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Plants are predominantly
photosynthetic Photosynthesis is a process used by plants and other organisms to convert Conversion or convert may refer to: Arts, entertainment, and media * Conversion (Doctor Who audio), "Conversion" (''Doctor Who'' audio), an episode of the audio drama ' ...
eukaryote Eukaryotes () are organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interact ...

eukaryote
s of the
kingdom Kingdom may refer to: Monarchy * A type of monarchy * A realm ruled by: **A king, during the reign of a male monarch **A queen regnant, during the reign of a female monarch Taxonomy * Kingdom (biology), a category in biological taxonomy Arts an ...
Plantae. Historically, the plant kingdom encompassed all living things that were not
animal Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells ...

animal
s, and included
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic Photosynthesis is a process used by plants and other organisms to convert Conversion or convert may refer to: Arts, entertainment, and media * Co ...

algae
and
fungi A fungus (plural The plural (sometimes abbreviated An abbreviation (from Latin ''brevis'', meaning ''short'') is a shortened form of a word or phrase, by any method. It may consist of a group of letters, or words taken from the full ...

fungi
; however, all current definitions of Plantae exclude the fungi and some algae, as well as the
prokaryote A prokaryote () is a single-celled organism A unicellular organism, also known as a single-celled organism, is an organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contig ...
s (the
archaea Archaea ( ; singular archaeon ) constitute a domain Domain may refer to: Mathematics *Domain of a function, the set of input values for which the (total) function is defined **Domain of definition of a partial function **Natural domain of a pa ...

archaea
and
bacteria Bacteria (; common noun 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. Typ ...

bacteria
). By one definition, plants form the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants - on a phylogenetic tree. R ...

clade
Viridiplantae Viridiplantae (literally "green plants") are a clade A clade (), also known as a monophyletic group or natural group, is a group of organism In biology, an organism () is any organic, life, living system that functions as an indi ...
(Latin name for "green plants"), a group that includes the
flowering plant Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greec ...

flowering plant
s,
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single e ...
s and other
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also ...
s,
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia art ...

fern
s and their allies,
hornwort Hornworts are a group of bryophyte Bryophytes are an informal group consisting of three divisions Division or divider may refer to: Mathematics *Division (mathematics) Division is one of the four basic operations of arithmetic, the ways th ...
s,
liverworts The Marchantiophyta () are a division of non-vascular plant, non-vascular embryophyte, land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plan ...

liverworts
,
moss Mosses are small, non-vascular flower A flower, sometimes known as a bloom or blossom Cherry blossoms in Paris in full bloom. In botany, blossoms are the flowers of stone fruit fruit tree, trees (genus ''Prunus'') and of some other plant ...

moss
es, and the
green algae The green algae (singular: green alga) are a large, informal grouping of algae consisting of the Chlorophyta and Charophyta/Streptophyta, which are now placed in separate Division (botany), divisions, together with the more basal Mesostigmatoph ...

green algae
, but excludes the
red Red is the color at the long wavelength end of the visible spectrum of light, next to orange and opposite violet. It has a dominant wavelength Image:dominant wavelength.png, frame, Dominant/complementary wavelength example on the CIE color ...
and
brown algae The brown algae (singular: alga), comprising the class (biology), class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Most brown algae live in marine enviro ...
. Most plants are
multicellular A multicellular organism is an organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular ...

multicellular
organisms. Green plants obtain most of their energy from
sunlight Sunlight is a portion of the given off by the , in particular , , and light. On , sunlight is and through , and is obvious as when the Sun is above the . When direct is not blocked by s, it is experienced as sunshine, a combination of b ...

sunlight
via
photosynthesis Photosynthesis is a process used by plants and other organisms to convert Conversion or convert may refer to: Arts, entertainment, and media * Conversion (Doctor Who audio), "Conversion" (''Doctor Who'' audio), an episode of the audio drama ' ...

photosynthesis
by primary
chloroplast A chloroplast is a type of membrane-bound organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structure ...

chloroplast
s 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 mutualism (biology), mutualistic relationship. (The term endosymbiosis is from the Greek language, Greek: ...

endosymbiosis
with
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the grammatical number, grammatical ...

cyanobacteria
. Their chloroplasts contain
chlorophyll Chlorophyll (also chlorophyl) is any of several related green 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 ...

chlorophyll
s a and b, which gives them their green color. Some plants are
parasitic Parasitism is a Symbiosis, symbiotic biological interactions, relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is adaptation (biology), ad ...

parasitic
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 Reproduction (or procreation or breeding) is the biological process Biological processes are those processes that are vital for an organism In biology, an organism (from Ancient Greek, ...
and
alternation of generations Alternation of generations (also known as metagenesis or heterogenesis) is the type of that occurs in those s and in the and the that have distinct haploid sexual and diploid asexual stages. In these groups, a with ''n'' s alternates with ...

alternation of generations
, although
asexual reproduction Asexual reproduction is a type of reproduction Reproduction (or procreation or breeding) is the biological process Biological processes are those processes that are vital for an organism In biology, an organism (from Ancient Gree ...
is also common. There are about 320,000
species In biology, a species is the basic unit of biological classification, 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 individu ...

species
of plants, of which the great majority, some 260–290 thousand, 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 A grain is a small, hard, dry seed A seed is an embryonic ''Embryonic'' is the twelfth studio album by experimental rock band the Flaming Lips released on October 13, 2009, on Warner Bros. Records, Warner Bros. The band's first double album ...

grain
,
fruit In 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 ...

fruit
, 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 A flower, som ...

vegetable
s also form basic human foods and have been
domesticated Domestication is a sustained multi-generational relationship in which one group of organisms assumes a significant degree of influence over the reproduction and care of another group to secure a more predictable supply of resources from that sec ...

domesticated
for millennia. Plants have many
cultural Culture () is an umbrella term which encompasses the social behavior Social behavior is behavior Behavior (American English) or behaviour (British English; American and British English spelling differences#-our, -or, see spelling diff ...
and other uses, as ornaments,
building materials Building material is material used for construction. Many naturally occurring substances, such as clay, rocks, sand, and wood, even twigs and leaves, have been used to construct buildings. Apart from naturally occurring materials, many man-mad ...
,
writing material Writing material refers to the materials that provide the surfaces on which humans use writing instruments A writing implement or writing instrument is an object used to produce writing Writing is a medium of human communication that involv ...
and, in great variety, they have been the
source of medicines
source of medicines
and
psychoactive drug A psychoactive drug, psychopharmaceutical, psychoactive agent, or psychotropic drug, is a chemical substance that changes nervous system function and results in alterations in perception, mood (psychology), mood, consciousness, cognition, or beha ...
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 wo ...

botany
, a branch of
biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanisms, Development ...

biology
.


