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Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are
plants Plants are the eukaryotes that form the kingdom Plantae; they are predominantly photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with cyanobacteria to produce sugars f ...
that have lignified tissues (the
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
) for conducting water and minerals throughout the plant. They also have a specialized non-lignified tissue (the
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
) to conduct products of
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
. The group includes most
land plants The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophy ...
( accepted known species) excluding
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
es. Vascular plants include the
clubmoss Lycopodiopsida is a class of vascular plants also known as lycopsids, lycopods, or lycophytes. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching stems bearing simple leaves ...
es, horsetails,
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s,
gymnosperm The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...
s (including
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 sin ...
s), and angiosperms (
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s). They are contrasted with nonvascular plants such as
mosses 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 ho ...
and
green algae The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...
. Scientific names for the vascular plants group include Tracheophyta, Tracheobionta and Equisetopsida ''sensu lato''. Some early land plants (the
rhyniophyte The rhyniophytes are a group of extinct early vascular plants that are considered to be similar to the genus ''Rhynia'', found in the Early Devonian (around ). Sources vary in the name and Taxonomic rank#Ranks in botany, rank used for this group, ...
s) had less developed vascular tissue; the term eutracheophyte has been used for all other vascular plants, including all living ones. Historically, vascular plants were known as "higher plants", as it was believed that they were further
evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
than other plants due to being more complex organisms. However, this is an antiquated remnant of the obsolete scala naturae, and the term is generally considered to be unscientific.


Characteristics

Botanists define vascular plants by three primary characteristics: # Vascular plants have
vascular tissue Vascular tissue is a complex transporting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. T ...
s which distribute resources through the plant. Two kinds of vascular tissue occur in plants:
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
and
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
. Phloem and xylem are closely associated with one another and are typically located immediately adjacent to each other in the plant. The combination of one xylem and one phloem strand adjacent to each other is known as a
vascular bundle 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 incl ...
. The
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
of vascular tissue in plants allowed them to evolve to larger sizes than
non-vascular plant Non-vascular plants are plants without a vascular system consisting of xylem and phloem. Instead, they may possess simpler tissues that have specialized functions for the internal transport of water. Non-vascular plants include two distantly rel ...
s, which lack these specialized conducting tissues and are thereby restricted to relatively small sizes. # In vascular plants, the principal generation or phase is the ''
sporophyte A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
'', which produces
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s and is
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. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
(having two sets of
chromosomes A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most importa ...
per cell). (By contrast, the principal generation phase in non-vascular plants is the ''
gametophyte A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the se ...
'', which produces
gametes A gamete ( ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. The name gamete was introduced by the Ge ...
and is
haploid Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
, with one set of chromosomes per cell.) # Vascular plants have true roots, leaves, and stems, even if some groups have secondarily lost one or more of these traits. Cavalier-Smith (1998) treated the Tracheophyta as a
phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
or botanical division encompassing two of these characteristics defined by the Latin phrase "facies diploida xylem et phloem instructa" (diploid phase with xylem and phloem). One possible mechanism for the presumed evolution from emphasis on haploid generation to emphasis on diploid generation is the greater efficiency in spore dispersal with more complex diploid structures. Elaboration of the spore stalk enabled the production of more spores and the development of the ability to release them higher and to broadcast them further. Such developments may include more photosynthetic area for the spore-bearing structure, the ability to grow independent roots, woody structure for support, and more branching.
Sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
in vascular land plants involves the process of meiosis. Meiosis provides a direct
DNA repair DNA repair is a collection of processes by which a cell (biology), cell identifies and corrects damage to the DNA molecules that encode its genome. A weakened capacity for DNA repair is a risk factor for the development of cancer. DNA is cons ...
capability for dealing with DNA damages, including oxidative DNA damages, in
germline In biology and genetics, the germline is the population of a multicellular organism's cells that develop into germ cells. In other words, they are the cells that form gametes ( eggs and sperm), which can come together to form a zygote. They dif ...
reproductive tissues.


Phylogeny

A proposed
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
of the vascular plants after Kenrick and Crane 1997 is as follows, with modification to the gymnosperms from Christenhusz ''et al.'' (2011a), Pteridophyta from Smith ''et al.'' and lycophytes and ferns by Christenhusz ''et al.'' (2011b) The cladogram distinguishes the
rhyniophyte The rhyniophytes are a group of extinct early vascular plants that are considered to be similar to the genus ''Rhynia'', found in the Early Devonian (around ). Sources vary in the name and Taxonomic rank#Ranks in botany, rank used for this group, ...
s from the "true" tracheophytes, the eutracheophytes. This phylogeny is supported by several molecular studies. Other researchers state that taking fossils into account leads to different conclusions, for example that the ferns (Pteridophyta) are not monophyletic. Hao and Xue presented an alternative phylogeny in 2013 for pre-
euphyllophyte The euphyllophytes are a clade of plants within the tracheophytes (the vascular plants). The group may be treated as an unranked clade, a division under the name Euphyllophyta or a subdivision under the name Euphyllophytina. The euphyllophytes ...
plants.


