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Plant

Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude the fungi and some algae, as well as the prokaryotes (the archaea and bacteria). By one definition, plants form the clade Viridiplantae (Latin name for "green plants") which is sister of the Glaucophyta, and consists of the green algae and Embryophyta (land plants). The latter includes the flowering plants, conifers and other gymnosperms, ferns and their allies, hornworts, liverworts, and mosses.

Plants
Temporal range:
Mesoproterozoic–present
Scientific classification
Domain: Eukaryota
(unranked): Diaphoretickes
(unranked): Archaeplastida
Kingdom: Plantae
sensu Copeland, 1956
Superdivisions
Synonyms
  • Viridiplantae Cavalier-Smith 1981[1]
  • Chlorobionta Jeffrey 1982, emend. Bremer 1985, emend. Lewis and McCourt 2004[2]
  • Chlorobiota Kenrick and Crane 1997[3]
  • Chloroplastida Adl et al., 2005 [4]
  • Phyta Barkley 1939 emend. Holt & Uidica 2007
  • Cormophyta Endlicher, 1836
  • Cormobionta Rothmaler, 1948
  • Euplanta Barkley, 1949
  • Telomobionta Takhtajan, 1964
  • Embryobionta Cronquist et al., 1966
  • Metaphyta Whittaker, 1969

Most plants are multicellular organisms. Green plants obtain most of their energy from sunlight via photosynthesis by primary chloroplasts that are derived from endosymbiosis with cyanobacteria. Their chloroplasts contain chlorophylls a and b, which gives them their green color. Some plants are 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 and alternation of generations, although asexual reproduction is also common.

There are about 320,000 known species of plants, of which the great majority, some 260,000–290,000, produce seeds.[5] Green plants provide a substantial proportion of the world's molecular oxygen,[6] and are the basis of most of Earth's ecosystems. Plants that produce grain, fruit, and vegetables also form basic human foods and have been domesticated for millennia. Plants have many cultural and other uses, as ornaments, building materials, writing material and, in great variety, they have been the source of medicines and psychoactive drugs. The scientific study of plants is known as botany, a branch of biology.

Definition

All living things were traditionally placed into one of two groups, plants and animals. This classification may date from Aristotle (384–322 BCE), 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 (1707–1778) created the basis of the modern system of scientific classification, these two groups became the kingdoms Vegetabilia (later Metaphyta or Plantae) and Animalia (also called Metazoa). Since then, it has become clear that the plant kingdom as originally defined included several unrelated groups, and the fungi and several groups of algae were removed to new kingdoms. However, these organisms are still sometimes considered plants, particularly in informal contexts.[citation needed]

The term "plant" generally implies the possession of the following traits: multicellularity, possession of cell walls containing cellulose, and the ability to carry out photosynthesis with primary chloroplasts.[7][8]

Current definitions of Plantae

When the name Plantae or plant is applied to a specific group of organisms or taxon, it usually refers to one of four concepts. From least to most inclusive, these four groupings are:

Name(s) Scope Description
Land plants, also known as Embryophyta Plantae sensu strictissimo Plants in the strictest sense include the liverworts, hornworts, mosses, and vascular plants, as well as fossil plants similar to these surviving groups (e.g., Metaphyta Whittaker, 1969,[9] Plantae Margulis, 1971[10]).
Green plants, also known as Viridiplantae, Viridiphyta, Chlorobionta or Chloroplastida Plantae sensu stricto Plants in a strict sense include the green algae, and land plants that emerged within them, including stoneworts. The relationships between plant groups are still being worked out, and the names given to them vary considerably. The clade Viridiplantae encompasses a group of organisms that have cellulose in their cell walls, possess chlorophylls a and b and have plastids bound by only two membranes that are capable of photosynthesis and of storing starch. This clade is the main subject of this article (e.g., Plantae Copeland, 1956[11]).
Archaeplastida, also known as Plastida or Primoplantae Plantae sensu lato Plants in a broad sense comprise the green plants listed above plus the red algae (Rhodophyta) and the glaucophyte algae (Glaucophyta) that store Floridean starch outside the plastids, in the cytoplasm. This clade includes all of the organisms that eons ago acquired their primary chloroplasts directly by engulfing cyanobacteria (e.g., Plantae Cavalier-Smith, 1981[12]).
Old definitions of plant (obsolete) Plantae sensu amplo Plants in the widest sense refers to older, obsolete classifications that placed diverse algae, fungi or bacteria in Plantae (e.g., Plantae or Vegetabilia Linnaeus,[13] Plantae Haeckel 1866,[14] Metaphyta Haeckel, 1894,[15] Plantae Whittaker, 1969[9]).

Another way of looking at the relationships between the different groups that have been called "plants" is through a cladogram, which shows their evolutionary relationships. These are not yet completely settled, but one accepted relationship between the three groups described above is shown below[clarification needed].[16][17][18][19][20][21][22][23][24][excessive citations] Those which have been called "plants" are in bold (some minor groups have been omitted).

Archaeplastida + cryptista 

Rhodophyta (red algae)

Rhodelphidia (predatorial)

Picozoa

Glaucophyta (glaucophyte algae)

green plants

Cryptista

groups traditionally
called green algae

The way in which the groups of green algae are combined and named varies considerably between authors.

Algae

Algae consist of several groups of organisms which produce food by photosynthesis and thus have traditionally been included in the plant kingdom. The seaweeds range from large multicellular algae to single-celled organisms and are classified into three groups, the green algae, red algae and brown algae. 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.[25][26]

The Viridiplantae, the green plants – green algae and land plants – form a clade, a group consisting of all the descendants of a common ancestor. With a few exceptions, the green plants have the following features in common; primary chloroplasts derived from cyanobacteria containing chlorophylls a and b, cell walls containing cellulose, and food stores in the form of starch contained within the plastids. They undergo closed mitosis without centrioles, and typically have mitochondria with flat cristae. The chloroplasts of green plants are surrounded by two membranes, suggesting they originated directly from endosymbiotic cyanobacteria.

Two additional groups, the Rhodophyta (red algae) and Glaucophyta (glaucophyte algae), also have primary chloroplasts that appear to be derived directly from endosymbiotic cyanobacteria, although they differ from Viridiplantae in the pigments which are used in photosynthesis and so are different in colour. These groups also differ from green plants in that the storage polysaccharide is floridean starch 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, 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, haptophytes, dinoflagellates, and euglenids) not only have different pigments but also have chloroplasts with three or four surrounding membranes. They are not close relatives of the Archaeplastida, presumably having acquired chloroplasts separately from ingested or symbiotic green and red algae. They are thus not included in even the broadest modern definition of the plant kingdom, although they were in the past.

The green plants or Viridiplantae were traditionally divided into the green algae (including the stoneworts) and the land plants. However, it is now known that the land plants evolved from within a group of green algae, so that the green algae by themselves are a paraphyletic group, that is, 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 and the Streptophyta (including the land plants and Charophyta).[27][28]

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,[29] mainly unicellular or multicellular marine organisms such as the sea lettuce, Ulva.

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 desmids and stoneworts. Streptophyte algae are either unicellular or form multicellular filaments, branched or unbranched.[28] The genus 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.[30] Growing immersed in fresh water, they consist of a central stalk with whorls of branchlets.

Fungi

 
A variety of fungi species

Linnaeus' original classification placed the fungi within the Plantae, since they were unquestionably neither animals or minerals and these were the only other alternatives. With 19th century developments in microbiology, Ernst Haeckel introduced the new kingdom Protista in addition to Plantae and Animalia, but whether fungi were best placed in the Plantae or should be reclassified as 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 (concestor), of the Fungi was probably more similar to that of the Animalia than to that of Plantae or any other kingdom.[31]

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 autotrophs, fungi do not possess chloroplasts and generally obtain carbon by breaking down and absorbing surrounding materials, and so are called heterotrophic saprotrophs. In addition, the substructure of multicellular fungi is different from that of plants, taking the form of many chitinous microscopic strands called hyphae, which may be further subdivided into cells or may form a syncytium containing many eukaryotic nuclei. Fruiting bodies, of which mushrooms are the most familiar example, are the reproductive structures of fungi, and are unlike any structures produced by plants.[citation needed]

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 both list about 391,000 species.[32][33][34]

Diversity of living green plant (Viridiplantae) divisions
Informal group Division name
[citation needed]
Common name No. of living species Approximate no. in informal group
Green algae Chlorophyta Green algae (chlorophytes) 3,800–4,300 [35][36] 8,500

(6,600–10,300)

Charophyta Green algae (e.g. desmids & stoneworts) 2,800–6,000 [37][38]
Bryophytes Marchantiophyta Liverworts 6,000–8,000 [39] 19,000

(18,100–20,200)

Anthocerotophyta Hornworts 100–200 [40]
Bryophyta Mosses 12,000 [41]
Pteridophytes Lycopodiophyta Clubmosses 1,200 [26] 12,000

(12,200)

Polypodiophyta Ferns, whisk ferns & horsetails 11,000 [26]
spermatophyte Cycadophyta Cycads 160 [42] 260,000

(259,511)

Ginkgophyta Ginkgo 1 [43]
Pinophyta Conifers 630 [26]
Gnetophyta Gnetophytes 70 [26]
Magnoliophyta Flowering plants 258,650 [44]

The naming of plants is governed by the International Code of Nomenclature for algae, fungi, and plants and International Code of Nomenclature for Cultivated Plants (see cultivated plant taxonomy).