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 A philosopher is someone who practices philosophy Philosophy (from , ) is the study of general and fundamental questio ...

Aristotle
(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 Nobility#Ennoblement, ennoblement as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomi ...

Linnaeus
(1707–1778) created the basis of the modern system of
scientific classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxon, taxa (singular: ...

scientific classification
, these two groups became the kingdoms Vegetabilia (later Metaphyta or Plantae) and
Animalia Animals (also called Metazoa) are multicellular A multicellular organism is an organism In biology, an organism () is any organic, life, living system that functions as an individual entity. All organisms are composed of cells ...

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 (plural The plural (sometimes abbreviated An abbreviation (from Latin ''brevis'', meaning ''short'') is a shortened form of a word or phrase, by any method. It may consist of a group of letters, or words taken from the full ...

fungi
and several groups of
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic Photosynthesis is a process used by plants and other organisms to convert Conversion or convert may refer to: Arts, entertainment, and media * Co ...

algae
were removed to new kingdoms. However, these organisms are still often 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 In chemistry Chemistry is the study of the properties and behavior of . It is a that covers the that make up matter to the composed of s, s and s: their composition, structure, properties, behavior ...

cellulose
, 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 Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechani ...
, 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 Cladistics (, from Greek language, Greek , ''kládos'', "branch") is an approach to Taxonomy (biology), biological classification in whi ...

cladogram
, 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
seaweed Seaweed, or macroalgae, refers to thousands of species of , , . The term includes some types of ' (red), ' (brown) and ' (green) macroalgae. Seaweed species such as s provide essential nursery habitat for fisheries and other marine species an ...

seaweed
s range from large multicellular algae to single-celled organisms and are classified into three groups, the
green algae The green algae (singular: green alga) are a large, informal grouping of algae consisting of the Chlorophyta and Charophyta/Streptophyta, which are now placed in separate Division (botany), divisions, together with the more basal Mesostigmatoph ...

green algae
,
red algae Red algae, or Rhodophyta ( , ; ), are one of the oldest groups of eukaryotic algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic Photosynthesis is a process used by plants and other organism ...

red algae
and
brown algae The brown algae (singular: alga), comprising the class (biology), class Phaeophyceae, are a large group of multicellular algae, including many seaweeds located in colder waters within the Northern Hemisphere. Most brown algae live in marine enviro ...
. 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 clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyly, monophyletic – that is, composed of a common ancestor and all its lineage (evolution), lineal descendants - on a phylogenetic tree. R ...

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
chloroplast A chloroplast is a type of membrane-bound organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structure ...

chloroplast
s derived from cyanobacteria containing
chlorophyll Chlorophyll (also chlorophyl) is any of several related green 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 ...

chlorophyll
s ''a'' and ''b'', cell walls containing
cellulose Cellulose is an organic compound In chemistry Chemistry is the study of the properties and behavior of . It is a that covers the that make up matter to the composed of s, s and s: their composition, structure, properties, behavior ...

cellulose
, and food stores in the form of
starch Starch or amylum is a polymeric A polymer (; Greek '' poly-'', "many" + '' -mer'', "part") is a substance Substance may refer to: * Substance (Jainism), a term in Jain ontology to denote the base or owner of attributes * Chemical substance ...
contained within the plastids. They undergo closed
mitosis In cell biology Cell biology (also cellular biology or cytology) is a branch of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical proce ...

mitosis
without
centriole In cell biology Cell biology (also cellular biology or cytology) is a branch of biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, ...

centriole
s, and typically have
mitochondria A mitochondrion (; ) is a double-membrane A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Biological membranes include cell membranes ...

mitochondria
with flat cristae. The
chloroplast A chloroplast is a type of membrane-bound organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structure ...

chloroplast
s of green plants are surrounded by two membranes, suggesting they originated directly from endosymbiotic
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the grammatical number, grammatical ...

cyanobacteria
. Two additional groups, the
Rhodophyta 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 majorit ...
(red algae) and
Glaucophyta The glaucophytes, also known as glaucocystophytes or glaucocystids, are a small group of freshwater unicellular algae, less common today than they were during the Proterozoic. Only 15 species have been described, but more species are likely to e ...
(glaucophyte algae), also have primary chloroplasts that appear to be derived directly from endosymbiotic
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ), in many languages, is one of the values of the grammatical number, grammatical ...

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 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 ') are a major group of s, comprising the (Rhodophyta), the , and the s, and some smaller groups such as the s. All of the lineages of Archaeplastida have become ic, except for the lineage , sister to the Rhodoph ...
, 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 th ...
s,
dinoflagellate The dinoflagellates (Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 m ...
s, and
euglenid Euglenids (euglenoids, or euglenophytes, formally Euglenida/Euglenoida, ICZN, or Euglenophyceae, ICBN) are one of the best-known groups of flagellate 's '' Artforms of Nature'', 1904 (''Giardia lamblia'') ('' Chlamydomonas'') A flagellate is ...
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 In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The group is said to be paraphyleti ...
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 Prasinophyte, 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, i ...
and the
Streptophyta Streptophyta (), informally the streptophytes (, from the Greek language, 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 t ...
(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 Isle of Mull, 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 ...
''. 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 Order or ORDER or Orders may refer to: * Orderliness Orderliness is associated with other qualities such as cleanliness Cleanliness is both the abstract st ...
s and . 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 filamentous charophyte The Charophyta () or charophytes () is a group of freshwater green algae, sometimes treated as a phylum, division, but also as a ...

