Plant Life At Castlewood Lake
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Plants are predominantly
photosynthetic Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in c ...
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
s of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
and fungi; however, all current definitions of Plantae exclude the fungi and some algae, as well as the prokaryotes (the
archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
and bacteria). By one definition, plants form the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Viridiplantae (Latin name for "green plants") which is sister of the Glaucophyta, and consists of the
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
and Embryophyta (land plants). The latter includes the flowering plants,
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s and other gymnosperms, ferns and their allies,
hornwort Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s,
liverworts The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ge ...
, and
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es. Most plants are multicellular organisms. Green plants obtain most of their energy from
sunlight Sunlight is a portion of the electromagnetic radiation given off by the Sun, in particular infrared, visible, and ultraviolet light. On Earth, sunlight is scattered and filtered through Earth's atmosphere, and is obvious as daylight when t ...
via photosynthesis by primary
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s that are derived from endosymbiosis with
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
. Their chloroplasts contain
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
s a and b, which gives them their green color. Some plants are parasitic or
mycotroph A mycotroph is a plant that gets all or part of its carbon, water, or nutrient supply through symbiotic association with fungi. The term can refer to plants that engage in either of two distinct symbioses with fungi: *Many mycotrophs have a mut ...
ic 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 Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of Biological life cycle, life cycle in plants and algae. It consists of a Multicellular organism, multicellular haploid sexual phase, the gametophy ...
, although
asexual reproduction Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the fu ...
is also common. There are about 320,000 known species of plants, of which the great majority, some 260,000–290,000, produce seeds. Green plants provide a substantial proportion of the world's molecular oxygen, and are the basis of most of Earth's ecosystems. Plants that produce
grain A grain is a small, hard, dry fruit (caryopsis) – with or without an attached hull layer – harvested for human or animal consumption. A grain crop is a grain-producing plant. The two main types of commercial grain crops are cereals and legum ...
, fruit, and vegetables also form basic human foods and have been domesticated for millennia. Plants have many
cultural Culture () is an umbrella term which encompasses the social behavior, institutions, and Social norm, norms found in human Society, societies, as well as the knowledge, beliefs, arts, laws, Social norm, customs, capabilities, and habits of the ...
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 BC – 322 BC), who made the distinction between plants, which generally do not move, and animals, which often are mobile to catch their food. Much later, when Linnaeus (1707–1778) created the basis of the modern system of scientific classification, these two groups became the
kingdoms Kingdom commonly refers to: * A monarchy ruled by a king or queen * Kingdom (biology), a category in biological taxonomy Kingdom may also refer to: Arts and media Television * ''Kingdom'' (British TV series), a 2007 British television drama s ...
Vegetabilia (later Metaphyta or Plantae) and
Animalia Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
(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 Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
were removed to new kingdoms. However, these organisms are still sometimes considered plants, particularly in informal contexts. The term "plant" generally implies the possession of the following traits: multicellularity, possession of cell walls containing cellulose, and the ability to carry out photosynthesis with primary chloroplasts.– Definition from the Merriam-Webster Online Dictionary">


Current definitions of Plantae

When the name Plantae or plant is applied to a specific group of organisms or taxon, it usually refers to one of four concepts. From least to most inclusive, these four groupings are: Another way of looking at the relationships between the different groups that have been called "plants" is through a cladogram, which shows their evolutionary relationships. These are not yet completely settled, but . Those which have been called "plants" are in bold (some minor groups have been omitted). The way in which the groups of green algae are combined and named varies considerably between authors.


Algae

Algae consist of several groups of organisms which produce food by photosynthesis and thus have traditionally been included in the plant kingdom. The
seaweed Seaweed, or macroalgae, refers to thousands of species of macroscopic, multicellular, marine algae. The term includes some types of '' Rhodophyta'' (red), ''Phaeophyta'' (brown) and ''Chlorophyta'' (green) macroalgae. Seaweed species such as ...
s range from large multicellular algae to single-celled organisms and are classified into three groups, the
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
,
red algae Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
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. The Viridiplantae, the green plants – green algae and land plants – form a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, a group consisting of all the descendants of a common ancestor. With a few exceptions, the green plants have the following features in common; primary
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s derived from cyanobacteria containing
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
s ''a'' and ''b'', cell walls containing cellulose, and food stores in the form of
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
contained within the plastids. They undergo closed
mitosis In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
without
centriole In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and a ...
s, and typically have
mitochondria A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
with flat cristae. The
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s of green plants are surrounded by two membranes, suggesting they originated directly from endosymbiotic
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
. Two additional groups, the
Rhodophyta Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
(red algae) and Glaucophyta (glaucophyte algae), also have primary chloroplasts that appear to be derived directly from endosymbiotic
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
, 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,
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s, 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 In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
group, i.e. a group that excludes some of the descendants of a common ancestor. Paraphyletic groups are generally avoided in modern classifications, so that in recent treatments the Viridiplantae have been divided into two clades, the Chlorophyta and the Streptophyta (including the land plants and Charophyta). The Chlorophyta (a name that has also been used for ''all'' green algae) are the sister group to the Charophytes, from which the land plants evolved. There are about 4,300 species, mainly unicellular or multicellular marine organisms such as the sea lettuce, ''
Ulva Ulva (; gd, Ulbha) is a small island in the Inner Hebrides of Scotland, off the west coast of Mull. It is separated from Mull by a narrow strait, and connected to the neighbouring island of Gometra by a bridge. Much of the island is formed from ...
''. 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. 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. Growing immersed in fresh water, they consist of a central stalk with whorls of branchlets.