Evolution

The evolution of plants has resulted in increasing levels of complexity, from the earliest algal mats, through bryophytes, lycopods, ferns to the complex gymnosperms and angiosperms of today. Plants in all of these groups continue to thrive, especially in the environments in which they evolved.

An algal scum formed on the land 1,200 million years ago, but it was not until the Ordovician Period, around 450 million years ago, that land plants appeared.[45] 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.[46] 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 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,[47] and that even the common ancestor of red and green algae, and the unicellular freshwater algae glaucophytes, originated in a terrestrial environment in freshwater biofilms or microbial mats.[48] Primitive land plants began to diversify in the late Silurian Period, around 420 million years ago, and the results of their diversification are displayed in remarkable detail in an early Devonian fossil assemblage from the 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.[49] 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 (~200 million years ago), 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 40 million years ago. The grasses, as well as many other groups, evolved new mechanisms of metabolism to survive the low CO2 and warm, dry conditions of the tropics over the last 10 million years.

A 1997 proposed phylogenetic tree of Plantae, after Kenrick and Crane,[50] is as follows, with modification to the Pteridophyta from Smith et al.[51] The Prasinophyceae are a paraphyletic assemblage of early diverging green algal lineages, but are treated as a group outside the Chlorophyta:[52] later authors have not followed this suggestion.

Prasinophyceae (micromonads)

Streptobionta
Embryophytes
Stomatophytes
Polysporangiates
Tracheophytes
Eutracheophytes
Euphyllophytina
Lignophyta

Spermatophytes (seed plants)

Progymnospermophyta †

Pteridophyta

Pteridopsida (true ferns)

Marattiopsida

Equisetopsida (horsetails)

Psilotopsida (whisk ferns & adders'-tongues)

Cladoxylopsida †

Lycophytina

Rhyniophyta †

Aglaophyton †

Horneophytopsida †

Bryophyta (mosses)

Anthocerotophyta (hornworts)

Marchantiophyta (liverworts)

Charophyta

A newer proposed classification follows Leliaert et al. 2011[53] and modified with Silar 2016[20][21][54][55] for the green algae clades and Novíkov & Barabaš-Krasni 2015[56] 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.[57] The placing of algal groups is supported by phylogenies based on genomes from the Mesostigmatophyceae and Chlorokybophyceae that have since been sequenced.[58][59] The classification of Bryophyta is supported both by Puttick et al. 2018,[60] and by phylogenies involving the hornwort genomes that have also since been sequenced.[61][62]

Embryophytes

The plants that are likely most familiar to us are the multicellular land plants, called embryophytes. Embryophytes include the vascular plants, such as ferns, conifers and flowering plants. They also include the bryophytes, of which mosses and liverworts are the most common.

All of these plants have eukaryotic cells with cell walls composed of cellulose, and most obtain their energy through photosynthesis, using light, water and carbon dioxide to synthesize food. About three hundred plant species do not photosynthesize but are parasites on other species of photosynthetic plants. Embryophytes are distinguished from green algae, which represent a mode of photosynthetic life similar to the kind modern plants are believed to have evolved from, by having specialized reproductive organs protected by non-reproductive tissues.

Bryophytes first appeared during the early Paleozoic. They mainly live in habitats where moisture is available for significant periods, although some species, such as Targionia, are desiccation-tolerant. Most species of bryophytes remain small throughout their life-cycle. This involves an alternation between two generations: a haploid stage, called the gametophyte, and a diploid stage, called the sporophyte. In bryophytes, the sporophyte is always unbranched and remains nutritionally dependent on its parent gametophyte. The embryophytes have the ability to secrete a cuticle on their outer surface, a waxy layer that confers resistance to desiccation. In the mosses and hornworts a cuticle is usually only produced on the sporophyte. Stomata are absent from liverworts, but occur on the sporangia of mosses and hornworts, allowing gas exchange.

Vascular plants first appeared during the Silurian period, and by the Devonian had diversified and spread into many different terrestrial environments. They developed a number of adaptations that allowed them to spread into increasingly more arid places, notably the vascular tissues xylem and 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 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. After fertilisation by means of sperm produced by pollen grains, an embryo sporophyte develops inside the ovule. The integument becomes a seed coat, and the ovule develops into a seed. Seed plants can survive and reproduce in extremely arid conditions, because they are not dependent on free water for the movement of sperm, or the development of free living gametophytes.

The first seed plants, pteridosperms (seed ferns), now extinct, appeared in the Devonian and diversified through the Carboniferous. They were the ancestors of modern gymnosperms, of which four surviving groups are widespread today, particularly the conifers, which are dominant trees in several biomes. The name gymnosperm comes from the Greek γυμνόσπερμος, a composite of γυμνός (gymnos lit.'naked') and σπέρμα (sperma lit.'seed'), 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

 
A petrified log in Petrified Forest National Park, Arizona

Plant fossils include roots, wood, leaves, seeds, fruit, pollen, spores, phytoliths, and amber (the fossilized resin produced by some plants). Fossil land plants are recorded in terrestrial, lacustrine, fluvial and nearshore marine sediments. Pollen, spores and algae (dinoflagellates and acritarchs) are used for dating sedimentary rock sequences. The remains of fossil plants are not as common as fossil animals, although plant fossils are locally abundant in many regions worldwide.

The earliest fossils clearly assignable to Kingdom Plantae are fossil green algae from the Cambrian. These fossils resemble calcified multicellular members of the Dasycladales. Earlier Precambrian fossils are known that resemble single-cell green algae, but definitive identity with that group of algae is uncertain.

The earliest fossils attributed to green algae date from the Precambrian (ca. 1200 mya).[63][64] The resistant outer walls of prasinophyte cysts (known as phycomata) are well preserved in fossil deposits of the Paleozoic (ca. 250–540 mya). A filamentous fossil (Proterocladus) from middle Neoproterozoic deposits (ca. 750 mya) has been attributed to the Cladophorales, while the oldest reliable records of the Bryopsidales, Dasycladales) and stoneworts are from the Paleozoic.[52][65]

The oldest known fossils of embryophytes date from the Ordovician, though such fossils are fragmentary. By the Silurian, fossils of whole plants are preserved, including the simple vascular plant Cooksonia in mid-Silurian and the much larger and more complex lycophyte Baragwanathia longifolia in late Silurian. From the early Devonian Rhynie chert, detailed fossils of lycophytes and rhyniophytes have been found that show details of the individual cells within the plant organs and the symbiotic association of these plants with fungi of the order Glomales. The Devonian period also saw the evolution of leaves and roots, and the first modern tree, Archaeopteris. This tree with fern-like foliage and a trunk with conifer-like wood was heterosporous producing spores of two different sizes, an early step in the evolution of seeds.[66]

The Coal measures are a major source of Paleozoic 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 itself is the remains of fossilised plants, though structural detail of the plant fossils is rarely visible in coal. In the Fossil Grove at Victoria Park in Glasgow, Scotland, the stumps of Lepidodendron trees are found in their original growth positions.

The fossilized remains of conifer and angiosperm roots, stems and branches may be locally abundant in lake and inshore sedimentary rocks from the Mesozoic and Cenozoic eras. Sequoia and its allies, magnolia, oak, and palms are often found.

 
Range of pangaea glossopteris

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. Petrified wood is often heavily silicified (the organic material replaced by silicon dioxide), and the impregnated tissue is often preserved in fine detail. Such specimens may be cut and polished using lapidary equipment. Fossil forests of petrified wood have been found in all continents.