Spirogyra
'' 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

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 Microbiology (from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is appro ...

microbiology
,
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that stud ...

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 protists 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 Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanis ...
(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 compound , CH4; is among the simplest organic compounds. In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen chemical bo ...
s, fungi do not possess chloroplasts and generally obtain carbon by breaking down and absorbing surrounding materials, and so are called
heterotroph A heterotroph (; from Ancient Greek Ancient Greek includes the forms of the Greek language Greek ( el, label=Modern Greek Modern Greek (, , or , ''Kiní Neoellinikí Glóssa''), generally referred to by speakers simply as Greek ...
ic
saprotrophs Mycelial cord made up of a collection of hyphae; an essential part in the process of saprotrophic nutrition, it is used for the intake of organic matter through its cell wall. The network of hyphae is referred to as a mycelium, which is fundamental ...
. In addition, the substructure of multicellular fungi is different from that of plants, taking the form of many chitinous microscopic strands called
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus A fungus (plural The plural (sometimes abbreviated An abbreviation (from Latin ''brevis'', meaning ''short'') is a shortened form of a word or phrase, by any ...
e, which may be further subdivided into cells or may form a
syncytium A syncytium or symplasm (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") 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 organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interact ...
nuclei ''Nucleus'' (plural nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA ...

nuclei
. Fruiting bodies, of which
mushroom A mushroom or toadstool is the fleshy, spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible befor ...

mushroom
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 and
World Plants In its most general sense, the term "world" refers to the totality of entities, to the whole of reality or to everything that is. The nature of the world has been conceptualized differently in different fields. Some conceptions see the worl ...
both list about 350,000 species. The naming of plants is governed by the
International Code of Nomenclature for algae, fungi, and plants The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical name A botanical name is a formal scientific name Science (from the Latin Latin (, ...
and
International Code of Nomenclature for Cultivated Plants The ''International Code of Nomenclature for Cultivated Plants'' (ICNCP), also known as the Cultivated Plant Code, is a guide to the rules and regulations for naming cultigens, plants whose origin or selection is primarily due to intentional huma ...
(see
cultivated plant taxonomy Cultivated plant taxonomy is the study of the theory A theory is a reason, rational type of abstraction, abstract thinking about a phenomenon, or the results of such thinking. The process of contemplative and rational thinking is often associated ...
).


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 Cyanobacteria , also known as Cyanophyta, are a phylum of Gram-negative bacteria Gram-neg ...
s, through
bryophyte Bryophytes are an informal group consisting of three divisions Division or divider may refer to: Mathematics *Division (mathematics) Division is one of the four basic operations of arithmetic, the ways that numbers are combined to make new n ...

bryophyte
s,
lycopod Lycopodiopsida is a class of herbaceous vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are called clubmosses, firmosses and quillworts. They have dichotomously branching stems bea ...
s,
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia art ...

fern
s to the complex
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also ...
s and
angiosperm Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ...

angiosperm
s 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 A system is a group of Interaction, interacting or interrelated elements that act according to a set of rules to form a unified whole. A system, surrounded and influenced by its environment, ...
, 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 Prof Frederick Orpen Bower FRSE FRS (4 November 1855 – 11 April 1948) was an English botanist Botany, also called , plant biology or phytology, is the science Science (from the Latin word ''scientia'', meaning "knowledge") is ...
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 freshwater unicellular algae, less common today than they were during the Proterozoic. Only 15 species have been described, but more species are likely to exi ...

glaucophyte
s, originated in a terrestrial environment in freshwater biofilms or microbial mats. Primitive land plants began to diversify in the late Silurian Period, around , and the results of their diversification are displayed in remarkable detail in an early
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic The Paleozoic (or Palaeozoic) Era ( ; from the Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the H ...
fossil assemblage from the
Rhynie chert The Rhynie chert is a Lower Devonian Sedimentary rock, 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 ...

Rhynie chert
. 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 A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree (graph theory), tree showing the evolutionary relationships among va ...

phylogenetic tree
of Plantae, after Kenrick and Crane, is as follows, with modification to the Pteridophyta from Smith ''et al.'' The
Prasinophyceae The Prasinophytes (incl. Tetraphytina) or chlorophyta Image:Taiwan 2009 East Coast ShihTiPing Giant Stone Steps Algae FRD 6581.jpg, Green algae on coastal rocks at :zh:石梯坪, Shihtiping in Taiwan Chlorophyta or Prasinophyte, Prasinophyta ...
are a
paraphyletic In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and all descendants of that ancestor excluding a few—typically only one or two—Monophyly, monophyletic subgroups. The group is said to be paraphyleti ...

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 In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular ...

multicellular
land plants, called
embryophyte The Embryophyta (), or land plants, are the most familiar group of green plant Plants are predominantly photosynthetic Photosynthesis is a process used by plants and other organisms to Energy transformation, convert light energy into ...

embryophyte
s. Embryophytes include the
vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia artēria'' 'windpipe' + φυτά ''phutá'' 'plants'), form a large group of plants ( 300,000 ...
s, such as ferns, conifers and flowering plants. They also include the ''
bryophyte Bryophytes are an informal group consisting of three divisions Division or divider may refer to: Mathematics *Division (mathematics) Division is one of the four basic operations of arithmetic, the ways that numbers are combined to make new n ...

bryophyte
s'', of which
moss Mosses are small, non-vascular flower A flower, sometimes known as a bloom or blossom Cherry blossoms in Paris in full bloom. In botany, blossoms are the flowers of stone fruit fruit tree, trees (genus ''Prunus'') and of some other plant ...

moss
es and
liverworts The Marchantiophyta () are a division of non-vascular plant, non-vascular embryophyte, land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plan ...
are the most common. All of these plants have
eukaryotic Eukaryotes () are organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interact ...

