Precise numbers are difficult to determine, but as of 2010[update], there are thought to be 300–315 thousand species of plants, of which the great majority, some 260–290 thousand, are seed plants (see the table below). Green plants provide most of the world's molecular oxygen and are the basis of most of the earth's ecologies, especially on land. Plants described as grains, fruits and vegetables form mankind's basic foodstuffs, and have been domesticated for millennia. Plants serve as ornaments and, until recently and in great variety, they have served as the source of most medicines and drugs. Their scientific study is known as botany, a branch of biology.
Plants are one of the two groups into which all living things were traditionally divided; the other is animals. The division goes back at least as far as Aristotle (384 BC – 322 BC) who distinguished 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 often considered plants, particularly in popular contexts.
Outside of formal scientific contexts, the term "plant" implies an association with certain traits, such as being multicellular, possessing cellulose, and having the ability to carry out photosynthesis.
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:
This group includes the land plants plus various groups of green algae, including stoneworts. The names given to these groups vary considerably as of July 2011[update]. Viridiplantae encompass a group of organisms that possess chlorophylla and b, have plastids that are bound by only two membranes, are capable of storing starch, and have cellulose in their cell walls. It is this clade which is mainly the subject of this article.
Old classifications placed diverse algae, fungi and bacteria in Plantae (e.g., Plantae Haeckel 1866 included land plants, some algae and fungi).
Another way of looking at the relationships between the different groups which have been called "plants" is through a cladogram, which shows their evolutionary relationships. The evolutionary history of plants is not yet completely settled, but one accepted relationship between the three groups described above is shown below. Those which have been called "plants" are in bold.
Algae comprise several different groups of organisms which produce energy through photosynthesis and for that reason have been included in the plant kingdom in the past. Most conspicuous among the algae are the seaweeds, multicellular algae that may roughly resemble land plants, but are classified among the brown, red and green algae. Each of these algal groups also includes various microscopic and single-celled organisms. There is good evidence that some of these algal groups arose independently from separate non-photosynthetic ancestors, with the result that many groups of algae are no longer classified within the plant kingdom as it is defined here.
The Viridiplantae, the green plants – green algae and land plants – form a clade, a group consisting of all the descendants of a common ancestor. With a few exceptions among the green algae, all green plants have many features in common, including cell walls containing cellulose, chloroplasts containing chlorophyllsa and b, and food stores in the form of starch. 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 chloroplasts which appear to be derived directly from endosymbiotic cyanobacteria, although they differ in the pigments which are used in photosynthesis and so are different in colour. All three groups together are generally believed to have a single common origin, and so are classified together in the taxon Archaeplastida, whose name implies that the chloroplasts or plastids of all the members of the taxon were derived from a single ancient endosymbiotic event. This is the broadest modern definition of the plants.
In contrast, most other algae (e.g. heterokonts, 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, i.e. a group which 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 (or Charophyta).
The Chlorophyta (a name that has also been used for all green algae) are the sister group to the group from which the land plants evolved. There are about 4,300 species of mainly marine organisms, both unicellular and multicellular. The latter include the sea lettuce, Ulva.
The other group within the Viridiplantae are the mainly freshwater or terrestrial Streptophyta (or Charophyta), which consist of several groups of green algae plus the stoneworts and land plants. (The names have been used differently, e.g. Streptophyta to mean the group which excludes the land plants and Charophyta for the stoneworts alone or the stoneworts plus the land plants.) Streptophyte algae are either unicellular or form multicellular filaments, branched or unbranched. 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" which pond-owners so dislike. The freshwater stoneworts strongly resemble land plants and are believed to be their closest relatives. Growing underwater, they consist of a central stalk with whorls of branchlets, giving them a superficial resemblance to horsetails, species of the genus Equisetum, which are true land plants.
The classification of fungi has been controversial until quite recently in the history of biology. Linnaeus' original classification placed the fungi within the Plantae, since they were unquestionably not animals or minerals and these were the only other alternatives. With later developments in microbiology, in the 19th century Ernst Haeckel felt that another kingdom was required to classify newly discovered micro-organisms. The introduction of the new kingdom Protista in addition to Plantae and Animalia, led to uncertainty as to whether fungi truly were best placed in the Plantae or whether they ought to be reclassified as protists. Haeckel himself found it difficult to decide and it was not until 1969 that a solution was found whereby 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.