Fossils of seed ferns such as Glossopteris are widely distributed throughout several continents of the Southern Hemisphere, a fact that gave support to Alfred Wegener's early ideas regarding Continental drift theory.

Structure, growth, and development

 
The leaf is usually the primary site of photosynthesis in plants.

Most of the solid material in a plant is taken from the atmosphere. Through the process of photosynthesis, most plants use the energy in sunlight to convert carbon dioxide from the atmosphere, plus water, into simple sugars. These sugars are then used as building blocks and form the main structural component of the plant. Chlorophyll, a green-colored, magnesium-containing pigment is essential to this process; it is generally present in plant leaves, and often in other plant parts as well. Parasitic plants, 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 of nitrogen, phosphorus, potassium, magnesium and other elemental nutrients from the soil. Epiphytic and lithophytic plants depend on air and nearby debris for nutrients, and carnivorous plants supplement their nutrient requirements, particularly for nitrogen and phosphorus, with insect prey that they capture. For the majority of plants to grow successfully they also require oxygen in the atmosphere and around their roots (soil gas) for respiration. Plants use oxygen and glucose (which may be produced from stored starch) to provide energy.[67] Some plants grow as submerged aquatics, using oxygen dissolved in the surrounding water, and a few specialized vascular plants, such as mangroves and reed (Phragmites australis),[68] can grow with their roots in anoxic conditions.

Factors affecting growth

The genome of a plant controls its growth. For example, selected varieties or genotypes of wheat grow rapidly, maturing within 110 days, whereas others, in the same environmental conditions, grow more slowly and mature within 155 days.[69]

Growth is also determined by environmental factors, such as temperature, available water, available light, carbon dioxide and available nutrients in the soil. Any change in the availability of these external conditions will be reflected in the plant's growth and the timing of its development.[citation needed]

Biotic factors also affect plant growth. Plants can be so crowded that no single individual produces normal growth, causing etiolation and chlorosis. Optimal plant growth can be hampered by grazing animals, suboptimal soil composition, lack of mycorrhizal fungi, and attacks by insects or plant diseases, including those caused by bacteria, fungi, viruses, and nematodes.[69]

 
There is no photosynthesis in deciduous leaves in autumn.

Simple plants like algae may have short life spans as individuals, but their populations are commonly seasonal. Annual plants grow and reproduce within one growing season, biennial plants grow for two growing seasons and usually reproduce in second year, and perennial plants live for many growing seasons and once mature will often reproduce annually. These designations often depend on climate and other environmental factors. Plants that are annual in alpine or temperate regions can be biennial or perennial in warmer climates. Among the vascular plants, perennials include both evergreens that keep their leaves the entire year, and deciduous plants that lose their leaves for some part of it. In temperate and boreal climates, they generally lose their leaves during the winter; many tropical plants lose their leaves during the dry season.[citation needed]

The growth rate of plants is extremely variable. Some mosses grow less than 0.001 millimeters per hour (mm/h), while most trees grow 0.025–0.250 mm/h. Some climbing species, such as kudzu, which do not need to produce thick supportive tissue, may grow up to 12.5 mm/h.[citation needed] Plants protect themselves from frost and dehydration stress with antifreeze proteins, heat-shock proteins and sugars (sucrose is common). LEA (Late Embryogenesis Abundant) protein expression is induced by stresses and protects other proteins from aggregation as a result of desiccation and freezing.[70]

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.[71] Intracellular freezing, which usually kills the cell[72] 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.[73] At rates of cooling of a few degrees Celsius per hour, segregation of ice occurs in intercellular spaces.[74] 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 −7 °C (19 °F).[71] 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 directly, or indirectly via the generation of reactive oxygen species.[75] Plants are capable of a DNA damage response that is a critical mechanism for maintaining genome stability.[76] The DNA damage response is particularly important during seed germination, since seed quality tends to deteriorate with age in association with DNA damage accumulation.[77] During germination repair processes are activated to deal with this accumulated DNA damage.[78] In particular, single- and double-strand breaks in DNA can be repaired.[79] Double-strand repair in plants often produce DNA junctions with structural alterations.[80]: 89  The DNA checkpoint kinase ATM has a key role in integrating progression through germination with repair responses to the DNA damages accumulated by the aged seed.[81]

Plant cells

 
Plant cell structure

Plant cells are typically distinguished by their large water-filled central vacuole, chloroplasts, and rigid cell walls that are made up of 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, storage, protective (e.g. epidermal layer), or reproductive tissues, with more primitive plants lacking some tissue types.[82]

Physiology

Photosynthesis

Plants photosynthesize, which means that they manufacture their own food molecules using energy obtained from light. The primary mechanism plants have for capturing light energy is the pigment chlorophyll. All green plants contain two forms of chlorophyll, chlorophyll a and chlorophyll b. The latter of these pigments is not found in red or brown algae. The simple equation of photosynthesis is as follows:

 

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.[83] 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)[84] and in Arabidopsis thaliana (FLS2, 2000).[85] 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 and phloem. They also have roots 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.[82]

Genomics

Plants have some of the largest genomes among all organisms.[86] The largest plant genome (in terms of gene number) is that of wheat (Triticum asestivum), predicted to encode ≈94,000 genes[87] 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.[88] In terms of sheer DNA sequence, the smallest published genome is that of the carnivorous bladderwort (Utricularia gibba) at 82 Mb (although it still encodes 28,500 genes)[89] while the largest, from the Norway Spruce (Picea abies), extends over 19,600 Mb (encoding about 28,300 genes).[90]

Ecology

The photosynthesis conducted by land plants and algae is the ultimate source of energy and organic material in nearly all ecosystems. Photosynthesis, at first by cyanobacteria and later by photosynthetic eukaryotes, radically changed the composition of the early Earth's anoxic atmosphere, which as a result is now 21% oxygen. Animals and most other organisms are aerobic, relying on oxygen; those that do not are confined to relatively rare anaerobic environments. Plants are the 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.[citation needed] Plants form about 80% of the world biomass at about 450 gigatonnes (4.4×1011 long tons; 5.0×1011 short tons) of carbon.[91]

Land plants are key components of the water cycle and several other biogeochemical cycles. Some plants have coevolved with 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.[citation needed]

Distribution

Plants are distributed almost worldwide. While they inhabit a multitude of biomes and ecoregions, few can be found beyond the tundras at the northernmost regions of continental shelves. At the southern extremes, plants of the Antarctic flora have adapted tenaciously to the prevailing conditions.[citation needed]

Plants are often the dominant physical and structural component of habitats where they occur. Many of the Earth's biomes are named for the type of vegetation because plants are the dominant organisms in those biomes, such as grasslands, taiga and tropical rainforest.[citation needed]

Ecological relationships

 

Numerous animals have coevolved with plants. Many animals pollinate flowers in exchange for food in the form of pollen or nectar. Many animals disperse seeds, often by eating 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 mutualistic symbiosis known as mycorrhiza. The fungi help the plants gain water and mineral nutrients from the soil, while the plant gives the fungi carbohydrates manufactured in photosynthesis. Some plants serve as homes for 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.[92]

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 broomrape and toothwort that acquire all their nutrients through connections to the roots of other plants, and so have no chlorophyll. Some plants, known as myco-heterotrophs, parasitize mycorrhizal fungi, and hence act as epiparasites on other plants.