Fungi

Linnaeus' original classification placed the fungi within the Plantae, since they were unquestionably neither animals or minerals and these were the only other alternatives. With 19th century developments in
microbiology Microbiology () is the scientific study of microorganisms, those being unicellular (single cell), multicellular (cell colony), or acellular (lacking cells). Microbiology encompasses numerous sub-disciplines including virology, bacteriology, prot ...
,
Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new sp ...
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 Robert Whitaker or Whittaker may refer to: *Robert Whittaker (fighter) (born 1990), Australian mixed martial artist *Robert Whitaker (equestrian) (born 1983), British showjumper *Robert Whitaker (author) (active since 1989), American author *Robert ...
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 do not possess chloroplasts and generally obtain carbon by breaking down and absorbing surrounding materials, and so are called heterotrophic
saprotrophs Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi (f ...
. In addition, the substructure of multicellular fungi is different from that of plants, taking the form of many chitinous microscopic strands called
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e, 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.


Diversity

The table below shows some species count estimates of different green plant (Viridiplantae) divisions. About 85–90% of all plants are flowering plants. Several projects are currently attempting to collect all plant species in online databases, e.g. the World Flora Online and
World Plants In its most general sense, the term "world" refers to the totality of entities, to the whole of reality or to everything that is. The nature of the world has been conceptualized differently in different fields. Some conceptions see the worl ...
both list about 391,000 species. The naming of plants is governed by the
International Code of Nomenclature for algae, fungi, and plants The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all those "trad ...
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
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s of today. Plants in all of these groups continue to thrive, especially in the environments in which they evolved. An algal scum formed on the land , but it was not until the Ordovician Period, around , that land plants appeared. However, new evidence from the study of carbon isotope ratios in Precambrian rocks has suggested that complex photosynthetic plants developed on the earth over 1000 m.y.a. For more than a century it has been assumed that the ancestors of land plants evolved in aquatic environments and then adapted to a life on land, an idea usually credited to botanist Frederick Orpen Bower in his 1908 book ''The Origin of a Land Flora''. A recent alternative view, supported by genetic evidence, is that they evolved from terrestrial single-celled algae, and that even the common ancestor of red and green algae, and the unicellular freshwater algae glaucophytes, originated in a terrestrial environment in freshwater biofilms or microbial mats. Primitive land plants began to diversify in the late Silurian Period, around , and the results of their diversification are displayed in remarkable detail in an early
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
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 in the Carboniferous and later geological periods and is ongoing today. Most plant groups were relatively unscathed by the Permo-Triassic extinction event, although the structures of communities changed. This may have set the scene for the evolution of flowering plants in the Triassic (~), which exploded in the Cretaceous and Tertiary. The latest major group of plants to evolve were the grasses, which became important in the mid Tertiary, from around . The grasses, as well as many other groups, evolved new mechanisms of metabolism to survive the low and warm, dry conditions of the tropics over the last . A 1997 proposed
phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
of Plantae, after Kenrick and Crane, is as follows, with modification to the Pteridophyta from Smith ''et al.'' The
Prasinophyceae The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapman ...
are a
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
assemblage of early diverging green algal lineages, but are treated as a group outside the Chlorophyta: later authors have not followed this suggestion. A newer proposed classification follows Leliaert et al. 2011 and modified with Silar 2016 for the green algae clades and Novíkov & Barabaš-Krasni 2015 for the land plants clade. Notice that the Prasinophyceae are here placed inside the Chlorophyta. Later, a phylogeny based on genomes and transcriptomes from 1,153 plant species was proposed. The placing of algal groups is supported by phylogenies based on genomes from the Mesostigmatophyceae and Chlorokybophyceae that have since been sequenced. The classification of Bryophyta is supported both by Puttick ''et al.'' 2018, and by phylogenies involving the hornwort genomes that have also since been sequenced.