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 generally obtain carbon by breaking down and absorbing surrounding materials, and so are called heterotrophicsaprotrophs. 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 eukaryoticnuclei. Fruiting bodies, of which mushrooms are the most familiar example, are the reproductive structures of fungi, and are unlike any structures produced by plants.
The table below shows some species count estimates of different green plant (Viridiplantae) divisions. It suggests there are about 300,000 species of living Viridiplantae, of which 85–90% are flowering plants. (Note: as these are from different sources and different dates, they are not necessarily comparable, and like all species counts, are subject to a degree of uncertainty in some cases.)
Diversity of living green plant (Viridiplantae) divisions
Evidence suggests that 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. 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. These began to diversify in the late Silurian Period, around 420 million years ago, and the fruits 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. Late Devonian plants had thereby reached a degree of sophistication that allowed them to form forests of tall trees. Evolutionary innovation continued after the Devonian period. 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 CO
2 and warm, dry conditions of the tropics over the last 10 million years.
A proposed phylogenetic tree of Plantae, after Kenrick and Crane, is as follows, with modification to the Pteridophyta from Smith et al. The Prasinophyceae are a paraphyletic assemblage of early diverging green algal lineages.
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. Plants 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 can only survive where moisture is available for significant periods, although some species 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 bryophytes have the ability to secrete a cuticle on their outer surface, a waxy layer that confers resistant 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 while controlling water loss.
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.
The first seed plants, Pteridosperms (seed ferns), now extinct, appeared in the Devonian and diversified through the Carboniferous. In these the microgametophyte is reduced to pollen and the megagametophyte remains inside the megasporangium, attached to the parent plant. A megasporangium invested in protective layer called an integument is known as an ovule. After fertilisation by means of sperm deposited by pollen grains, an embryo 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.
Early seed plants are gymnosperms, as the ovules and subsequent seeds are not enclosed in a protective structure (carpels or fruit), but are found naked, typically on cone scales. Pollen typically lands directly on the ovule. Four surviving groups remain widespread now, particularly the conifers, which are dominant trees in several biomes.
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 calcifiedmulticellular members of the Dasycladales. Earlier Precambrian fossils are known which resemble single-cell green algae, but definitive identity with that group of algae is uncertain.
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 lycophyteBaragwanathia longifolia. From the Devonian, detailed fossils of rhyniophytes have been found. Early fossils of these ancient plants show the individual cells within the plant tissue. The Devonian period also saw the evolution of what many believe to be the first modern tree, Archaeopteris. This fern-like tree combined a woody trunk with the fronds of a fern, but produced no seeds.
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 Forest at Victoria Park in Glasgow, Scotland, the stumps of Lepidodendron trees are found in their original growth positions.
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.
The earliest fossils attributed to green algae date from the Precambrian (ca. 1200 mya). 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.
Most of the solid material in a plant is taken from the atmosphere. Through a process known as photosynthesis, most plants use the energy in sunlight to convert carbon dioxide from the atmosphere, plus water, into simple sugars. (Parasitic plants, on the other hand, use the resources of its host to grow.) 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.
Plants usually rely on soil primarily for support and water (in quantitative terms), but also obtain compounds of nitrogen, phosphorus, potassium, magnesium and other elemental nutrients. Epiphytic and lithophytic plants depend on air and nearby debris for nutrients, and carnivorous plants supplement their nutrient requirements 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. Some plants grow as submerged aquatics, using oxygen dissolved in the surrounding water, and a few specialized vascular plants, such as mangroves, can grow with their roots in anoxic conditions.
There is no photosynthesis in deciduous leaves in autumn.
Factors affecting growth
The genotype of a plant affects its growth. For example, selected varieties of wheat grow rapidly, maturing within 110 days, whereas others, in the same environmental conditions, grow more slowly and mature within 155 days.
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.
Biotic factors are also capable of affecting plant growth. Plants compete with other plants for space, water, light and nutrients. 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.