Many plants are epiphytes, meaning they grow on other plants, usually trees, without parasitizing them. Epiphytes may indirectly harm their host plant by intercepting mineral nutrients and light that the host would otherwise receive. The weight of large numbers of epiphytes may break tree limbs. Hemiepiphytes like the strangler fig begin as epiphytes but eventually set their own roots and overpower and kill their host. Many orchids, bromeliads, ferns and mosses often grow as epiphytes. Bromeliad epiphytes accumulate water in leaf axils to form phytotelmata that may contain complex aquatic food webs.[93]

Approximately 630 plants are carnivorous, such as the Venus Flytrap (Dionaea muscipula) and sundew (Drosera species). They trap small animals and digest them to obtain mineral nutrients, especially nitrogen and phosphorus.[94]

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.[95] Eventually, one species will out-compete the other, which will push the disadvantaged species to extinction.[95]

In regard to plants, competition tends to negatively affect their growth when competing for shared resources.[96] These shared resources commonly include space for growth, sunlight, water and nutrients. Light is an important resource because it is necessary for photosynthesis.[96] Plants use their leaves to shade other plants from sunlight and grow quickly to maximize their own expose.[96] Water is also important for photosynthesis, and plants have different root systems to maximize water uptake from soil.[97] 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.[97]

Minerals are also important for plant growth and development, where deficiencies can occur if nutrient needs are not met.[98] Common nutrients competed for amongst plants include nitrogen and phosphorus. Space is also extremely important for a growing and developing plant.[99] 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.[99] If an old tree dies, then competition arises amongst a number of trees to replace it.[96] Those that are less effective competitors are less likely to contribute to the next generation of offspring.[96]

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.[100]

Importance

Cultivation

The study of plant uses by people is called economic botany or ethnobotany.[101] Human cultivation of plants is part of agriculture, which is the basis of human civilization.[102] Plant agriculture is subdivided into agronomy, horticulture and forestry.[103]

Food

 
Mechanical harvest of oats

Humans depend on plants for food, either directly or as feed for domestic animals. Agriculture deals with the production of food crops, and has 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.[104] 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 staples include cereals such as rice and wheat, starchy roots and tubers such as cassava and potato, and legumes such as peas and beans. Vegetable oils such as olive oil and palm oil provide lipids, while fruit and vegetables contribute vitamins and minerals to the diet.[105]

Medicines

 
Melocactus plants being used as medicine

Medicinal plants are a primary source of organic compounds, both for their medicinal and physiological effects, and for the industrial synthesis of a vast array of organic chemicals.[106] Many hundreds of medicines are derived from plants, both traditional medicines used in herbalism[107][108] and chemical substances purified from plants or first identified in them, sometimes by ethnobotanical search, and then synthesised for use in modern medicine. Modern medicines derived from plants include aspirin, taxol, morphine, quinine, reserpine, colchicine, digitalis and vincristine. Plants used in herbalism include 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 CE and remained in use in Europe and the Middle East until around 1600 CE; it was the precursor of all modern pharmacopoeias.[109][110][111]

Nonfood products

 
Timber in storage for later processing at a sawmill

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.[112] Nonfood products include essential oils, natural dyes, pigments, waxes, resins, tannins, alkaloids, amber and cork. Products derived from plants include soaps, shampoos, perfumes, cosmetics, paint, varnish, turpentine, rubber, latex, lubricants, linoleum, plastics, inks, and gums. Renewable fuels from plants include firewood, peat and other biofuels.[113][114] The fossil fuels coal, petroleum and natural gas are derived from the remains of aquatic organisms including phytoplankton in geological time.[115]

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 pulped to make paper and cardboard.[116] Cloth is often made from cotton, flax, ramie or synthetic fibres such as rayon and acetate derived from plant cellulose. Thread used to sew cloth likewise comes in large part from cotton.[117]

Aesthetic uses

 
A rose espalier at Niedernhall in Germany

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 historic gardens, national parks, rainforests, forests with colorful autumn leaves, and festivals such as Japan's[118] and America's cherry blossom festivals.[119]

 
Capitals of ancient Egyptian columns decorated to resemble papyrus plants (at Luxor, Egypt)

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.[citation needed]

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 tulipomania.[120]

Architectural designs resembling plants appear in the capitals of Ancient Egyptian columns, which were carved to resemble either the Egyptian white lotus or the papyrus.[121] Images of plants are often used in painting and photography, as well as on textiles, money, stamps, flags and coats of arms.[citation needed]

Scientific and cultural uses

 
Barbara McClintock (1902–1992) was a pioneering cytogeneticist who used maize (corn) to study the mechanism of inheritance of traits.

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,[122] and examination of chromosomes in maize allowed Barbara McClintock to demonstrate their connection to inherited traits.[123] The plant Arabidopsis thaliana is used in laboratories as a model organism to understand how genes control the growth and development of plant structures.[124] NASA predicts that space stations or space colonies will one day rely on plants for life support.[125]

Ancient trees are revered and many are famous. Tree rings themselves are an important method of dating in archeology, and serve as a record of past climates.[126]

Plants figure prominently in mythology, religion and literature.[127][128][129] They are used as national and state emblems, including 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 messages.[citation needed]

Negative effects

Weeds are commercially or aesthetically undesirable plants growing in managed environments such as farms, urban areas, gardens, lawns, and parks. People have spread plants beyond their native ranges and some of these introduced plants become invasive, damaging existing ecosystems by displacing native species, and sometimes becoming serious weeds of cultivation.[citation needed]

Plants may cause harm to animals, including people. Plants that produce windblown pollen invoke allergic reactions in people who suffer from hay fever. A wide variety of plants are 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 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.[130][131] Both illegal and legal drugs derived from plants may have negative effects on the economy, affecting worker productivity and law enforcement costs.[132][133]

See also

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Further reading

General:

  • Evans, L.T. (1998). Feeding the Ten Billion – Plants and Population Growth. Cambridge University Press. Paperback, 247 pages. ISBN 0-521-64685-5.
  • Kenrick, Paul & Crane, Peter R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, D.C.: Smithsonian Institution Press. ISBN 1-56098-730-8.
  • Raven, Peter H.; Evert, Ray F.; & Eichhorn, Susan E. (2005). Biology of Plants (7th ed.). New York: W.H. Freeman and Company. ISBN 0-7167-1007-2.
  • Taylor, Thomas N. & Taylor, Edith L. (1993). The Biology and Evolution of Fossil Plants. Englewood Cliffs, NJ: Prentice Hall. ISBN 0-13-651589-4.
  • Trewavas A (2003). "Aspects of Plant Intelligence". Annals of Botany. 92 (1): 1–20. doi:10.1093/aob/mcg101. PMC 4243628. PMID 12740212.

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.
  • Prance G.T. (2001). "Discovering the Plant World". Taxon. 50 (2, Golden Jubilee Part 4): 345–359. doi:10.2307/1223885. JSTOR 1223885.

External links

  • Jones, T.M.; Reid, C.S.; Urbatsch, L.E. "Visual study of divisional Plantae". {{cite journal}}: Cite journal requires |journal= (help) (requires Microsoft Silverlight)
  • Chaw, S.-M.; et al. (1997). (PDF). Mol. Biol. Evol. 14 (1): 56–68. doi:10.1093/oxfordjournals.molbev.a025702. PMID 9000754. Archived from the original (PDF) on 24 January 2005.
  • Index Nominum Algarum
  • 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 Europaea
  • Flora of Central Europe (in German)
  • Flora of North America
  • List of Japanese Wild Plants Online
  • 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