Embryophytes

The plants that are likely most familiar to us are the multicellular land plants, called embryophytes. Embryophytes include the vascular plants, such as ferns, conifers and flowering plants. They also include the '' bryophytes'', of which
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es and
liverworts The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of ge ...
are the most common. All of these plants have eukaryotic cells with
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
s 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 The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as ...
, 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 Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
stage, called the gametophyte, and a
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
stage, called the
sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote pr ...
. In bryophytes, the sporophyte is always unbranched and remains nutritionally dependent on its parent gametophyte. The embryophytes have the ability to secrete a
cuticle A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
on their outer surface, a waxy layer that confers resistance to desiccation. In the
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es and
hornwort Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s a cuticle is usually only produced on the sporophyte.
Stomata In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
are absent from liverworts, but occur on the sporangia of mosses and hornworts, allowing gas exchange. Vascular plants first appeared during the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
period, and by the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
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 A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sex ...
is reduced from a multicellular free-living organism to a few cells in a pollen grain and the miniaturised
megagametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the ...
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 Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
grains, an embryo sporophyte develops inside the ovule. The integument becomes a seed coat, and the ovule develops into a seed. Seed plants can survive and reproduce in extremely arid conditions, because they are not dependent on free water for the movement of sperm, or the development of free living gametophytes. The first seed plants,
pteridosperms The term Pteridospermatophyta (or "seed ferns" or "Pteridospermatopsida") is a polyphyletic group of extinct seed-bearing plants (spermatophytes). The earliest fossil evidence for plants of this type is the genus ''Elkinsia'' of the late Devonian ...
(seed ferns), now extinct, appeared in the Devonian and diversified through the Carboniferous. They were the ancestors of modern gymnosperms, of which four surviving groups are widespread today, particularly the
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
s, which are dominant trees in several biomes. The name gymnosperm comes from the Greek , a composite of ( ) and ( ), as the ovules and subsequent seeds are not enclosed in a protective structure (carpels or fruit), but are borne naked, typically on cone scales.


Fossils

Plant fossils include roots, wood, leaves, seeds, fruit,
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
,
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s, phytoliths, and amber (the fossilized resin produced by some plants). Fossil land plants are recorded in terrestrial, lacustrine, fluvial and nearshore marine sediments.
Pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophyt ...
, spores and algae (
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s 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 The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
. These fossils resemble calcified multicellular members of the Dasycladales. Earlier
Precambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of the ...
fossils are known that resemble single-cell green algae, but definitive identity with that group of algae is uncertain. The earliest fossils attributed to green algae date from the
Precambrian The Precambrian (or Pre-Cambrian, sometimes abbreviated pꞒ, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of the ...
(ca. 1200 mya). The resistant outer walls of
prasinophyte The prasinophytes are a group of unicellular green algae. Prasinophytes mainly include marine planktonic species, as well as some freshwater representatives.Sym, S. D. and Pienaar, R. N. 1993. The class Prasinophyceae. In Round, F. E. and Chapman ...
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. The oldest known fossils of embryophytes date from the Ordovician, though such fossils are fragmentary. By the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
, 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 ''Baragwanathia'' is a genus of extinct lycopsid plants of Late Silurian to Early Devonian age (), fossils of which have been found in Australia, Canada, China and Czechia. The name derives from William Baragwanath who discovered the first spec ...
'' in late Silurian. From the early Devonian Rhynie chert, 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 Glomerales is an order of symbiotic fungi within the phylum Glomeromycota. Biology These fungi are all biotrophic mutualists. Most employ the arbuscular mycorrhizal method of nutrient exchange with plants. They produce large (.1-.5mm) spores ...
. The Devonian period also saw the evolution of leaves and roots, and the first modern tree, ''
Archaeopteris ''Archaeopteris'' is an extinct genus of progymnosperm tree with fern-like leaves. A useful index fossil, this tree is found in strata dating from the Upper Devonian to Lower Carboniferous (), the oldest fossils being 385 million years old, ...
''. 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. 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 The Fossil Grove is a group of plant fossils located within Victoria Park, Glasgow, Scotland. It was discovered in 1887 and contains the fossilised stumps and roots of eleven extinct ''Lepidodendron'' trees, which are sometimes described as " ...
at
Victoria Park Victoria Park may refer to: Places Australia * Victoria Park Nature Reserve, a protected area in Northern Rivers region, New South Wales * Victoria Park, Adelaide, a park and racecourse * Victoria Park, Brisbane, a public park and former golf ...
in Glasgow, Scotland, the stumps of '' Lepidodendron'' trees are found in their original growth positions. The fossilized remains of
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
and
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
roots, stems and branches may be locally abundant in lake and inshore sedimentary rocks from the Mesozoic and
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
eras. Sequoia and its allies, magnolia, oak, and palms are often found. 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

Most of the solid material in a plant is taken from the
atmosphere An atmosphere () is a layer of gas or layers of gases that envelop a planet, and is held in place by the gravity of the planetary body. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A s ...
. Through the process of photosynthesis, most plants use the energy in
sunlight Sunlight is a portion of the electromagnetic radiation given off by the Sun, in particular infrared, visible, and ultraviolet light. On Earth, sunlight is scattered and filtered through Earth's atmosphere, and is obvious as daylight when t ...
to convert carbon dioxide from the atmosphere, plus water, into simple
sugar Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double ...
s. These sugars are then used as building blocks and form the main structural component of the plant.
Chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
, a green-colored, magnesium-containing pigment is essential to this process; it is generally present in plant
leaves A leaf (plural, : leaves) is any of the principal appendages of a vascular plant plant stem, stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", wh ...
, 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
nutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...
s from the soil.
Epiphytic An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
and
lithophytic Lithophytes are plants that grow in or on rocks. They can be classified as either epilithic (or epipetric) or endolithic; epilithic lithophytes grow on the surfaces of rocks, while endolithic lithophytes grow in the crevices of rocks (and are als ...
plants depend on air and nearby debris for nutrients, and
carnivorous plant Carnivorous plants are plants that derive some or most of their nutrients from trapping and consuming animals or protozoans Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryot ...
s supplement their nutrient requirements, particularly for nitrogen and phosphorus, with insect prey that they capture. For the majority of plants to grow successfully they also require oxygen in the atmosphere and around their roots (
soil gas Soil gases (soil atmosphere) are the gases found in the air space between soil components. The spaces between the solid soil particles, if they do not contain water, are filled with air. The primary soil gases are nitrogen, carbon dioxide and oxygen ...
) for respiration. Plants use oxygen and glucose (which may be produced from stored
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
) to provide energy. Some plants grow as submerged aquatics, using oxygen dissolved in the surrounding water, and a few specialized vascular plants, such as mangroves and reed ('' Phragmites australis''), can grow with their roots in anoxic conditions.