Simple plants like algae may have short life spans as individuals, but their populations are commonly seasonal. Other plants may be organized according to their seasonal growth pattern: annual plants live and reproduce within one growing season, biennial plants live for two growing seasons and usually reproduce in second year, and perennial plants live for many growing seasons and continue to reproduce once they are mature. 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 which lose their leaves for some part of it. In temperate and boreal climates, they generally lose their leaves during the winter; many tropical plants lose their leaves during the dry season.
The growth rate of plants is extremely variable. Some mosses grow less than 0.001 millimeters per hour (mm/h), while most trees grow 0.025-0.250 mm/h. Some climbing species, such as kudzu, which do not need to produce thick supportive tissue, may grow up to 12.5 mm/h.
Plants are photosynthetic, 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 pigmentchlorophyll. 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.
By means of cells that behave like nerves, plants receive and distribute within their systems information about incident light intensity and quality. Incident light which stimulates a chemical reaction in one leaf, will cause a chain reaction of signals to the entire plant via a type of cell termed a bundle sheath cell. Researchers from the Warsaw University of Life Sciences in Poland, found that plants have a specific memory for varying light conditions which prepares their immune systems against seasonal pathogens. Plants use pattern-recognition receptors to recognize conserved microbial signatures. This recognition triggers an immune response. The first plant receptors of conserved microbial signatures were identified in rice (XA21, 1995) and in Arabidopsis thaliana (FLS2, 2000). Plants also carry immune receptors that recognize highly variable pathogen effectors. These include the NBS-LRR class of proteins.
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.
The photosynthesis conducted by land plants and algae is the ultimate source of energy and organic material in nearly all ecosystems. Photosynthesis radically changed the composition of the early Earth's 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.
Plants are distributed worldwide in varying numbers. 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 have adapted tenaciously to the prevailing conditions. (See Antarctic flora.)
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 and forests.
Numerous animals have coevolved with plants. Many animals pollinateflowers 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 mutualisticsymbiosis 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.
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, complex aquatic food webs.
A section of a Yew branch showing 27 annual growth rings, pale sapwood and dark heartwood, and pith (centre dark spot). The dark radial lines are longitudinal sections of small branches which became included by growth of the tree.
The study of plant uses by people is termed economic botany or ethnobotany; some consider economic botany to focus on modern cultivated plants, while ethnobotany focuses on indigenous plants cultivated and used by native peoples. Human cultivation of plants is part of agriculture, which is the basis of human civilization. Plant agriculture is subdivided into agronomy, horticulture and forestry.
Tree rings are an important method of dating in archeology and serve as a record of past climates. Basic biological research has often been done with plants, such as the pea plants used to derive Gregor Mendel's laws of genetics. Space stations or space colonies may one day rely on plants for life support. Plants are used as national and state emblems, including state trees and state flowers. Ancient trees are revered and many are famous. Numerous world records are held by plants. Plants are often used as memorials, gifts and to mark special occasions such as births, deaths, weddings and holidays. Plants figure prominently in mythology, religion and literature. The field of ethnobotany studies plant use by indigenous cultures which helps to conserve endangered species as well as discover new medicinal plants. Gardening is the most popular leisure activity in the U.S. Working with plants or horticulture therapy is beneficial for rehabilitating people with disabilities. Certain plants contain psychotropic chemicals which are extracted and ingested, including tobacco, cannabis (marijuana), and opium.
Weeds are uncultivated and usually unwanted plants by gardeners. People have spread plants beyond their native ranges and some of these introduced plants become invasive, damaging existing ecosystems by displacing native species. Invasive plants cause costly damage in crop losses annually by displacing crop plants, they further increase the cost of production and the use of chemicals to control them, which in turn affects the environment.
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 psychotropicchemicals, which are extracted and ingested or smoked, including tobacco, cannabis (marijuana), cocaine and opium. Smoking causes damage to health or even death, while some drugs may also be harmful or fatal to people. Both illegal and legal drugs derived from plants may have negative effects on the economy, affecting worker productivity and law enforcement costs. Some plants cause allergic reactions when ingested, while other plants cause food intolerances that negatively affect health.
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