plant, other, uses, disambiguation, predominantly, photosynthetic, eukaryotes, kingdom, historically, plant, kingdom, encompassed, living, things, that, were, animals, included, algae, fungi, however, current, definitions, exclude, fungi, some, algae, well, pr. For other uses see Plant disambiguation Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae Historically the plant kingdom encompassed all living things that were not animals and included algae and fungi however all current definitions of Plantae exclude the fungi and some algae as well as the prokaryotes the archaea and bacteria By one definition plants form the clade Viridiplantae Latin name for green plants which is sister of the Glaucophyta and consists of the green algae and Embryophyta land plants The latter includes the flowering plants conifers and other gymnosperms ferns and their allies hornworts liverworts and mosses PlantsTemporal range Mesoproterozoic present Pha Proterozoic Archean Had nScientific classificationDomain Eukaryota unranked Diaphoretickes unranked ArchaeplastidaKingdom Plantaesensu Copeland 1956SuperdivisionsChlorokybophyta Mesostigmatophyta Spirotaenia Chlorobionta Kenrick amp Crane 1997 Chlorophyta Streptobionta Kenrick amp Crane 1997 Klebsormidiophyceae Charophyta stoneworts Chaetosphaeridiales Coleochaetophyta Zygnematophyta Embryophyta Engler 1892 land plants Marchantiophyta liverworts Bryophyta mosses Anthocerotophyta hornworts Horneophyta Aglaophyta Tracheophyta vascular plants SynonymsViridiplantae Cavalier Smith 1981 1 Chlorobionta Jeffrey 1982 emend Bremer 1985 emend Lewis and McCourt 2004 2 Chlorobiota Kenrick and Crane 1997 3 Chloroplastida Adl et al 2005 4 Phyta Barkley 1939 emend Holt amp Uidica 2007 Cormophyta Endlicher 1836 Cormobionta Rothmaler 1948 Euplanta Barkley 1949 Telomobionta Takhtajan 1964 Embryobionta Cronquist et al 1966 Metaphyta Whittaker 1969Most plants are multicellular organisms Green plants obtain most of their energy from sunlight via photosynthesis by primary chloroplasts that are derived from endosymbiosis with cyanobacteria Their chloroplasts contain chlorophylls a and b which gives them their green color Some plants are 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 and alternation of generations although asexual reproduction is also common There are about 320 000 known species of plants of which the great majority some 260 000 290 000 produce seeds 5 Green plants provide a substantial proportion of the world s molecular oxygen 6 and are the basis of most of Earth s ecosystems Plants that produce grain fruit and vegetables also form basic human foods and have been domesticated for millennia Plants have many cultural and other uses as ornaments building materials writing material and in great variety they have been the source of medicines and psychoactive drugs The scientific study of plants is known as botany a branch of biology Contents 1 Definition 1 1 Current definitions of Plantae 1 2 Algae 1 3 Fungi 2 Diversity 2 1 Evolution 2 2 Embryophytes 2 3 Fossils 3 Structure growth and development 3 1 Factors affecting growth 3 1 1 Effects of freezing 3 2 DNA damage and repair 3 3 Plant cells 4 Physiology 4 1 Photosynthesis 4 2 Immune system 4 3 Internal distribution 5 Genomics 6 Ecology 6 1 Distribution 6 2 Ecological relationships 6 3 Competition 7 Importance 7 1 Cultivation 7 1 1 Food 7 1 2 Medicines 7 1 3 Nonfood products 7 1 4 Aesthetic uses 7 2 Scientific and cultural uses 7 3 Negative effects 8 See also 9 References 10 Further reading 11 External linksDefinitionAll living things were traditionally placed into one of two groups plants and animals This classification may date from Aristotle 384 322 BCE 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 1707 1778 created the basis of the modern system of scientific classification these two groups became the kingdoms Vegetabilia later Metaphyta or Plantae and Animalia also called Metazoa Since then it has become clear that the plant kingdom as originally defined included several unrelated groups and the fungi and several groups of algae were removed to new kingdoms However these organisms are still sometimes considered plants particularly in informal contexts citation needed The term plant generally implies the possession of the following traits multicellularity possession of cell walls containing cellulose and the ability to carry out photosynthesis with primary chloroplasts 7 8 Current definitions of Plantae When the name Plantae or plant is applied to a specific group of organisms or taxon it usually refers to one of four concepts From least to most inclusive these four groupings are Name s Scope DescriptionLand plants also known as Embryophyta Plantae sensu strictissimo Plants in the strictest sense include the liverworts hornworts mosses and vascular plants as well as fossil plants similar to these surviving groups e g Metaphyta Whittaker 1969 9 Plantae Margulis 1971 10 Green plants also known as Viridiplantae Viridiphyta Chlorobionta or Chloroplastida Plantae sensu stricto Plants in a strict sense include the green algae and land plants that emerged within them including stoneworts The relationships between plant groups are still being worked out and the names given to them vary considerably The clade Viridiplantae encompasses a group of organisms that have cellulose in their cell walls possess chlorophylls a and b and have plastids bound by only two membranes that are capable of photosynthesis and of storing starch This clade is the main subject of this article e g Plantae Copeland 1956 11 Archaeplastida also known as Plastida or Primoplantae Plantae sensu lato Plants in a broad sense comprise the green plants listed above plus the red algae Rhodophyta and the glaucophyte algae Glaucophyta that store Floridean starch outside the plastids in the cytoplasm This clade includes all of the organisms that eons ago acquired their primary chloroplasts directly by engulfing cyanobacteria e g Plantae Cavalier Smith 1981 12 Old definitions of plant obsolete Plantae sensu amplo Plants in the widest sense refers to older obsolete classifications that placed diverse algae fungi or bacteria in Plantae e g Plantae or Vegetabilia Linnaeus 13 Plantae Haeckel 1866 14 Metaphyta Haeckel 1894 15 Plantae Whittaker 1969 9 Another way of looking at the relationships between the different groups that have been called plants is through a cladogram which shows their evolutionary relationships These are not yet completely settled but one accepted relationship between the three groups described above is shown below clarification needed 16 17 18 19 20 21 22 23 24 excessive citations Those which have been called plants are in bold some minor groups have been omitted Archaeplastida cryptista Rhodophyta red algae Rhodelphidia predatorial PicozoaGlaucophyta glaucophyte algae green plants Prasinodermophyta Palmophyllophyceae prasinophyte clade VI PrasinodermophyceaeChlorophytaStreptophyta Charales stoneworts land plants or embryophytesCryptista groups traditionallycalled green algaeThe way in which the groups of green algae are combined and named varies considerably between authors Algae Green algae from Ernst Haeckel s Kunstformen der Natur 1904 Main article Algae Algae consist of several groups of organisms which produce food by photosynthesis and thus have traditionally been included in the plant kingdom The seaweeds range from large multicellular algae to single celled organisms and are classified into three groups the green algae red algae and brown algae 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 25 26 The Viridiplantae the green plants green algae and land plants form a clade a group consisting of all the descendants of a common ancestor With a few exceptions the green plants have the following features in common primary chloroplasts derived from cyanobacteria containing chlorophylls a and b cell walls containing cellulose and food stores in the form of starch contained within the plastids They undergo closed mitosis without centrioles and typically have mitochondria with flat cristae The chloroplasts of green plants are surrounded by two membranes suggesting they originated directly from endosymbiotic cyanobacteria Two additional groups the Rhodophyta red algae and Glaucophyta glaucophyte algae also have primary chloroplasts that appear to be derived directly from endosymbiotic cyanobacteria although they differ from Viridiplantae in the pigments which are used in photosynthesis and so are different in colour These groups also differ from green plants in that the storage polysaccharide is floridean starch 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 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 haptophytes dinoflagellates and euglenids not only have different pigments but also have chloroplasts with three or four surrounding membranes They are not close relatives of the Archaeplastida presumably having acquired chloroplasts separately from ingested or symbiotic green and red algae They are thus not included in even the broadest modern definition of the plant kingdom although they were in the past The green plants or Viridiplantae were traditionally divided into the green algae including the stoneworts and the land plants However it is now known that the land plants evolved from within a group of green algae so that the green algae by themselves are a paraphyletic group that is 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 and the Streptophyta including the land plants and Charophyta 27 28 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 29 mainly unicellular or multicellular marine organisms such as the sea lettuce Ulva 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 desmids and stoneworts Streptophyte algae are either unicellular or form multicellular filaments branched or unbranched 28 The genus 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 30 Growing immersed in fresh water they consist of a central stalk with whorls of branchlets Fungi This section relies largely or entirely upon a single source Relevant discussion may be found on the talk page Please help improve this article by introducing citations to additional sources June 2021 Learn how and when to remove this template message Main article Fungi A variety of fungi species Linnaeus original classification placed the fungi within the Plantae since they were unquestionably neither animals or minerals and these were the only other alternatives With 19th century developments in microbiology Ernst Haeckel introduced the new kingdom Protista in addition to Plantae and Animalia but whether fungi were best placed in the Plantae or should be reclassified as 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 concestor of the Fungi was probably more similar to that of the Animalia than to that of Plantae or any other kingdom 31 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 autotrophs fungi do not possess chloroplasts and generally obtain carbon by breaking down and absorbing surrounding materials and so are called heterotrophic saprotrophs In addition the substructure of