Factors affecting growth

The genome of a plant controls its growth. For example, selected varieties or genotypes of wheat grow rapidly, maturing within 110 days, whereas others, in the same environmental conditions, grow more slowly and mature within 155 days.Robbins, W.W.; Weier, T.E.; ''et al.''., ''Botany: Plant Science'', 3rd edition, Wiley International, New York, 1965. Growth is also determined by environmental factors, such as temperature, available water, available light, carbon dioxide and available
nutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...
s in the soil. Any change in the availability of these external conditions will be reflected in the plant's growth and the timing of its development. Biotic factors also affect plant growth. Plants can be so crowded that no single individual produces normal growth, causing etiolation and chlorosis. Optimal plant growth can be hampered by grazing animals, suboptimal soil composition, lack of
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
l 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.
Annual plant An annual plant is a plant that completes its life cycle, from germination to the production of seeds, within one growing season, and then dies. The length of growing seasons and period in which they take place vary according to geographical ...
s grow and reproduce within one
growing season A season is a division of the year marked by changes in weather, ecology, and the amount of daylight. The growing season is that portion of the year in which local conditions (i.e. rainfall, temperature, daylight) permit normal plant growth. Whil ...
,
biennial plant A biennial plant is a flowering plant that, generally in a temperate climate, takes two years to complete its biological life cycle. Life cycle In its first year, the biennal plant undergoes primary growth, during which its vegetative structures ...
s 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 Alpine may refer to any mountainous region. It may also refer to: Places Europe * Alps, a European mountain range ** Alpine states, which overlap with the European range Australia * Alpine, New South Wales, a Northern Village * Alpine National Pa ...
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 climate The subarctic climate (also called subpolar climate, or boreal climate) is a climate with long, cold (often very cold) winters, and short, warm to cool summers. It is found on large landmasses, often away from the moderating effects of an ocean, ge ...
s, 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 Kudzu (; also called Japanese arrowroot or Chinese arrowroot) is a group of climbing, coiling, and trailing deciduous perennial vines native to much of East Asia, Southeast Asia, and some Pacific islands, but invasive species, invasive in many ...
, which do not need to produce thick supportive tissue, may grow up to 12.5 mm/h. Plants protect themselves from
frost Frost is a thin layer of ice on a solid surface, which forms from water vapor in an above-freezing atmosphere coming in contact with a solid surface whose temperature is below freezing, and resulting in a phase change from water vapor (a gas) ...
and dehydration stress with antifreeze proteins,
heat-shock proteins Heat shock proteins (HSP) are a family of proteins produced by cells in response to exposure to stressful conditions. They were first described in relation to heat shock, but are now known to also be expressed during other stresses including expo ...
and sugars (
sucrose Sucrose, a disaccharide, is a sugar composed of glucose and fructose subunits. It is produced naturally in plants and is the main constituent of white sugar. It has the molecular formula . For human consumption, sucrose is extracted and refined ...
is common). LEA ( Late Embryogenesis Abundant) protein expression is induced by stresses and protects other proteins from aggregation as a result of
desiccation Desiccation () is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. ...
and freezing.


Effects of freezing

When water freezes in plants, the consequences for the plant depend very much on whether the freezing occurs within cells (intracellularly) or outside cells in intercellular spaces.Glerum, C. 1985. Frost hardiness of coniferous seedlings: principles and applications. pp. 107–123 ''in ''Duryea, M.L. (Ed.). Proceedings: Evaluating seedling quality: principles, procedures, and predictive abilities of major tests. Workshop, October 1984, Oregon State Univ., For. Res. Lab., Corvallis OR. Intracellular freezing, which usually kills the cellLyons, J.M.; Raison, J.K.; Steponkus, P.L. 1979. The plant membrane in response to low temperature: an overview. pp. 1–24 ''in'' Lyons, J.M.; Graham, D.; Raison, J.K. (Eds.). Low Temperature Stress in Crop Plants. Academic Press, New York NY. regardless of the hardiness of the plant and its tissues, seldom occurs in nature because rates of cooling are rarely high enough to support it. Rates of cooling of several degrees Celsius per minute are typically needed to cause intracellular formation of ice.Mazur, P. 1977. The role of intracellular freezing in the death of cells cooled at supraoptimal rates. Cryobiology 14:251–272. At rates of cooling of a few degrees Celsius per hour, segregation of ice occurs in intercellular spaces.Sakai, A.; Larcher, W. (Eds.) 1987. Frost Survival of Plants. Springer-Verlag, New York. 321 p. This may or may not be lethal, depending on the hardiness of the tissue. At freezing temperatures, water in the intercellular spaces of plant tissue freezes first, though the water may remain unfrozen until temperatures drop below . After the initial formation of intercellular ice, the cells shrink as water is lost to the segregated ice, and the cells undergo freeze-drying. This dehydration is now considered the fundamental cause of freezing injury.