eukaryotic
cells with
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane cell membrane vs. Prokaryotes The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to a ...
s composed of
cellulose Cellulose is an organic compound In chemistry Chemistry is the study of the properties and behavior of . It is a that covers the that make up matter to the composed of s, s and s: their composition, structure, properties, behavior ...

cellulose
, and most obtain their energy through
photosynthesis Photosynthesis is a process used by plants and other organisms to convert Conversion or convert may refer to: Arts, entertainment, and media * Conversion (Doctor Who audio), "Conversion" (''Doctor Who'' audio), an episode of the audio drama ' ...

photosynthesis
, using
light Light or visible light is electromagnetic radiation within the portion of the electromagnetic spectrum that is visual perception, perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nan ...

light
, water and
carbon dioxide Carbon dioxide (chemical formula A chemical formula is a way of presenting information about the chemical proportions of s that constitute a particular or molecule, using symbols, numbers, and sometimes also other symbols, such as pare ...

carbon dioxide
to synthesize food. About three hundred plant species do not photosynthesize but are
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is adaptation (biology), adapted structurally to this w ...
s on other species of photosynthetic plants. Embryophytes are distinguished from
green algae The green algae (singular: green alga) are a large, informal grouping of algae consisting of the Chlorophyta and Charophyta/Streptophyta, which are now placed in separate Division (botany), divisions, together with the more basal Mesostigmatoph ...

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 ( ; from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Eu ...
. 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 Ploidy () is the number of complete sets of chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. Most eukaryotic chromosomes include packaging proteins called histones which, aided by ...
stage, called the
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the plant kingdom encompassed all ...
, and a
diploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Sets of chromosomes refer to the number of mate ...
stage, called the
sporophyte 350px, Sporophytes of moss during spring A sporophyte () is the diploid Ploidy () is the number of complete sets of chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. Most eukaryo ...
. In bryophytes, the sporophyte is always unbranched and remains nutritionally dependent on its parent gametophyte. The embryophytes have the ability to secrete a
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non-homology (biology), homologous, differing in the ...
on their outer surface, a waxy layer that confers resistance to desiccation. In the
moss Mosses are small, non-vascular flower A flower, sometimes known as a bloom or blossom Cherry blossoms in Paris in full bloom. In botany, blossoms are the flowers of stone fruit fruit tree, trees (genus ''Prunus'') and of some other plant ...

moss
es and
hornwort Hornworts are a group of bryophyte Bryophytes are an informal group consisting of three divisions Division or divider may refer to: Mathematics *Division (mathematics) Division is one of the four basic operations of arithmetic, the ways th ...
s a cuticle is usually only produced on the sporophyte.
Stomata File:LeafUndersideWithStomata.jpg, The underside of a leaf. In this species (''Tradescantia zebrina'') the guard cells of the stomata are green because they contain chlorophyll while the epidermal cells are chlorophyll-free and contain red pigme ...

Stomata
are absent from liverworts, but occur on the sporangia of mosses and hornworts, allowing gas exchange. Vascular plants first appeared during the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician The Ordovician ( ) is a geologic period A geological period is one of the several subdivisions of geologic time enabling cross-refere ...
period, and by the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic The Paleozoic (or Palaeozoic) Era ( ; from the Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the H ...
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 Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North Ame ...

xylem
and
phloem Phloem (, ) is the living tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North America * ''Triphosa ...

phloem
, 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 is reduced from a multicellular free-living organism to a few cells in a pollen grain and the miniaturised
megagametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plant Plants are mainly multicellular organisms, predominantly photosynthetic Photosynthesis is a process used by plants and other organisms ...
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 plant The spermatophytes, also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plants that produce seeds, hence the alternative name seed plants. They are a subset of the embryophytes or l ...

ovule
. After fertilisation by means of sperm produced by
pollen Pollen is a powdery substance consisting of pollen grains which are microsporophytes of seed plants The spermatophytes (; ), also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plant Plant ...

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 The term Pteridospermatophyta (or "seed ferns" or "Pteridospermatopsida") is a polyphyletic group of extinct seed-bearing plants (spermatophytes). The earliest fossil evidence for plants of this type is the genus ''Elkinsia'' of the late Devonian ...
(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 Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also ...
s, of which four surviving groups are widespread today, particularly the
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single e ...
s, which are dominant
tree In botany, a tree is a perennial plant with an elongated Plant stem, stem, or trunk (botany), trunk, supporting branches and leaves in most species. In some usages, the definition of a tree may be narrower, including only wood plants with se ...

tree
s in several
biome A biome is a collection of plants Plants are predominantly photosynthetic Photosynthesis is a process used by plants and other organisms to Energy transformation, convert light energy into chemical energy that, through cellular respi ...
s. The name gymnosperm comes from the
Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 million as of ...
, 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 Classical Latin is the form of Latin language Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communication used by humans, inc ...

fossil
s include roots, wood, leaves, seeds, fruit,
pollen Pollen is a powdery substance consisting of pollen grains which are microsporophytes of seed plants The spermatophytes (; ), also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plant Plant ...

pollen
,
spore )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible before dispersion of their spores: the calyptra (1) is ...
s,
phytolith Phytoliths (from Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approxim ...
s, and
amber Amber is fossil A fossil (from Classical Latin Classical Latin is the form of Latin language Latin (, or , ) is a classical language A classical language is a language A language is a structured system of communi ...

amber
(the fossilized resin produced by some plants). Fossil land plants are recorded in terrestrial, lacustrine, fluvial and nearshore marine sediments.
Pollen Pollen is a powdery substance consisting of pollen grains which are microsporophytes of seed plants The spermatophytes (; ), also known as phanerogams (taxon Phanerogamae) or phaenogams (taxon Phaenogamae), comprise those plant Plant ...