multicellular fungi is different from that of plants taking the form of many chitinous microscopic strands called hyphae which may be further subdivided into cells or may form a syncytium containing many eukaryotic nuclei Fruiting bodies of which mushrooms are the most familiar example are the reproductive structures of fungi and are unlike any structures produced by plants citation needed DiversityThe 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 both list about 391 000 species 32 33 34 Diversity of living green plant Viridiplantae divisions Informal group Division name citation needed Common name No of living species Approximate no in informal groupGreen algae Chlorophyta Green algae chlorophytes 3 800 4 300 35 36 8 500 6 600 10 300 Charophyta Green algae e g desmids amp stoneworts 2 800 6 000 37 38 Bryophytes Marchantiophyta Liverworts 6 000 8 000 39 19 000 18 100 20 200 Anthocerotophyta Hornworts 100 200 40 Bryophyta Mosses 12 000 41 Pteridophytes Lycopodiophyta Clubmosses 1 200 26 12 000 12 200 Polypodiophyta Ferns whisk ferns amp horsetails 11 000 26 spermatophyte Cycadophyta Cycads 160 42 260 000 259 511 Ginkgophyta Ginkgo 1 43 Pinophyta Conifers 630 26 Gnetophyta Gnetophytes 70 26 Magnoliophyta Flowering plants 258 650 44 The naming of plants is governed by the International Code of Nomenclature for algae fungi and plants and International Code of Nomenclature for Cultivated Plants see cultivated plant taxonomy Evolution Further information Evolutionary history of plants The evolution of plants has resulted in increasing levels of complexity from the earliest algal mats through bryophytes lycopods ferns to the complex gymnosperms and angiosperms of today Plants in all of these groups continue to thrive especially in the environments in which they evolved An algal scum formed on the land 1 200 million years ago but it was not until the Ordovician Period around 450 million years ago that land plants appeared 45 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 46 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 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 47 and that even the common ancestor of red and green algae and the unicellular freshwater algae glaucophytes originated in a terrestrial environment in freshwater biofilms or microbial mats 48 Primitive land plants began to diversify in the late Silurian Period around 420 million years ago and the results of their diversification are displayed in remarkable detail in an early Devonian fossil assemblage from the 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 49 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 200 million years ago 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 40 million years ago The grasses as well as many other groups evolved new mechanisms of metabolism to survive the low CO2 and warm dry conditions of the tropics over the last 10 million years A 1997 proposed phylogenetic tree of Plantae after Kenrick and Crane 50 is as follows with modification to the Pteridophyta from Smith et al 51 The Prasinophyceae are a paraphyletic assemblage of early diverging green algal lineages but are treated as a group outside the Chlorophyta 52 later authors have not followed this suggestion Prasinophyceae micromonads Streptobionta Embryophytes Stomatophytes Polysporangiates Tracheophytes Eutracheophytes Euphyllophytina Lignophyta Spermatophytes seed plants Progymnospermophyta Pteridophyta Pteridopsida true ferns MarattiopsidaEquisetopsida horsetails Psilotopsida whisk ferns amp adders tongues Cladoxylopsida Lycophytina LycopodiophytaZosterophyllophyta Rhyniophyta Aglaophyton Horneophytopsida Bryophyta mosses Anthocerotophyta hornworts Marchantiophyta liverworts CharophytaChlorophyta Trebouxiophyceae Pleurastrophyceae ChlorophyceaeUlvophyceaeA newer proposed classification follows Leliaert et al 2011 53 and modified with Silar 2016 20 21 54 55 for the green algae clades and Novikov amp Barabas Krasni 2015 56 for the land plants clade Notice that the Prasinophyceae are here placed inside the Chlorophyta Viridiplantae MesostigmatophyceaeChlorokybophyceaeSpirotaeniaChlorophyta inc PrasinophyceaeStreptobionta StreptofilumKlebsormidiophytaPhragmoplastophyta Charophyta Rabenhorst 1863 emend Lewis amp McCourt 2004 Stoneworts ColeochaetophytaZygnematophytaEmbryophyta Marchantiophyta Liverworts Stomatophyta Bryophyta True mosses Anthocerotophyta Non flowering hornworts Polysporangiophyta Horneophyta AglaophytaTracheophyta Vascular Plants Green algaeLater a phylogeny based on genomes and transcriptomes from 1 153 plant species was proposed 57 The placing of algal groups is supported by phylogenies based on genomes from the Mesostigmatophyceae and Chlorokybophyceae that have since been sequenced 58 59 The classification of Bryophyta is supported both by Puttick et al 2018 60 and by phylogenies involving the hornwort genomes that have also since been sequenced 61 62 RhodophytaGlaucophytaViridiplantae ChlorophytaPrasinococcales MesostigmatophyceaeChlorokybophyceaeSpirotaeniaKlebsormidialesCharaColeochaetalesZygnematophyceaeBryophytes HornwortsLiverwortsMossesLycophytesFernsSpermatophytes GymnospermsAngiosperms chlorophyte algae gradestreptophyte algae gradeEmbryophytes Main article Embryophyte This section does not cite any sources Please help improve this section by adding citations to reliable sources Unsourced material may be challenged and removed November 2018 Learn how and when to remove this template message Dicksonia antarctica a species of tree fernThe plants that are likely most familiar to us are the multicellular land plants called embryophytes Embryophytes include the vascular plants such as ferns conifers and flowering plants They also include the bryophytes of which mosses and liverworts are the most common All of these plants have eukaryotic cells with cell walls composed of cellulose and most obtain their energy through photosynthesis using light water and carbon dioxide to synthesize food About three hundred plant species do not photosynthesize but are parasites on other species of photosynthetic plants Embryophytes are distinguished from green algae which represent a mode of photosynthetic life similar to the kind modern plants are believed to have evolved from by having specialized reproductive organs protected by non reproductive tissues Bryophytes first appeared during the early Paleozoic They mainly live in habitats where moisture is available for significant periods although some species such as Targionia are desiccation tolerant Most species of bryophytes remain small throughout their life cycle This involves an alternation between two generations a haploid stage called the gametophyte and a diploid stage called the sporophyte In bryophytes the sporophyte is always unbranched and remains nutritionally dependent on its parent gametophyte The embryophytes have the ability to secrete a cuticle on their outer surface a waxy layer that confers resistance to desiccation In the mosses and hornworts a cuticle is usually only produced on the sporophyte Stomata are absent from liverworts but occur on the sporangia of mosses and hornworts allowing gas exchange Vascular plants first appeared during the Silurian period and by the Devonian had diversified and spread into many different terrestrial environments They developed a number of adaptations that allowed them to spread into increasingly more arid places notably the vascular tissues xylem and 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 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 After fertilisation by means of sperm produced by pollen grains an embryo sporophyte develops inside the ovule The integument becomes a seed coat and the ovule develops into a seed Seed plants can survive and reproduce in extremely arid conditions because they are not dependent on free water for the movement of sperm or the development of free living gametophytes The first seed plants pteridosperms seed ferns now extinct appeared in the Devonian and diversified through the Carboniferous They were the ancestors of modern gymnosperms of which four surviving groups are widespread today particularly the conifers which are dominant trees in several biomes The name gymnosperm comes from the Greek gymnospermos a composite of gymnos gymnos lit naked and sperma sperma lit seed 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 Main articles Paleobotany and Evolutionary history of plants A petrified log in Petrified Forest National Park Arizona This section needs additional citations for verification Please help improve this article by adding citations to reliable sources Unsourced material may be challenged and removed November 2018 Learn how and when to remove this template message Plant fossils include roots wood leaves seeds fruit pollen spores phytoliths and amber the fossilized resin produced by some plants Fossil land plants are recorded in terrestrial lacustrine fluvial and nearshore marine sediments Pollen spores and algae dinoflagellates and acritarchs are used for dating sedimentary rock sequences The remains of fossil plants are not as common as fossil animals although plant fossils are locally abundant in many regions worldwide The earliest fossils clearly assignable to Kingdom Plantae are fossil green algae from the Cambrian These fossils resemble calcified multicellular members of the Dasycladales Earlier Precambrian fossils are known that resemble single cell green algae but definitive identity with that group of algae is uncertain The earliest fossils attributed to green algae date from the Precambrian ca 1200 mya 63 64 The resistant outer walls of prasinophyte cysts known as phycomata are well preserved in fossil deposits of the Paleozoic ca 250 540 mya A filamentous fossil Proterocladus from middle Neoproterozoic deposits ca 750 mya has been attributed to the Cladophorales while the oldest reliable records of the Bryopsidales Dasycladales and stoneworts are from the Paleozoic 52 65 The oldest known fossils of embryophytes date from the Ordovician though such fossils are fragmentary By the Silurian fossils of whole plants are preserved including the simple vascular plant Cooksonia in mid Silurian and the much larger and more complex lycophyte Baragwanathia longifolia in late Silurian From the early Devonian Rhynie chert detailed fossils of lycophytes and rhyniophytes have been found that show details of the individual cells within the plant organs and the symbiotic association of these plants with fungi of the order Glomales The Devonian period also saw the evolution of leaves and roots and the first modern tree Archaeopteris This tree with fern like foliage and a trunk with conifer like wood was heterosporous producing spores of two different sizes an early step in the evolution of seeds 66 The Coal measures are a major source of Paleozoic 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 itself is the remains of fossilised plants though structural detail of the plant fossils is rarely visible in coal In the Fossil Grove at Victoria Park in Glasgow Scotland the stumps of Lepidodendron