DNA damage and repair

Plants are continuously exposed to a range of biotic and abiotic stresses. These stresses often cause
DNA damage DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA da ...
directly, or indirectly via the generation of reactive oxygen species. Plants are capable of a DNA damage response that is a critical mechanism for maintaining genome stability. The DNA damage response is particularly important during seed
germination Germination is the process by which an organism grows from a seed or spore. The term is applied to the sprouting of a seedling from a seed of an angiosperm or gymnosperm, the growth of a sporeling from a spore, such as the spores of fungi, fer ...
, since seed quality tends to deteriorate with age in association with DNA damage accumulation. During germination repair processes are activated to deal with this accumulated DNA damage. In particular, single- and
double A double is a look-alike or doppelgänger; one person or being that resembles another. Double, The Double or Dubble may also refer to: Film and television * Double (filmmaking), someone who substitutes for the credited actor of a character * Th ...
-strand breaks in DNA can be repaired. Double-strand repair in plants often produce DNA junctions with structural alterations. 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.


Plant cells

Plant cells are typically distinguished by their large water-filled central vacuole,
chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in ...
s, and rigid
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering mech ...
s that are made up of cellulose, hemicellulose, and
pectin Pectin ( grc, πηκτικός ': "congealed" and "curdled") is a heteropolysaccharide, a structural acid contained in the primary lamella, in the middle lamella, and in the cell walls of terrestrial plants. The principal, chemical component of ...
. Cell division is also characterized by the development of a
phragmoplast image:Phragmoplast.png, 300px, Phragmoplast and cell plate formation in a plant cell during cytokinesis. Left side: Phragmoplast forms and cell plate starts to assemble in the center of the cell. Towards the right: Phragmoplast enlarges in a donut- ...
for the construction of a
cell plate image:Phragmoplast.png, 300px, Phragmoplast and cell plate formation in a plant cell during cytokinesis. Left side: Phragmoplast forms and cell plate starts to assemble in the center of the cell. Towards the right: Phragmoplast enlarges in a donut ...
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 The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are als ...
tissues, with more primitive plants lacking some tissue types.


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 Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
. 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: :6CO2 + 6H2O -> textC6H12O6 + 6O2


Immune system

By means of cells that behave like nerves, plants receive and distribute within their systems information about incident light intensity and quality. Incident light that stimulates a chemical reaction in one leaf, will cause a chain reaction of signals to the entire plant via a type of cell termed a ''
bundle sheath cell A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will inclu ...
''. 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 ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A winter a ...
'' (FLS2, 2000). Plants also carry immune receptors that recognize highly variable pathogen effectors. These include the NBS-LRR class of proteins.


Internal distribution

Vascular plants differ from other plants in that nutrients are transported between their different parts through specialized structures, called xylem 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.


Genomics

Plants have some of the largest genomes among all organisms. The largest plant genome (in terms of gene number) is that of wheat (''Triticum asestivum''), predicted to encode ≈94,000 genes and thus almost 5 times as many as the human genome. The first plant genome sequenced was that of ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A winter a ...
'' which encodes about 25,500 genes. 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) while the largest, from the
Norway Spruce ''Picea abies'', the Norway spruce or European spruce, is a species of spruce native to Northern, Central and Eastern Europe. It has branchlets that typically hang downwards, and the largest cones of any spruce, 9–17 cm long. It is very close ...
(''Picea abies''), extends over 19,600 Mb (encoding about 28,300 genes).


Ecology

The photosynthesis conducted by land plants and algae is the ultimate source of energy and organic material in nearly all ecosystems. Photosynthesis, at first by cyanobacteria and later by photosynthetic eukaryotes, radically changed the composition of the early Earth's anoxic atmosphere, which as a result is now 21% oxygen. Animals and most other organisms are aerobic, relying on oxygen; those that do not are confined to relatively rare
anaerobic environment Hypoxia refers to low oxygen conditions. Normally, 20.9% of the gas in the atmosphere is oxygen. The partial pressure of oxygen in the atmosphere is 20.9% of the total barometric pressure. In water, oxygen levels are much lower, approximately 7 p ...
s. 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 form about 80% of the world
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bi ...
at about of carbon. Land plants are key components of the water cycle and several other
biogeochemical cycle A biogeochemical cycle (or more generally a cycle of matter) is the pathway by which a chemical substance cycles (is turned over or moves through) the biotic and the abiotic compartments of Earth. The biotic compartment is the biosphere and the ...
s. Some plants have
coevolve In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
d with
nitrogen fixing Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the Atmosphere of Earth, air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but al ...
bacteria, making plants an important part of the
nitrogen cycle The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmospheric, terrestrial, and marine ecosystems. The conversion of nitrogen can be carried out through both biologi ...
. Plant roots play an essential role in soil development and the prevention of
soil erosion Soil erosion is the denudation or wearing away of the upper layer of soil. It is a form of soil degradation. This natural process is caused by the dynamic activity of erosive agents, that is, water, ice (glaciers), snow, air (wind), plants, and ...
.