Pollen
,
spores In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mecha ...
and algae (
dinoflagellate The dinoflagellates (Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its population is approximately 10.7 m ...
s and
acritarch Acritarchs are organic microfossils, known from approximately 1800 million years ago to the present. Their diversity reflects major ecological events such as the appearance of predation and the Cambrian explosion. Definition Acritarchs were ori ...
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 The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period A geological period is one of the several subdivisions of geologic time enabling cross-referencing of rocks and geologic events from place to place. These peri ...
. These fossils resemble
calcified Calcification is the accumulation of calcium salts in a Tissue (biology), 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 origi ...
multicellular A multicellular organism is an organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular ...

multicellular
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 substrate ...
. Earlier
Precambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic The Phanerozoic Eon is the current geologic eon in the geologic time scale The geologi ...

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 The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic The Phanerozoic Eon is the current geologic eon in the geologic time scale The geologi ...

Precambrian
(ca. 1200 mya). The resistant outer walls of
prasinophyte The Prasinophytes (incl. Tetraphytina) or chlorophyta Image:Taiwan 2009 East Coast ShihTiPing Giant Stone Steps Algae FRD 6581.jpg, Green algae on coastal rocks at :zh:石梯坪, Shihtiping in Taiwan Chlorophyta or Prasinophyte, Prasinophyta ...
cysts (known as phycomata) are well preserved in fossil deposits of the
Paleozoic The Paleozoic (or Palaeozoic) Era ( ; from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Eu ...
(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 Molecular ...
, while the oldest reliable records of the
Bryopsidales Bryopsidales is an order of green algae, in the class Ulvophyceae. Characteristics The thallus Thallus (plural: thalli), from Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-Eur ...
,
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 substrate ...
) and are from the
Paleozoic The Paleozoic (or Palaeozoic) Era ( ; from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Eu ...
. The oldest known fossils of embryophytes date from the
Ordovician The Ordovician ( ) is a geologic period The geologic time scale (GTS) is a system of chronological dating that classifies Geology, geological strata (stratigraphy) in time. It is used by geologists, paleontology, paleontologists, and other eart ...

Ordovician
, though such fossils are fragmentary. By the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician The Ordovician ( ) is a geologic period A geological period is one of the several subdivisions of geologic time enabling cross-refere ...
, fossils of whole plants are preserved, including the simple vascular plant ''
Cooksonia ''Cooksonia'' is an extinct group of primitive land plant The Embryophyta (), or land plants, are the most familiar group of green plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Histor ...

Cooksonia
'' 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 Plant Plants are mainly multicellular organisms, predominantly photosynthetic Photosynthesis is a process used by plants and ot ...
''
Baragwanathia longifolia ''Baragwanathia'' is a genus of extinct lycopsid Lycopodiopsida is a class of herbaceous vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are called clubmosses, firmosses and qui ...
'' in late Silurian. From the early Devonian
Rhynie chert The Rhynie chert is a Lower Devonian Sedimentary rock, 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 ...

Rhynie chert
, detailed fossils of lycophytes and
rhyniophyte The rhyniophytes are a group of extinct early vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from the Greek ''trācheia''), form a large group of plants ( 300,000 accepted known sp ...
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 Glomerales is an order of symbiotic fungi within the phylum Glomeromycota. Biology These fungi are all biotrophic mutualists. Most employ the arbuscular mycorrhizal method of nutrient exchange with plants. They produce large (.1-.5mm) spores ( ...
. The
Devonian period The Devonian ( ) is a period (geology), geologic period and system of the Paleozoic, 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, ...
also saw the evolution of leaves and roots, and the first modern tree, ''
Archaeopteris ''Archaeopteris'' is an extinct Extinction is the termination of a kind of organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies the Life ...

Archaeopteris
''. This tree with fern-like foliage and a trunk with conifer-like wood was
heterosporous Heterospory is the production of spores )'', growing on a thinning, thinned hybrid black poplar ''(populus, Populus x canadensis)''. The last stage of the moss#Life cycle, moss lifecycle is shown, where the sporophytes are visible before dispe ...
producing spores of two different sizes, an early step in the evolution of seeds. The
Coal measure The coal measures is a lithostratigraphical term for the coal-bearing part of the Upper Carboniferous The Carboniferous ( ) is a geologic period and system A system is a group of Interaction, interacting or interrelated elements that act acco ...

Coal measure
s are a major source of
Paleozoic The Paleozoic (or Palaeozoic) Era ( ; from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Eu ...
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 , Germany ) , image_map = , map_caption = , map_width = 250px , capital = Berlin , coordinates = , largest_city = capital , languages_type = Official language , languages = German language, German , ...

coal
itself is the remains of fossilised plants, though structural detail of the plant fossils is rarely visible in coal. In the
Fossil Grove The Fossil Grove is a group of fossils located within Victoria Park, Glasgow Glasgow, (, also , ; sco, Glesca or ; gd, Glaschu ) with an estimated city A city is a large human settlement.Goodall, B. (1987) ''The Penguin Dictionary of ...
at Victoria Park in
Glasgow Glasgow ( ; sco, Glesga; gd, Glaschu) is the most populous city A city is a large .Goodall, B. (1987) ''The Penguin Dictionary of Human Geography''. London: Penguin.Kuper, A. and Kuper, J., eds (1996) ''The Social Science Encyclopedia'' ...

Glasgow
, Scotland, the stumps of ''
Lepidodendron ''Lepidodendron'' is an extinct Extinction is the termination of a kind of organism In biology, an organism (from Ancient Greek, Greek: ὀργανισμός, ''organismos'') is any individual contiguous system that embodies the Life# ...