Nutrient distribution

Water and
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 form of inorganic solutes are drawn up from the soil by the roots and transported throughout the plant by the
xylem Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
.
Organic compound Some chemical authorities define an organic compound as a chemical compound that contains a carbon–hydrogen or carbon–carbon bond; others consider an organic compound to be any chemical compound that contains carbon. For example, carbon-co ...
s such as
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 ...
produced by
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
in leaves are distributed by the
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
sieve-tube elements. The xylem consists of vessels in
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s and of
tracheid A tracheid is a long and tapered Lignin, lignified cell in the xylem of Tracheophyta, vascular plants. It is a type of conductive cell called a tracheary element. Angiosperms also use another type of conductive cell, called vessel elements, to t ...
s in other vascular plants. Xylem cells are dead, hard-walled hollow cells arranged to form files of tubes that function in water transport. A tracheid cell wall usually contains the polymer
lignin Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidit ...
. The phloem, on the other hand, consists of living cells called sieve-tube members. Between the sieve-tube members are sieve plates, which have pores to allow molecules to pass through. Sieve-tube members lack such organs as nuclei or
ribosome Ribosomes () are molecular machine, macromolecular machines, found within all cell (biology), cells, that perform Translation (biology), biological protein synthesis (messenger RNA translation). Ribosomes link amino acids together in the order s ...
s, but cells next to them, the companion cells, function to keep the sieve-tube members alive.


Transpiration

The most abundant compound in all plants, as in all cellular organisms, is
water Water is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...
, which has an important structural role and a vital role in plant metabolism.
Transpiration Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. It is a passive process that requires no energy expense by the plant. Transpiration also cools plants, c ...
is the main process of water movement within plant tissues. Plants constantly transpire water through their
stoma In botany, a stoma (: stomata, from Greek language, Greek ''στόμα'', "mouth"), also called a stomate (: stomates), is a pore found in the Epidermis (botany), epidermis of leaves, stems, and other organs, that controls the rate of gas exc ...
ta to the atmosphere and replace that water with soil moisture taken up by their roots. When the stomata are closed at night, water pressure can build up in the plant. Excess water is excreted through pores known as hydathodes. The movement of
water Water is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...
out of the leaf stomata sets up transpiration pull or tension in the water column in the xylem vessels or tracheids. The pull is the result of water
surface tension Surface tension is the tendency of liquid surfaces at rest to shrink into the minimum surface area possible. Surface tension (physics), tension is what allows objects with a higher density than water such as razor blades and insects (e.g. Ge ...
within the cell walls of the mesophyll cells, from the surfaces of which evaporation takes place when the stomata are open.
Hydrogen bond In chemistry, a hydrogen bond (H-bond) is a specific type of molecular interaction that exhibits partial covalent character and cannot be described as a purely electrostatic force. It occurs when a hydrogen (H) atom, Covalent bond, covalently b ...
s exist between water
molecule A molecule is a group of two or more atoms that are held together by Force, attractive forces known as chemical bonds; depending on context, the term may or may not include ions that satisfy this criterion. In quantum physics, organic chemi ...
s, causing them to line up; as the molecules at the top of the plant evaporate, each pulls the next one up to replace it, which in turn pulls on the next one in line. The draw of water upwards may be entirely passive and can be assisted by the movement of water into the roots via
osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively permeable membrane, selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of ...
. Consequently, transpiration requires the plant to expend very little energy on water movement. Transpiration assists the plant in absorbing nutrients from the soil as soluble
salts In chemistry, a salt or ionic compound is a chemical compound consisting of an assembly of positively charged ions ( cations) and negatively charged ions (anions), which results in a compound with no net electric charge (electrically neutral). ...
. Transpiration plays an important role in the absorption of nutrients from the soil as soluble salts are transported along with the water from the soil to the leaves. Plants can adjust their transpiration rate to optimize the balance between water loss and nutrient absorption.


Absorption

Living root cells passively absorb water. Pressure within the root increases when transpiration demand via
osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively permeable membrane, selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of ...
is low and decreases when water demand is high. No water movement towards the shoots and leaves occurs when
evapotranspiration Evapotranspiration (ET) refers to the combined processes which move water from the Earth's surface (open water and ice surfaces, bare soil and vegetation) into the Atmosphere of Earth, atmosphere. It covers both water evaporation (movement of w ...
is absent. This condition is associated with high temperature, high
humidity Humidity is the concentration of water vapor present in the air. Water vapor, the gaseous state of water, is generally invisible to the human eye. Humidity indicates the likelihood for precipitation (meteorology), precipitation, dew, or fog t ...
, darkness, and drought.


Conduction

Xylem is the water-conducting tissue, and the secondary xylem provides the raw material for the forest products industry. Xylem and
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
tissues each play a part in the conduction processes within plants. Sugars are conveyed throughout the plant in the phloem; water and other nutrients pass through the xylem. Conduction occurs from a source to a sink for each separate nutrient. Sugars are produced in the leaves (a source) by
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
and transported to the growing shoots and roots (sinks) for use in growth,
cellular respiration Cellular respiration is the process of oxidizing biological fuels using an inorganic electron acceptor, such as oxygen, to drive production of adenosine triphosphate (ATP), which stores chemical energy in a biologically accessible form. Cell ...
or storage. Minerals are absorbed in the roots (a source) and transported to the shoots to allow
cell division Cell division is the process by which a parent cell (biology), cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukar ...
and growth.


See also

*
Fern allies Fern allies are a diverse group of seedless vascular plants that are not true ferns. Like ferns, a fern ally disperses by shedding spores to initiate an alternation of generations. Classification Originally, three or four groups of plants w ...
*
Bryophyte Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s *
Non-vascular plant Non-vascular plants are plants without a vascular system consisting of xylem and phloem. Instead, they may possess simpler tissues that have specialized functions for the internal transport of water. Non-vascular plants include two distantly rel ...
*
Pteridophyte A pteridophyte is a vascular plant (with xylem and phloem) that reproduces by means of spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as " cryptogams", meaning that their means of reproduction is ...


References


Bibliography

* * * * , in


External links


"Higher plants" or "vascular plants"?
{{Authority control Plants Extant Silurian first appearances