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 A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
. At the southern extremes, plants of the Antarctic flora have adapted tenaciously to the prevailing conditions. Plants are often the dominant physical and structural component of habitats where they occur. Many of the Earth's biomes are named for the type of vegetation because plants are the dominant organisms in those biomes, such as grasslands, taiga and tropical rainforest.


Ecological relationships

Numerous animals have coevolved with plants. Many animals pollinate flowers in exchange for food in the form of pollen or
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
. Many animals disperse seeds, often by eating fruit and passing the seeds in their
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
. 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
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s 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 Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
known as
mycorrhiza   A mycorrhiza (from Greek μύκης ', "fungus", and ῥίζα ', "root"; pl. mycorrhizae, mycorrhiza or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant ...
. 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 An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; h ...
fungi that protect the plant from herbivores by producing toxins. The fungal endophyte, '' Neotyphodium coenophialum'', in tall fescue (''Festuca arundinacea'') does tremendous economic damage to the cattle industry in the U.S. Many legume plants have nitrogen fixing bacteria in the genus '' Rhizobium'', found in nodules of their roots, that fix nitrogen from the air for the plant to use. In exchange, the plants supply sugars to the bacteria. Various forms of parasitism are also fairly common among plants, from the semi-parasitic
mistletoe Mistletoe is the common name for obligate hemiparasitic plants in the order Santalales. They are attached to their host tree or shrub by a structure called the haustorium, through which they extract water and nutrients from the host plant. ...
that merely takes some nutrients from its host, but still has photosynthetic leaves, to the fully parasitic
broomrape ''Orobanche'', commonly known as broomrape, is a genus of over 200 species of small Parasitic plant, parasitic herbaceous plants, mostly native to the temperate climate, temperate Northern Hemisphere. It is the type genus of the Orobanchaceae, ...
and
toothwort Toothwort is a common name for several plants and may refer to: *''Cardamine'', a genus of plants in the mustard family, Brassicaceae *''Lathraea'', a genus of parasitic plants in the family Orobanchaceae Orobanchaceae, the broomrapes, is a fam ...
that acquire all their nutrients through connections to the roots of other plants, and so have no
chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to a ...
. Some plants, known as myco-heterotrophs, parasitize mycorrhizal fungi, and hence act as
epiparasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
s on other plants. Many plants are
epiphyte An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
s, 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.
Hemiepiphyte A hemiepiphyte is a plant that spends part of its life cycle as an epiphyte. The seeds of primary hemiepiphytes germinate in the canopy and initially live epiphytically. They send roots downward, and these roots eventually make contact with t ...
s like the
strangler fig Strangler fig is the common name for a number of tropical and subtropical plant species in the genus ''Ficus'', including those that are commonly known as banyans. Some of the more well-known species are: * ''Ficus altissima'' * ''Ficus aurea'', ...
begin as epiphytes but eventually set their own roots and overpower and kill their host. Many orchids,
bromeliad The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, ...
s, ferns and
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es often grow as epiphytes. Bromeliad epiphytes accumulate water in leaf axils to form phytotelmata that may contain complex aquatic food webs. Approximately 630 plants are
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
, such as the Venus Flytrap (''Dionaea muscipula'') and sundew (''Drosera'' species). They trap small animals and digest them to obtain mineral nutrients, especially nitrogen and phosphorus.Barthlott, W.; Porembski, S.; Seine, R.; Theisen, I. 2007. ''The Curious World of Carnivorous Plants: A Comprehensive Guide to Their Biology and Cultivation.'' Timber Press: Portland, Oregon.


Competition

Competition occurs when members of the same species, or several different species, compete for shared resources in a given habitat. According to the competitive exclusion principle, when environmental resources are limited, species cannot occupy nor be supported by identical niches. Eventually, one species will out-compete the other, which will push the disadvantaged species to extinction. In regard to plants, competition tends to negatively affect their growth when competing for shared resources. These shared resources commonly include space for growth, sunlight, water and nutrients. Light is an important resource because it is necessary for photosynthesis. Plants use their leaves to shade other plants from sunlight and grow quickly to maximize their own expose. Water is also important for photosynthesis, and plants have different root systems to maximize water uptake from soil. Some plants have deep roots that are able to locate water stored deep underground, and others have shallower roots that are capable of extending longer distances to collect recent rainwater. Minerals are also important for plant growth and development, where deficiencies can occur if nutrient needs are not met. Common nutrients competed for amongst plants include nitrogen and phosphorus. Space is also extremely important for a growing and developing plant. Having optimal space makes it more likely that leaves are exposed to sufficient amounts of sunlight and are not overcrowded in order for photosynthesis to occur. If an old tree dies, then competition arises amongst a number of trees to replace it. Those that are less effective competitors are less likely to contribute to the next generation of offspring. Contrary to the belief that plants are always in competition, new research has found that in a harsh environment mature plants sheltering seedlings help the smaller plant survive.


