomes among all organisms. The largest plant genome (in terms of gene number) is that of wheat (''Triticum asestivum''), predicted to encode ≈94,000 genes and thus almost 5 times as many as the human genome. The first plant genome sequenced was that of ''Arabidopsis thaliana'' which encodes about 25,500 genes. In terms of sheer DNA sequence, the smallest published genome is that of the carnivorous Utricularia gibba, bladderwort (''Utricularia gibba)'' at 82 Mb (although it still encodes 28,500 genes) while the largest, from the Picea abies, Norway Spruce (''Picea abies''), extends over 19,600 Mb (encoding about 28,300 genes).
Ecology
The photosynthesis conducted by land plants and algae is the ultimate source of energy and organic material in nearly all ecosystems. Photosynthesis, at first by cyanobacteria and later by photosynthetic eukaryotes, radically changed the composition of the early Earth's anoxic atmosphere, which as a result is now 21% oxygen. Animals and most other organisms are Aerobic organism, aerobic, relying on oxygen; those that do not are confined to relatively rare anaerobic environments. Plants are the Autotroph, primary producers in most terrestrial ecosystems and form the basis of the food web in those ecosystems. Many animals rely on plants for shelter as well as oxygen and food. Plants form about 80% of the world Biomass (ecology), biomass at about of carbon.
Land plants are key components of the water cycle and several other biogeochemical cycles. Some plants have coevolved with nitrogen fixation, nitrogen fixing bacteria, making plants an important part of the nitrogen cycle. Plant roots play an essential role in soil development and the prevention of soil erosion.
Distribution
Plants are distributed almost worldwide. While they inhabit a multitude of
biome
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 ...
s and ecoregions, few can be found beyond the tundras at the northernmost regions of continental shelf, continental shelves. At the southern extremes, plants of the Antarctic flora have adapted tenaciously to the prevailing conditions.
Plants are often the dominant physical and structural component of habitats where they occur. Many of the Earth's
biome
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 ...
s are named for the type of vegetation because plants are the dominant organisms in those biomes, such as grasslands, taiga and tropical rainforest.
Ecological relationships
Numerous animals have coevolved with plants. Many animals pollination, pollinate flowers in exchange for food in the form of pollen or nectar. Many animals biological dispersal, disperse seeds, often by eating
fruit
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
and passing the seeds in their feces. Myrmecophytes are plants that have coevolved with ants. The plant provides a home, and sometimes food, for the ants. In exchange, the ants defend the plant from herbivores and sometimes competing plants. Ant wastes provide organic fertilizer.
The majority of plant species have various kinds of fungi associated with their root systems in a kind of Mutualism (biology), mutualistic symbiosis known as mycorrhiza. The fungi help the plants gain water and mineral nutrients from the soil, while the plant gives the fungi carbohydrates manufactured in photosynthesis. Some plants serve as homes for endophyte, endophytic fungi that protect the plant from herbivores by producing toxins. The fungal endophyte, ''Neotyphodium coenophialum'', in tall fescue (''Festuca arundinacea'') does tremendous economic damage to the cattle industry in the U.S. Many legume plants have nitrogen fixing bacteria in the genus ''Rhizobium'', found in nodules of their roots, that fix nitrogen from the air for the plant to use. In exchange, the plants supply sugars to the bacteria.
Various forms of parasitism are also fairly common among plants, from the semi-parasitic mistletoe that merely takes some nutrients from its host, but still has photosynthetic leaves, to the fully parasitic Orobanche, broomrape and Lathraea, toothwort that acquire all their nutrients through connections to the roots of other plants, and so have no
chlorophyll. Some plants, known as myco-heterotrophs, parasitize mycorrhizal fungi, and hence act as epiparasites on other plants.
Many plants are epiphytes, meaning they grow on other plants, usually trees, without parasitizing them. Epiphytes may indirectly harm their host plant by intercepting mineral nutrients and light that the host would otherwise receive. The weight of large numbers of epiphytes may break tree limbs. Hemiepiphytes like the strangler fig begin as epiphytes but eventually set their own roots and overpower and kill their host. Many orchids, bromeliads,
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s and
mosses often grow as epiphytes. Bromeliad epiphytes accumulate water in leaf axils to form phytotelmata that may contain complex aquatic food webs.
Approximately 630 plants are carnivorous plant, carnivorous, such as the Venus Flytrap (''Dionaea muscipula'') and sundew (''Drosera'' species). They trap small animals and digest them to obtain mineral nutrients, especially nitrogen and phosphorus.
[Barthlott, W.; Porembski, S.; Seine, R.; Theisen, I. 2007. ''The Curious World of Carnivorous Plants: A Comprehensive Guide to Their Biology and Cultivation.'' Timber Press: Portland, Oregon.]
Competition
Competition occurs when members of the same species, or several different species, compete for shared resources in a given habitat. According to the competitive exclusion principle, when environmental resources are limited, species cannot occupy nor be supported by identical niches.