trees are found in their original growth positions The fossilized remains of conifer and angiosperm roots stems and branches may be locally abundant in lake and inshore sedimentary rocks from the Mesozoic and Cenozoic eras Sequoia and its allies magnolia oak and palms are often found Range of pangaea glossopteris 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 Petrified wood is often heavily silicified the organic material replaced by silicon dioxide and the impregnated tissue is often preserved in fine detail Such specimens may be cut and polished using lapidary equipment Fossil forests of petrified wood have been found in all continents Fossils of seed ferns such as Glossopteris are widely distributed throughout several continents of the Southern Hemisphere a fact that gave support to Alfred Wegener s early ideas regarding Continental drift theory Structure growth and developmentFurther information Plant morphology The leaf is usually the primary site of photosynthesis in plants Most of the solid material in a plant is taken from the atmosphere Through the process of photosynthesis most plants use the energy in sunlight to convert carbon dioxide from the atmosphere plus water into simple sugars These sugars are then used as building blocks and form the main structural component of the plant Chlorophyll a green colored magnesium containing pigment is essential to this process it is generally present in plant leaves and often in other plant parts as well Parasitic plants 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 of nitrogen phosphorus potassium magnesium and other elemental nutrients from the soil Epiphytic and lithophytic plants depend on air and nearby debris for nutrients and carnivorous plants supplement their nutrient requirements particularly for nitrogen and phosphorus with insect prey that they capture For the majority of plants to grow successfully they also require oxygen in the atmosphere and around their roots soil gas for respiration Plants use oxygen and glucose which may be produced from stored starch to provide energy 67 Some plants grow as submerged aquatics using oxygen dissolved in the surrounding water and a few specialized vascular plants such as mangroves and reed Phragmites australis 68 can grow with their roots in anoxic conditions Factors affecting growth The genome of a plant controls its growth For example selected varieties or genotypes of wheat grow rapidly maturing within 110 days whereas others in the same environmental conditions grow more slowly and mature within 155 days 69 Growth is also determined by environmental factors such as temperature available water available light carbon dioxide and available nutrients in the soil Any change in the availability of these external conditions will be reflected in the plant s growth and the timing of its development citation needed Biotic factors also affect plant growth Plants can be so crowded that no single individual produces normal growth causing etiolation and chlorosis Optimal plant growth can be hampered by grazing animals suboptimal soil composition lack of mycorrhizal fungi and attacks by insects or plant diseases including those caused by bacteria fungi viruses and nematodes 69 There is no photosynthesis in deciduous leaves in autumn Simple plants like algae may have short life spans as individuals but their populations are commonly seasonal Annual plants grow and reproduce within one growing season biennial plants grow for two growing seasons and usually reproduce in second year and perennial plants live for many growing seasons and once mature will often reproduce annually These designations often depend on climate and other environmental factors Plants that are annual in alpine or temperate regions can be biennial or perennial in warmer climates Among the vascular plants perennials include both evergreens that keep their leaves the entire year and deciduous plants that lose their leaves for some part of it In temperate and boreal climates they generally lose their leaves during the winter many tropical plants lose their leaves during the dry season citation needed The growth rate of plants is extremely variable Some mosses grow less than 0 001 millimeters per hour mm h while most trees grow 0 025 0 250 mm h Some climbing species such as kudzu which do not need to produce thick supportive tissue may grow up to 12 5 mm h citation needed Plants protect themselves from frost and dehydration stress with antifreeze proteins heat shock proteins and sugars sucrose is common LEA Late Embryogenesis Abundant protein expression is induced by stresses and protects other proteins from aggregation as a result of desiccation and freezing 70 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 71 Intracellular freezing which usually kills the cell 72 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 73 At rates of cooling of a few degrees Celsius per hour segregation of ice occurs in intercellular spaces 74 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 7 C 19 F 71 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 directly or indirectly via the generation of reactive oxygen species 75 Plants are capable of a DNA damage response that is a critical mechanism for maintaining genome stability 76 The DNA damage response is particularly important during seed germination since seed quality tends to deteriorate with age in association with DNA damage accumulation 77 During germination repair processes are activated to deal with this accumulated DNA damage 78 In particular single and double strand breaks in DNA can be repaired 79 Double strand repair in plants often produce DNA junctions with structural alterations 80 89 The DNA checkpoint kinase ATM has a key role in integrating progression through germination with repair responses to the DNA damages accumulated by the aged seed 81 Plant cells Plant cell structureThis section relies largely or entirely upon a single source Relevant discussion may be found on the talk page Please help improve this article by introducing citations to additional sources June 2021 Learn how and when to remove this template message Main article Plant cell Plant cells are typically distinguished by their large water filled central vacuole chloroplasts and rigid cell walls that are made up of 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 storage protective e g epidermal layer or reproductive tissues with more primitive plants lacking some tissue types 82 PhysiologyMain article Plant physiology Photosynthesis This section does not cite any sources Please help improve this section by adding citations to reliable sources Unsourced material may be challenged and removed June 2021 Learn how and when to remove this template message Main articles Photosynthesis and Biological pigment Plants photosynthesize which means that they manufacture their own food molecules using energy obtained from light The primary mechanism plants have for capturing light energy is the pigment chlorophyll All green plants contain two forms of chlorophyll chlorophyll a and chlorophyll b The latter of these pigments is not found in red or brown algae The simple equation of photosynthesis is as follows 6 CO 2 6 H 2 O light C 6 H 12 O 6 6 O 2 displaystyle ce 6CO2 6H2O gt text light C6H12O6 6O2 Immune system See also Immune system and Plant disease resistance 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 83 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 84 and in Arabidopsis thaliana FLS2 2000 85 Plants also carry immune receptors that recognize highly variable pathogen effectors These include the NBS LRR class of proteins Internal distribution This section relies largely or entirely upon a single source Relevant discussion may be found on the talk page Please help improve this article by introducing citations to additional sources June 2021 Learn how and when to remove this template message Main article Vascular tissue Vascular plants differ from other plants in that nutrients are transported between their different parts through specialized structures called xylem and phloem They also have roots 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 82 GenomicsPlants have some of the largest genomes among all organisms 86 The largest plant genome in terms of gene number is that of wheat Triticum asestivum predicted to encode 94 000 genes 87 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 88 In terms of sheer DNA sequence the smallest published genome is that of the carnivorous bladderwort Utricularia gibba at 82 Mb although it still encodes 28 500 genes 89 while the largest from the Norway Spruce Picea abies extends over 19 600 Mb encoding about 28 300 genes 90 EcologyMain article Plant ecology The photosynthesis conducted by land plants and algae is the ultimate source of energy and organic material in nearly all ecosystems Photosynthesis at first by cyanobacteria and later by photosynthetic eukaryotes radically changed the composition of the early Earth s anoxic atmosphere which as a result is now 21 oxygen Animals and most other organisms are aerobic relying on oxygen those that do not are confined to relatively rare anaerobic environments Plants are the 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 citation needed Plants form about 80 of the world biomass at about 450 gigatonnes 4 4 1011 long tons 5 0 1011 short tons of carbon 91 Land plants are key components of the water cycle and several other biogeochemical cycles Some plants have coevolved with 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 citation needed Distribution This section does not cite any sources Please help improve this section by adding citations to reliable sources Unsourced material may be challenged and removed June 2021 Learn how and when to remove this template message Plants are distributed almost worldwide While they inhabit a multitude of biomes and ecoregions few can be found beyond the tundras at the northernmost regions of continental shelves At the southern extremes plants of the Antarctic flora have adapted tenaciously to the prevailing conditions citation needed Plants are often the dominant physical and structural component of habitats where they occur Many of the Earth s biomes are named for the type of vegetation because plants are the dominant organisms in those biomes such as grasslands taiga and tropical rainforest citation needed Ecological relationships The Venus flytrap a species of carnivorous plant This section needs additional citations for verification Please help improve this article by adding citations to reliable sources Unsourced material may be challenged and removed November 2018 Learn how and when to remove this template message Numerous animals have coevolved with plants Many animals pollinate flowers in exchange for food in the form of pollen or nectar Many animals disperse seeds often by eating 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 mutualistic symbiosis known as mycorrhiza The fungi help the plants gain water and mineral nutrients from the soil while the plant gives the fungi carbohydrates