Importance


Cultivation

The study of plant uses by people is called economic botany or
ethnobotany Ethnobotany is the study of a region's plants and their practical uses through the traditional knowledge of a local culture and people. An ethnobotanist thus strives to document the local customs involving the practical uses of local flora for m ...
. Human cultivation of plants is part of agriculture, which is the basis of human civilization. ''
Plant agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people ...
'' is subdivided into
agronomy Agronomy is the science and technology of producing and using plants by agriculture for food, fuel, fiber, chemicals, recreation, or land conservation. Agronomy has come to include research of plant genetics, plant physiology, meteorology, and ...
, horticulture and forestry.


Food

Humans depend on plants for
food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is inge ...
, 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 Agronomy is the science and technology of producing and using plants by agriculture for food, fuel, fiber, chemicals, recreation, or land conservation. Agronomy has come to include research of plant genetics, plant physiology, meteorology, and ...
for arable crops, horticulture for vegetables and fruit, and forestry for timber. About 7,000 species of plant have been used for food, though most of today's food is derived from only 30 species. The major staples include
cereal A cereal is any Poaceae, grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, Cereal germ, germ, and bran. Cereal Grain, grain crops are grown in greater quantit ...
s such as rice and wheat, starchy roots and tubers such as cassava and potato, and
legume A legume () is a plant in the family Fabaceae (or Leguminosae), or the fruit or seed of such a plant. When used as a dry grain, the seed is also called a pulse. Legumes are grown agriculturally, primarily for human consumption, for livestock f ...
s such as peas and
bean A bean is the seed of several plants in the family Fabaceae, which are used as vegetables for human or animal food. They can be cooked in many different ways, including boiling, frying, and baking, and are used in many traditional dishes th ...
s. Vegetable oils such as
olive oil Olive oil is a liquid fat obtained from olives (the fruit of ''Olea europaea''; family Oleaceae), a traditional tree crop of the Mediterranean Basin, produced by pressing whole olives and extracting the oil. It is commonly used in cooking: f ...
and
palm oil Palm oil is an edible vegetable oil derived from the mesocarp (reddish pulp) of the fruit of the oil palms. The oil is used in food manufacturing, in beauty products, and as biofuel. Palm oil accounted for about 33% of global oils produced from ...
provide lipids, while fruit and vegetables contribute vitamins and minerals to the diet.


Medicines

Medicinal plants Medicinal plants, also called medicinal herbs, have been discovered and used in traditional medicine practices since prehistoric times. Plants synthesize hundreds of chemical compounds for various functions, including defense and protection ag ...
are a primary source of organic compounds, both for their medicinal and physiological effects, and for the industrial organic synthesis, synthesis of a vast array of organic chemicals. Many hundreds of medicines are derived from plants, both traditional medicines used in herbalism and chemical substances purified from plants or first identified in them, sometimes by ethnobotany, ethnobotanical search, and then organic synthesis, synthesised for use in modern medicine. Modern medicines derived from plants include aspirin, taxol, morphine, quinine, reserpine, colchicine, digitalis and vincristine. List of plants used in herbalism, Plants used in herbalism include Ginkgo biloba, ginkgo, echinacea, feverfew, and Saint John's wort. The pharmacopoeia of Dioscorides, ''De Materia Medica'', describing some 600 medicinal plants, was written between 50 and 70 AD and remained in use in Europe and the Middle East until around 1600 AD; it was the precursor of all modern pharmacopoeias.


Nonfood products

Plants grown as industrial crops are the source of a wide range of products used in manufacturing, sometimes so intensively as to risk harm to the environment. Nonfood products include essential oils, natural dyes, pigments, waxes, resins, tannins, alkaloids, amber and Cork material, cork. Products derived from plants include soaps, shampoos, perfumes, cosmetics, paint, varnish, turpentine, rubber, latex, lubricants, linoleum, plastics, inks, and Gum (botany), gums. Renewable fuels from plants include firewood, peat and other biofuels. The fossil fuels coal, petroleum and natural gas are derived from the remains of aquatic organisms including phytoplankton in geological time. Structural resources and fibres from plants are used to construct dwellings and to manufacture clothing. Wood is used not only for buildings, boats, and furniture, but also for smaller items such as musical instruments and sports equipment. Wood is Pulp (paper), pulped to make paper and cardboard. Cloth is often made from cotton, flax, ramie or synthetic fibres such as rayon and acetate derived from plant cellulose. Thread (yarn), Thread used to sew cloth likewise comes in large part from cotton.