Eventually, one species will out-compete the other, which will push the disadvantaged species to extinction.
In regard to plants, competition tends to negatively affect their growth when competing for shared resources.
These shared resources commonly include space for growth, sunlight, water and nutrients. Light is an important resource because it is necessary for photosynthesis.
Plants use their leaves to shade other plants from sunlight and grow quickly to maximize their own expose.
Water is also important for photosynthesis, and plants have different root systems to maximize water uptake from soil.
Some plants have deep roots that are able to locate water stored deep underground, and others have shallower roots that are capable of extending longer distances to collect recent rainwater.
Minerals are also important for plant growth and development, where deficiencies can occur if nutrient needs are not met. Common nutrients competed for amongst plants include nitrogen and phosphorus. Space is also extremely important for a growing and developing plant.
Having optimal space makes it more likely that leaves are exposed to sufficient amounts of sunlight and are not overcrowded in order for photosynthesis to occur.
If an old tree dies, then competition arises amongst a number of trees to replace it.
Those that are less effective competitors are less likely to contribute to the next generation of offspring.
Contrary to the belief that plants are always in competition, new research has found that in a harsh environment mature plants sheltering seedlings help the smaller plant survive.
Importance
Cultivation
The study of plant uses by people is called economic botany or ethnobotany. Human cultivation of plants is part of agriculture, which is the basis of human civilization. ''Plant agriculture'' is subdivided into agronomy, horticulture and forestry.
Food
Humans depend on plants for food, either directly or as feed for domestic animals. Agriculture deals with the production of food crops, and has History of agriculture, played a key role in the history of world civilizations. Agriculture includes agronomy for arable crops, horticulture for vegetables and fruit, and forestry for timber.
About 7,000 species of plant have been used for food, though most of today's food is derived from only 30 species. The major staple crop, staples include cereals such as rice and wheat, starchy roots and tubers such as cassava and potato, and legumes such as peas and beans. Vegetable oils such as olive oil and palm oil provide lipids, while
fruit
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
and
vegetable
Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems, ...
s contribute vitamins and minerals to the diet.
Medicines
Medicinal plants are a primary source of organic compounds, both for their medicinal and physiological effects, and for the industrial organic synthesis, synthesis of a vast array of organic chemicals. Many hundreds of medicines are derived from plants, both traditional medicines used in herbalism
and chemical substances purified from plants or first identified in them, sometimes by ethnobotany, ethnobotanical search, and then organic synthesis, synthesised for use in modern medicine. Modern medicines derived from plants include aspirin, taxol, morphine, quinine, reserpine, colchicine, digitalis and vincristine. List of plants used in herbalism, Plants used in herbalism include Ginkgo biloba, ginkgo, echinacea, feverfew, and Saint John's wort. The pharmacopoeia of Dioscorides, ''De Materia Medica'', describing some 600 medicinal plants, was written between 50 and 70 AD and remained in use in Europe and the Middle East until around 1600 AD; it was the precursor of all modern pharmacopoeias.
Nonfood products
Plants grown as industrial crops are the source of a wide range of products used in manufacturing, sometimes so intensively as to risk harm to the environment. Nonfood products include essential oils, natural dyes, pigments, waxes, resins, tannins, alkaloids,
amber
Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In ...
and Cork material, cork. Products derived from plants include soaps, shampoos, perfumes, cosmetics, paint, varnish, turpentine, rubber, latex, lubricants, linoleum, plastics, inks, and Gum (botany), gums. Renewable fuels from plants include firewood, peat and other biofuels.
The fossil fuels
coal
Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen.
Coal is formed when ...
, petroleum and natural gas are derived from the remains of aquatic organisms including phytoplankton in geological time.
Structural resources and fibres from plants are used to construct dwellings and to manufacture clothing. Wood is used not only for buildings, boats, and furniture, but also for smaller items such as musical instruments and sports equipment. Wood is Pulp (paper), pulped to make paper and cardboard. Cloth is often made from cotton, flax, ramie or synthetic fibres such as rayon and acetate derived from plant
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 w ...
. Thread (yarn), Thread used to sew cloth likewise comes in large part from cotton.
Aesthetic uses
Thousands of plant species are cultivated for aesthetic purposes as well as to provide shade, modify temperatures, reduce wind, abate noise, provide privacy, and prevent soil erosion. Plants are the basis of a multibillion-dollar per year tourism industry, which includes travel to garden tourism, historic gardens, national parks, rainforests, forests with colorful autumn leaves, and festivals such as Hanami, Japan's and National Cherry Blossom Festival, America's cherry blossom festivals.