manufactured in photosynthesis Some plants serve as homes for 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 92 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 broomrape and toothwort that acquire all their nutrients through connections to the roots of other plants and so have no chlorophyll Some plants known as myco heterotrophs parasitize mycorrhizal fungi and hence act as epiparasites on other plants Many plants are epiphytes meaning they grow on other plants usually trees without parasitizing them Epiphytes may indirectly harm their host plant by intercepting mineral nutrients and light that the host would otherwise receive The weight of large numbers of epiphytes may break tree limbs Hemiepiphytes like the strangler fig begin as epiphytes but eventually set their own roots and overpower and kill their host Many orchids bromeliads ferns and mosses often grow as epiphytes Bromeliad epiphytes accumulate water in leaf axils to form phytotelmata that may contain complex aquatic food webs 93 Approximately 630 plants are carnivorous such as the Venus Flytrap Dionaea muscipula and sundew Drosera species They trap small animals and digest them to obtain mineral nutrients especially nitrogen and phosphorus 94 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 95 Eventually one species will out compete the other which will push the disadvantaged species to extinction 95 In regard to plants competition tends to negatively affect their growth when competing for shared resources 96 These shared resources commonly include space for growth sunlight water and nutrients Light is an important resource because it is necessary for photosynthesis 96 Plants use their leaves to shade other plants from sunlight and grow quickly to maximize their own expose 96 Water is also important for photosynthesis and plants have different root systems to maximize water uptake from soil 97 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 97 Minerals are also important for plant growth and development where deficiencies can occur if nutrient needs are not met 98 Common nutrients competed for amongst plants include nitrogen and phosphorus Space is also extremely important for a growing and developing plant 99 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 99 If an old tree dies then competition arises amongst a number of trees to replace it 96 Those that are less effective competitors are less likely to contribute to the next generation of offspring 96 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 100 ImportanceMain article Plants in culture Cultivation The study of plant uses by people is called economic botany or ethnobotany 101 Human cultivation of plants is part of agriculture which is the basis of human civilization 102 Plant agriculture is subdivided into agronomy horticulture and forestry 103 Food Mechanical harvest of oats Main article Agriculture Humans depend on plants for food either directly or as feed for domestic animals Agriculture deals with the production of food crops and has 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 104 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 staples include cereals such as rice and wheat starchy roots and tubers such as cassava and potato and legumes such as peas and beans Vegetable oils such as olive oil and palm oil provide lipids while fruit and vegetables contribute vitamins and minerals to the diet 105 Medicines Main article Medicinal plants Melocactus plants being used as medicine Medicinal plants are a primary source of organic compounds both for their medicinal and physiological effects and for the industrial synthesis of a vast array of organic chemicals 106 Many hundreds of medicines are derived from plants both traditional medicines used in herbalism 107 108 and chemical substances purified from plants or first identified in them sometimes by ethnobotanical search and then synthesised for use in modern medicine Modern medicines derived from plants include aspirin taxol morphine quinine reserpine colchicine digitalis and vincristine Plants used in herbalism include 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 CE and remained in use in Europe and the Middle East until around 1600 CE it was the precursor of all modern pharmacopoeias 109 110 111 Nonfood products Timber in storage for later processing at a sawmill Main article Non food crop 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 112 Nonfood products include essential oils natural dyes pigments waxes resins tannins alkaloids amber and cork Products derived from plants include soaps shampoos perfumes cosmetics paint varnish turpentine rubber latex lubricants linoleum plastics inks and gums Renewable fuels from plants include firewood peat and other biofuels 113 114 The fossil fuels coal petroleum and natural gas are derived from the remains of aquatic organisms including phytoplankton in geological time 115 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 pulped to make paper and cardboard 116 Cloth is often made from cotton flax ramie or synthetic fibres such as rayon and acetate derived from plant cellulose Thread used to sew cloth likewise comes in large part from cotton 117 Aesthetic uses A rose espalier at Niedernhall in Germany Main article Ornamental plant 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 historic gardens national parks rainforests forests with colorful autumn leaves and festivals such as Japan s 118 and America s cherry blossom festivals 119 Capitals of ancient Egyptian columns decorated to resemble papyrus plants at Luxor Egypt 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 citation needed 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 tulipomania 120 Architectural designs resembling plants appear in the capitals of Ancient Egyptian columns which were carved to resemble either the Egyptian white lotus or the papyrus 121 Images of plants are often used in painting and photography as well as on textiles money stamps flags and coats of arms citation needed Scientific and cultural uses Barbara McClintock 1902 1992 was a pioneering cytogeneticist who used maize corn to study the mechanism of inheritance of traits 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 122 and examination of chromosomes in maize allowed Barbara McClintock to demonstrate their connection to inherited traits 123 The plant Arabidopsis thaliana is used in laboratories as a model organism to understand how genes control the growth and development of plant structures 124 NASA predicts that space stations or space colonies will one day rely on plants for life support 125 Ancient trees are revered and many are famous Tree rings themselves are an important method of dating in archeology and serve as a record of past climates 126 Plants figure prominently in mythology religion and literature 127 128 129 They are used as national and state emblems including 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 messages citation needed Negative effects The musk thistle is an invasive species in Texas Weeds are commercially or aesthetically undesirable plants growing in managed environments such as farms urban areas gardens lawns and parks People have spread plants beyond their native ranges and some of these introduced plants become invasive damaging existing ecosystems by displacing native species and sometimes becoming serious weeds of cultivation citation needed Plants may cause harm to animals including people Plants that produce windblown pollen invoke allergic reactions in people who suffer from hay fever A wide variety of plants are 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 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 130 131 Both illegal and legal drugs derived from plants may have negative effects on the economy affecting worker productivity and law enforcement costs 132 133 See also Plants portalBiosphere Evolutionary history of plants Plant defense against herbivory Plant identification Plant reproduction Plant to plant communication via mycorrhizal networks The Plant ListReferences Cavalier Smith T 1981 Eukaryote kingdoms Seven or nine BioSystems 14 3 4 461 481 doi 10 1016 0303 2647 81 90050 2 PMID 7337818 Lewis L A McCourt R M 2004 Green algae and the origin of land plants American Journal of Botany 91 10 1535 1556 doi 10 3732 ajb 91 10 1535 PMID 21652308 Kenrick Paul Crane Peter R 1997 The origin and early diversification of land plants A cladistic study Washington D C Smithsonian Institution Press ISBN 978 1 56098 730 7 Adl S M et al 2005 The new higher level classification of eukaryotes with emphasis on the taxonomy of protists The Journal of Eukaryotic Microbiology 52 5 399 451 doi 10 1111 j 1550 7408 2005 00053 x PMID 16248873 S2CID 8060916 a href Template Cite journal html title Template Cite journal cite journal a CS1 maint uses authors parameter link Numbers of threatened species by major groups of organisms 1996 2010 PDF International Union for Conservation of Nature 11 March 2010 Archived PDF from the original on 21 July 2011 Retrieved 27 April 2011 Field C B Behrenfeld M J Randerson J T Falkowski P 1998 Primary production of the biosphere Integrating terrestrial and oceanic components Science 281 5374 237 240 Bibcode 1998Sci 281 237F doi 10 1126 science 281 5374 237 PMID 9657713 Archived from the original on 25 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Smithsonian Institution Press ISBN 1 56098 730 8 Raven Peter H Evert Ray F amp Eichhorn Susan E 2005 Biology of Plants 7th ed New York W H Freeman and Company ISBN 0 7167 1007 2 Taylor Thomas N amp Taylor Edith L 1993 The Biology and Evolution of Fossil Plants Englewood Cliffs NJ Prentice Hall ISBN 0 13 651589 4 Trewavas A 2003 Aspects of Plant Intelligence Annals of Botany 92 1 1 20 doi 10 1093 aob mcg101 PMC 4243628 PMID 12740212 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 Prance G T 2001 Discovering the Plant World Taxon 50 2 Golden Jubilee Part 4 345 359 doi 10 2307 1223885 JSTOR 1223885 External links The Wikibook Dichotomous Key has a page on the topic of Plantae Jones T M Reid C S Urbatsch L E Visual study of divisional Plantae a href Template Cite journal html title Template Cite journal cite journal a Cite journal requires journal help requires Microsoft Silverlight Chaw S M et al 1997 Molecular Phylogeny of Extant Gymnosperms and Seed Plant Evolution Analysis of Nuclear 18s rRNA Sequences PDF Mol Biol Evol 14 1 56 68 doi 10 1093 oxfordjournals molbev a025702 PMID 9000754 Archived from the original PDF on 24 January 2005 Index Nominum Algarum Interactive Cronquist classification Plant Resources of Tropical Africa Tree of LifeBotanical and vegetation databasesAfrican Plants Initiative database Australia Chilean plants at Chilebosque e Floras Flora of China Flora of North America and others Flora Europaea Flora of Central Europe in German Flora of North America List of Japanese Wild Plants Online 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 Retrieved from https en wikipedia org w index php title Plant amp oldid 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