Lepidodendron
'' trees are found in their original growth positions. The fossilized remains of
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single e ...
and
angiosperm Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek#REDIRECT Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ...

angiosperm
root In vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia artēria'' 'windpipe' + φυτά ''phutá'' 'plants'), form a large grou ...

root
s, and
branch A branch ( or , ) or tree branch (sometimes referred to in 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 spe ...

branch
es may be locally abundant in lake and inshore
sedimentary rock Sedimentary rocks are types of rock Rock most often refers to: * Rock (geology) A rock is any naturally occurring solid mass or aggregate of minerals or mineraloid matter. It is categorized by the minerals included, its Chemical compoun ...

sedimentary rock
s from the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles and the Age of Conifers, is the second-to-last era An era is a span of time defined for the purposes of chronology or historiography, as in the regnal eras in the history of a given monarchy ...
and
Cenozoic The Cenozoic ( ; ) is Earth's current geological era An era is a span of time defined for the purposes of chronology or historiography, as in the regnal eras in the history of a given monarchy, a calendar era used for a given calendar, or the ge ...

Cenozoic
eras. and its allies,
magnolia ''Magnolia'' is a large genus Genus /ˈdʒiː.nəs/ (plural genera /ˈdʒen.ər.ə/) is a taxonomic rank In biological classification In biology, taxonomy () is the scientific study of naming, defining (Circumscription (taxonomy), ...

magnolia
,
oak An oak is a tree In botany, a tree is a perennial plant with an elongated Plant stem, stem, or trunk (botany), trunk, supporting branches and leaves in most species. In some usages, the definition of a tree may be narrower, including on ...

oak
, and are often found.
Petrified wood File:PetrifiedWood.jpg, Petrified log at the Petrified Forest National Park Petrified wood (from Ancient Greek meaning 'rock' or 'stone'; literally 'wood turned into stone') is the name given to a special type of fossilized remains of terrestr ...

Petrified wood
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 In earth science Earth science or geoscience includes all fields of natural science Natural science is a branch of science Science (from the Latin word ''scientia'', meaning "knowledge") is a systematic enterprise that Scientific ...

erosion
. Petrified wood is often heavily silicified (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 In classical physics and general chem ...
replaced by
silicon dioxide Silicon dioxide, also known as silica, is an oxide of rutile. Ti(IV) centers are grey; oxygen centers are red. Notice that oxygen forms three bonds to titanium and titanium forms six bonds to oxygen. An oxide () is a chemical compound that con ...
), 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 Rock (geology), stone, minerals, or gemstones into decorative items such as cabochons, engraved gems (including cameo (carving), cameos), and faceted designs. A person who practices lapidary ...
equipment. Fossil forests of petrified wood have been found in all continents. Fossils of seed ferns such as ''
Glossopteris ''Glossopteris'' ( grc, γλώσσα , meaning "tongue", because the leaves were tongue-shaped, and ''pteris'', Greek for fern or feathery) is the largest and best-known genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is th ...

Glossopteris
'' are widely distributed throughout several continents of the
Southern Hemisphere The Southern Hemisphere is the half (hemisphere Hemisphere may refer to: * A half of a sphere As half of the Earth * A hemispheres of Earth, hemisphere of Earth ** Northern Hemisphere ** Southern Hemisphere ** Eastern Hemisphere ** Western He ...

Southern Hemisphere
, a fact that gave support to
Alfred Wegener Alfred Lothar Wegener (; ; 1 November 1880 – November 1930) was a German polar researcher, geophysicist and meteorologist. During his lifetime he was primarily known for his achievements in meteorology and as a pioneer of polar researc ...

Alfred Wegener
's early ideas regarding
Continental drift Continental drift is the hypothesis that the Earth's continent A continent is one of several large landmasses. Generally identified by convention (norm), convention rather than any strict criteria, up to seven regions are commonly reg ...
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 Conversion or convert may refer to: Arts, entertainment, and media * Conversion (Doctor Who audio), "Conversion" (''Doctor Who'' audio), an episode of the audio drama ' ...

photosynthesis
, most plants use the energy in
sunlight Sunlight is a portion of the given off by the , in particular , , and light. On , sunlight is and through , and is obvious as when the Sun is above the . When direct is not blocked by s, it is experienced as sunshine, a combination of b ...

sunlight
to convert
carbon dioxide Carbon dioxide (chemical formula A chemical formula is a way of presenting information about the chemical proportions of s that constitute a particular or molecule, using symbols, numbers, and sometimes also other symbols, such as pare ...

carbon dioxide
from the atmosphere, plus
water Water (chemical formula H2O) 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 li ...

water
, into simple
sugar Sugar is the generic name for sweet-tasting, soluble carbohydrate is a disaccharide A disaccharide (also called a double sugar or ''biose'') is the sugar formed when two monosaccharides are joined by glycosidic linkage. Like monosacc ...

sugar
s. These sugars are then used as building blocks and form the main structural component of the plant.
Chlorophyll Chlorophyll (also chlorophyl) is any of several related green 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 ...

Chlorophyll
, a green-colored,
magnesium Magnesium is a chemical element upright=1.0, 500px, The chemical elements ordered by link=Periodic table In chemistry Chemistry is the science, scientific study of the properties and behavior of matter. It is a natural science ...

magnesium
-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 compound , CH4; is among the simplest organic compou ...
is essential to this process; it is generally present in plant
leaves A leaf (plural leaves) is the principal lateral appendage of the vascular plant plant stem, stem, usually borne above ground and specialized for photosynthesis. The leaves, stem, flower and fruit together form the shoot system. Leaves are ...

leaves
, and often in other plant parts as well.
Parasitic plant A parasitic plant is a plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; how ...

Parasitic plant
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
compounds Compound may refer to: Architecture and built environments * Compound (enclosure), a cluster of buildings having a shared purpose, usually inside a fence or wall ** Compound (fortification), a version of the above fortified with defensive structu ...
of
nitrogen Nitrogen is the chemical element upright=1.0, 500px, The chemical elements ordered by link=Periodic table In chemistry Chemistry is the science, scientific study of the properties and behavior of matter. It is a natural science ...

nitrogen
,
phosphorus Phosphorus is a chemical element In chemistry, an element is a pure Chemical substance, substance consisting only of atoms that all have the same numbers of protons in their atomic nucleus, nuclei. Unlike chemical compounds, chemical el ...

phosphorus
,
potassium Potassium is a chemical element In chemistry Chemistry is the study of the properties and behavior of . It is a that covers the that make up matter to the composed of s, s and s: their composition, structure, properties, b ...

potassium
, magnesium and other elemental
nutrient A nutrient is a substance Substance may refer to: * Substance (Jainism), a term in Jain ontology to denote the base or owner of attributes * Chemical substance, a material with a definite chemical composition * Matter, anything that has mass and t ...
s from the soil.
Epiphytic 200px, '' Tillandsia bourgaei'' growing on an oak tree in Mexico An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating a ...