While some gardens are planted with food crops, many are planted for aesthetic, ornamental, or conservation purposes. Arboretums and botanical gardens are public collections of living plants. In private outdoor gardens, lawn grasses, shade trees, ornamental trees, shrubs, vines, herbaceous perennials and bedding plants are used. Gardens may cultivate the plants in a naturalistic state, or may sculpture their growth, as with topiary or espalier. Gardening is the most popular leisure activity in the U.S., and working with plants or horticulture therapy is beneficial for rehabilitating people with disabilities.
Plants may also be grown or kept indoors as houseplants, or in specialized buildings such as greenhouses that are designed for the care and cultivation of living plants. Venus Flytrap, sensitive plant and resurrection plant are examples of plants sold as novelties. There are also art forms specializing in the arrangement of cut or living plant, such as bonsai, ikebana, and the arrangement of cut or dried flowers. Ornamental plants have sometimes changed the course of history, as in tulip mania, tulipomania.
Architectural designs resembling plants appear in the capitals of Ancient Egyptian columns, which were carved to resemble either the Nymphaea lotus, Egyptian white lotus or the Cyperus papyrus, papyrus. Images of plants are often used in painting and photography, as well as on textiles, money, stamps, flags and coats of arms.
Scientific and cultural uses
Basic biological research has often been done with plants. In genetics, the breeding of pea plants allowed Gregor Mendel to derive the basic laws governing inheritance, and examination of chromosomes in maize allowed Barbara McClintock to demonstrate their connection to inherited traits. The plant ''Arabidopsis thaliana'' is used in laboratories as a model organism to understand how genes control the growth and development of plant structures. NASA predicts that space stations or space colonies will one day rely on plants for Controlled Ecological Life Support System, life support.
Ancient trees are revered and many are List of famous trees, famous. Tree rings themselves are an important method of dating in archeology, and serve as a record of past climates.
Plants figure prominently in Trees in mythology, mythology, religion and List of fictional plants, literature. They are used as National emblem, national and state emblems, including List of U.S. state trees, state trees and state flowers. Plants are often used as memorials, gifts and to mark special occasions such as births, deaths, weddings and holidays. The arrangement of flowers may be used to send hidden Language of flowers, messages.
Negative effects
Weeds are commercially or aesthetically undesirable plants growing in managed environments such as agriculture, farms, urban areas, gardens, lawns, and parks. People have spread plants beyond their native ranges and some of these introduced plants become invasive species, invasive, damaging existing ecosystems by displacing native species, and sometimes becoming serious weeds of cultivation.
Plants may cause harm to animals, including people. Plants that produce anemophily, windblown pollen invoke allergic reactions in people who suffer from hay fever. A wide variety of plants are List of poisonous plants, poisonous. Toxalbumins are plant poisons fatal to most mammals and act as a serious deterrent to consumption. Several plants cause skin irritations when touched, such as poison ivy. Certain plants contain psychotropic secondary metabolite, chemicals, which are extracted and ingested or smoked, including nicotine from tobacco, cannabinoids from Cannabis sativa, cocaine from Erythroxylon coca and opium from opium poppy. Smoking causes damage to health or even death, while some drugs may also be harmful or fatal to people. Both illegal and legal drugs derived from plants may have negative effects on the economy, affecting worker productivity and law enforcement costs.
See also
* Biosphere
* Evolutionary history of plants
* Plant defense against herbivory
* Plant identification
* Plant reproduction
* Plant to plant communication via mycorrhizal networks
* The Plant List
References
Further reading
;General:
* Evans, L.T. (1998). ''Feeding the Ten Billion – Plants and Population Growth''. Cambridge University Press. Paperback, 247 pages. .
* Kenrick, Paul & Crane, Peter R. (1997). ''The Origin and Early Diversification of Land Plants: A Cladistic Study''. Washington, D.C.: Smithsonian Institution Press. .
* Raven, Peter H.; Evert, Ray F.; & Eichhorn, Susan E. (2005). ''Biology of Plants'' (7th ed.). New York: W.H. Freeman and Company. .
* Taylor, Thomas N. & Taylor, Edith L. (1993). ''The Biology and Evolution of Fossil Plants''. Englewood Cliffs, NJ: Prentice Hall. .
*
;Species estimates and counts:
* International Union for Conservation of Nature and Natural Resources (IUCN) Species Survival Commission (2004). IUCN Red List]
The IUCN Red List of Threatened Species
*
External links
* (requires Microsoft Silverlight)
*
Index Nominum AlgarumPlant Resources of Tropical AfricaTree of Life
;Botanical and vegetation databases
African Plants Initiative databaseAustraliaChilean plants at ''Chilebosque''e-Floras (Flora of China, Flora of North America and others)Flora of Central Europe
Flora of North AmericaMeet the Plants-National Tropical Botanical GardenLady Bird Johnson Wildflower Center – Native Plant Information Network at University of Texas, AustinThe Plant ListUnited States Department of Agriculturenot limited to continental US species
{{Authority control
Plants,
Kingdoms (biology), Plants