Aesthetic uses

Thousands of plant species are cultivated for aesthetic purposes as well as to provide shade, modify temperatures, reduce wind, abate noise, provide privacy, and prevent soil erosion. Plants are the basis of a multibillion-dollar per year tourism industry, which includes travel to garden tourism, historic gardens, national parks, rainforests, forests with colorful autumn leaves, and festivals such as Hanami, Japan's and National Cherry Blossom Festival, America's cherry blossom festivals. While some gardens are planted with food crops, many are planted for aesthetic, ornamental, or conservation purposes. Arboretums and botanical gardens are public collections of living plants. In private outdoor gardens, lawn grasses, shade trees, ornamental trees, shrubs, vines, herbaceous perennials and bedding plants are used. Gardens may cultivate the plants in a naturalistic state, or may sculpture their growth, as with topiary or espalier. Gardening is the most popular leisure activity in the U.S., and working with plants or horticulture therapy is beneficial for rehabilitating people with disabilities. Plants may also be grown or kept indoors as houseplants, or in specialized buildings such as greenhouses that are designed for the care and cultivation of living plants. Venus Flytrap, sensitive plant and resurrection plant are examples of plants sold as novelties. There are also art forms specializing in the arrangement of cut or living plant, such as bonsai, ikebana, and the arrangement of cut or dried flowers. Ornamental plants have sometimes changed the course of history, as in tulip mania, tulipomania. Architectural designs resembling plants appear in the capitals of Ancient Egyptian columns, which were carved to resemble either the Nymphaea lotus, Egyptian white lotus or the Cyperus papyrus, papyrus. Images of plants are often used in painting and photography, as well as on textiles, money, stamps, flags and coats of arms.


Scientific and cultural uses

Basic biological research has often been done with plants. In genetics, the breeding of pea plants allowed Gregor Mendel to derive the basic laws governing inheritance, and examination of chromosomes in maize allowed Barbara McClintock to demonstrate their connection to inherited traits. The plant ''
Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small flowering plant native to Eurasia and Africa. ''A. thaliana'' is considered a weed; it is found along the shoulders of roads and in disturbed land. A winter a ...
'' is used in laboratories as a model organism to understand how genes control the growth and development of plant structures. NASA predicts that space stations or space colonies will one day rely on plants for Controlled Ecological Life Support System, life support. Ancient trees are revered and many are List of famous trees, famous. Tree rings themselves are an important method of dating in archeology, and serve as a record of past climates. Plants figure prominently in Trees in mythology, mythology, religion and List of fictional plants, literature. They are used as National emblem, national and state emblems, including List of U.S. state trees, state trees and state flowers. Plants are often used as memorials, gifts and to mark special occasions such as births, deaths, weddings and holidays. The arrangement of flowers may be used to send hidden Language of flowers, messages.


Negative effects

Weeds are commercially or aesthetically undesirable plants growing in managed environments such as agriculture, farms, urban areas, gardens, lawns, and parks. People have spread plants beyond their native ranges and some of these introduced plants become invasive species, invasive, damaging existing ecosystems by displacing native species, and sometimes becoming serious weeds of cultivation. Plants may cause harm to animals, including people. Plants that produce anemophily, windblown pollen invoke allergic reactions in people who suffer from hay fever. A wide variety of plants are List of poisonous plants, poisonous. Toxalbumins are plant poisons fatal to most mammals and act as a serious deterrent to consumption. Several plants cause skin irritations when touched, such as poison ivy. Certain plants contain psychotropic secondary metabolite, chemicals, which are extracted and ingested or smoked, including nicotine from tobacco, cannabinoids from Cannabis sativa, cocaine from Erythroxylon coca and opium from opium poppy. Smoking causes damage to health or even death, while some drugs may also be harmful or fatal to people. Both illegal and legal drugs derived from plants may have negative effects on the economy, affecting worker productivity and law enforcement costs.


See also

* Biosphere * Evolutionary history of plants * Plant defense against herbivory * Plant identification * Plant reproduction * Plant to plant communication via mycorrhizal networks * The Plant List


References


Further reading

;General: * Evans, L.T. (1998). ''Feeding the Ten Billion – Plants and Population Growth''. Cambridge University Press. Paperback, 247 pages. . * Kenrick, Paul & Crane, Peter R. (1997). ''The Origin and Early Diversification of Land Plants: A Cladistic Study''. Washington, D.C.: Smithsonian Institution Press. . * Raven, Peter H.; Evert, Ray F.; & Eichhorn, Susan E. (2005). ''Biology of Plants'' (7th ed.). New York: W.H. Freeman and Company. . * Taylor, Thomas N. & Taylor, Edith L. (1993). ''The Biology and Evolution of Fossil Plants''. Englewood Cliffs, NJ: Prentice Hall. . * ;Species estimates and counts: * International Union for Conservation of Nature and Natural Resources (IUCN) Species Survival Commission (2004). IUCN Red List]
The IUCN Red List of Threatened Species
*


External links

* (requires Microsoft Silverlight) *
Index Nominum Algarum



Plant Resources of Tropical Africa

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

Australia

Chilean plants at ''Chilebosque''

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



Flora of Central Europe

Flora of North America



Meet the Plants-National Tropical Botanical Garden

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

The Plant List

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