Epiphytic
and
lithophytic Lithophytes are plants that grow in or on Rock (geology), rocks. They can be classified as either epilithic (or epipetric) or endolithic, with the former being plants that grow on the surfaces of rocks. Endolithic lithophytes grow in the crevices o ...
plants depend on air and nearby debris for nutrients, and
carnivorous plant Carnivorous plants are plants that derive some or most of their Plant nutrition, nutrients from trapping and consuming animals or protozoans, typically insects and other arthropods. However, carnivorous plants generate energy from photosynthesis. ...

carnivorous plant
s 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 Oxygen is the chemical element Image:Simple Periodic Table Chart-blocks.svg, 400px, Periodic table, The periodic table of the chemical elements In chemistry, an element is a pure substance consisting only of atoms that all have the same ...

oxygen
in the atmosphere and around their roots (
soil gas Soil gases are the gases found in the air space between soil Surface-water- gley developed in glacial till, Northern Ireland.">Northern_Ireland.html" ;"title="glacial till, Northern Ireland">glacial till, Northern Ireland. Soil is a mixture o ...
) for
respiration Respiration may refer to: Biology * Cellular respiration, the process in which nutrients are converted into useful energy in a cell ** Anaerobic respiration, cellular respiration without oxygen ** Maintenance respiration, the amount of cellular ...

respiration
. Plants use oxygen and
glucose Glucose is a simple with the . Glucose is the most abundant , a subcategory of s. Glucose is mainly made by and most during from water and carbon dioxide, using energy from sunlight, where it is used to make in s, the most abundant carbohydr ...

glucose
(which may be produced from stored
starch Starch or amylum is a polymeric A polymer (; Greek '' poly-'', "many" + '' -mer'', "part") is a substance Substance may refer to: * Substance (Jainism), a term in Jain ontology to denote the base or owner of attributes * Chemical substance ...
) to provide energy. Some plants grow as submerged aquatics, using oxygen dissolved in the surrounding water, and a few specialized vascular plants, such as
mangrove A mangrove is a shrub A shrub (often called a bush) is a small- to medium-sized perennial A perennial plant or simply perennial is a plant Plants are predominantly photosynthetic eukaryotes of the Kingdom (biology), kingdom Pl ...

mangrove
s and reed (''
Phragmites australis ''Phragmites australis'', known as common reed, is a broadly distributed wetland grass growing nearly tall. Taxonomy Recent studies have characterized morphological distinctions between the introduced and native stands of ''Phragmites australi ...

Phragmites australis
''), can grow with their roots in
anoxic The term anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of disso ...
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 Temperature ( ) is a physical quantity that expresses hot and cold. It is the manifestation of thermal energy Thermal radiation in visible light can be seen on this hot metalwork. Thermal energy refers to several distinct physical concept ...

temperature
, available
water Water (chemical formula H2O) 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 li ...

water
, available
light Light or visible light is electromagnetic radiation within the portion of the electromagnetic spectrum that is visual perception, perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nan ...

light
,
carbon dioxide Carbon dioxide (chemical formula A chemical formula is a way of presenting information about the chemical proportions of s that constitute a particular or molecule, using symbols, numbers, and sometimes also other symbols, such as pare ...

carbon dioxide
and available
nutrient A nutrient is a substance Substance may refer to: * Substance (Jainism), a term in Jain ontology to denote the base or owner of attributes * Chemical substance, a material with a definite chemical composition * Matter, anything that has mass and t ...
s 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 Etiolation is a process in flowering plant Flowering plants include multiple members of the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek Greek may refer to: Greece Anything of, from, ...
and
chlorosis 300px, A corn plant with severe chlorosis (left) beside a normal plant (right) In botany Botany, also called , plant biology or phytology, is the science Science (from the Latin word ''scientia'', meaning "knowledge") is a systematic e ...

chlorosis
. Optimal plant growth can be hampered by grazing animals, suboptimal soil composition, lack of
mycorrhiza A mycorrhiza (from Ancient Greek, Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a mutual symbiosis, symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role ...

mycorrhiza
l fungi, and attacks by insects or
plant diseases Plant pathology (also phytopathology) is the scientific study of disease A disease is a particular abnormal condition that negatively affects the structure A structure is an arrangement and organization of interrelated elements in ...
, 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 plant An annual plant is a plant that completes its life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia *Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending w ...
s grow and reproduce within one
growing season A season is a division of the year marked by changes in weather, ecology, and the amount of daylight. The growing season is that portion of the year in which local conditions (i.e. rainfall, temperature, daylight) permit normal plant morphology#Grow ...
,
biennial plant 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 ...
s grow for two growing seasons and usually reproduce in second year, and
perennial plant A perennial plant or simply perennial is a plant that lives more than two years. The term (''wikt:per-#Prefix, per-'' + ''wikt:-ennial#Suffix, -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annual p ...
s 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 Alpine may refer to: Places * Alps, a European mountain range * Alpine states, associated with the mountain range, or relating to any lofty mountain areas * Mountainous or alpine; the mountains. Australia * Alpine, New South Wales, a Northern Vill ...
or
temperate In geography Geography (from Greek Greek may refer to: Greece Anything of, from, or related to Greece Greece ( el, Ελλάδα, , ), officially the Hellenic Republic, is a country located in Southeast Europe. Its populati ...
regions can be biennial or perennial in warmer climates. Among the vascular plants, perennials include both
evergreen In 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 Anc ...

evergreen
s that keep their leaves the entire year, and
deciduous In the fields of horticulture Horticulture is the art of cultivating plants in gardens to produce food and medicinal ingredients, or for comfort and ornamental purposes. Horticulturists are agriculturists who grow flowers, fruits and nuts, ...
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 The tropics are the region of Earth surrounding the Equator. They are delimited in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S; these latitudes correspond to ...

tropical
plants lose their leaves during the
dry season The dry season is a yearly period of low rainfall, especially in the tropics The tropics are the region of Earth Earth is the third planet from the Sun and the only astronomical object known to harbour and support life. 29.2% of ...
. 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 Kudzu (also called Japanese arrowroot or Chinese arrowroot) is a group of climbing, coiling, and trailing perennial vines native to much of East Asia East Asia is the eastern region of Asia Asia () is Earth's largest and most po ...

kudzu
, which do not need to produce thick supportive tissue, may grow up to 12.5 mm/h. Plants protect themselves from
frost Frost is a thin layer of ice Ice is water Water (chemical formula H2O) is an , transparent, tasteless, odorless, and , which is the main constituent of 's and the s of all known living organisms (in which it acts as a ). It is ...

frost
and
dehydration In physiology, dehydration is a lack of total body water In physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology Biology is the natural science that studie ...

dehydration
stress with
antifreeze protein Antifreeze proteins (AFPs) or ice structuring proteins (ISPs) refer to a class of polypeptides produced by certain animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological ...
s, heat-shock proteins and sugars (
sucrose Sucrose is a type of sugar Sugar is the generic name for , soluble s, many of which are used in food. Simple sugars, also called s, include , , and . Compound sugars, also called s or double sugars, are molecules made of two monosacchari ...

sucrose
is common). LEA ( Late Embryogenesis Abundant) protein expression is induced by stresses and protects other proteins from aggregation as a result of
desiccation Desiccation (from Latin de- "thoroughly" + siccare "to dry") is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its lo ...
and
freezing Freezing is a phase transition where a liquid turns into a solid when its temperature is lowered below its freezing point. In accordance with the internationally established definition, freezing means the solidification phase change of a liquid o ...

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 cellLyons, 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 DNA repair is a collection of processes by which a identifies and corrects damage to the molecules that encode its . In human cells, both normal activities and environmental factors such as can cause DNA damage, resulting in tens of thousan ...
directly, or indirectly via the generation of
reactive oxygen species Reactive oxygen species (ROS) are highly Reactivity (chemistry), reactive chemicals formed from O2. Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen. The reduction of molecular oxygen (O2) produce ...
. 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 A seed is an embryonic ''Embryonic'' is the twelfth studio album by experimental rock band the Flaming Lips released on October 13, 2009, on Warner Bros. Records, Warner Bros. The band's first double album, it was released to generally positi ...

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- and double-strand breaks in DNA can be DNA repair, repaired. 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,
chloroplast A chloroplast is a type of membrane-bound organelle In cell biology, an organelle is a specialized subunit, usually within a cell (biology), cell, that has a specific function. The name ''organelle'' comes from the idea that these structure ...

chloroplast
s, and rigid
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane cell membrane vs. Prokaryotes The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to a ...
s that are made up of
cellulose Cellulose is an organic compound In chemistry Chemistry is the study of the properties and behavior of . It is a that covers the that make up matter to the composed of s, s and s: their composition, structure, properties, behavior ...

cellulose
, 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 within the portion of the electromagnetic spectrum that is visual perception, perceived by the human eye. Visible light is usually defined as having wavelengths in the range of 400–700 nan ...

light
. 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 compound , CH4; is among the simplest organic compou ...
chlorophyll Chlorophyll (also chlorophyl) is any of several related green 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 ...

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 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 ''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 Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North Ame ...

xylem
and
phloem Phloem (, ) is the living tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in North America * ''Triphosa ...

phloem
. They also have
root In vascular plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia artēria'' 'windpipe' + φυτά ''phutá'' 'plants'), form a large grou ...

root
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 Oxygen is the chemical element Image:Simple Periodic Table Chart-blocks.svg, 400px, Periodic table, The periodic table of the chemical elements In chemistry, an element is a pure substance consisting only of atoms that all have the same ...

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 collection of plants Plants are predominantly photosynthetic Photosynthesis is a process used by plants and other organisms to Energy transformation, convert light energy into chemical energy that, through cellular respi ...
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 collection of plants Plants are predominantly photosynthetic Photosynthesis is a process used by plants and other organisms to Energy transformation, convert light energy into chemical energy that, through cellular respi ...
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 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 ...

fruit
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 A mycorrhiza (from Ancient Greek, Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a mutual symbiosis, symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role ...

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 Chlorophyll (also chlorophyl) is any of several related green 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 ...

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 plant Vascular plants (from Latin ''vasculum'': duct), also known as Tracheophyta (the tracheophytes , from Greek τραχεῖα ἀρτηρία ''trācheia art ...

fern
s and
moss Mosses are small, non-vascular flower A flower, sometimes known as a bloom or blossom Cherry blossoms in Paris in full bloom. In botany, blossoms are the flowers of stone fruit fruit tree, trees (genus ''Prunus'') and of some other plant ...

moss
es 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 Nitrogen is the chemical element upright=1.0, 500px, The chemical elements ordered by link=Periodic table In chemistry Chemistry is the science, scientific study of the properties and behavior of matter. It is a natural science ...

nitrogen
and
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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
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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 A flower, som ...

vegetable
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,
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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
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, 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
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cellulose
. 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 Algarum



Plant Resources of Tropical Africa

Tree of Life
;Botanical and vegetation databases
African Plants Initiative database

Australia

Chilean plants at ''Chilebosque''

e-Floras (Flora of China, Flora of North America and others)



Flora of Central Europe

Flora of North America



Meet the Plants-National Tropical Botanical Garden

Lady Bird Johnson Wildflower Center – Native Plant Information Network at University of Texas, Austin

The Plant List

United States Department of Agriculture
not limited to continental US species {{Authority control Plants, Kingdoms (biology), Plants