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A vessel element or vessel member (also called
trachea The trachea, also known as the windpipe, is a Cartilage, cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air-breathing animals with lungs. The trachea extends ...
or xylem vessel) is one of the cell types found in
xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
, the water conducting tissue of plants. Vessel elements are typically found in angiosperms (
flowering plant 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) but absent from most
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό� ...
s such as
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...
. Vessel elements are the main feature distinguishing the "
hardwood Hardwood is wood from dicot trees. These are usually found in broad-leaved temperate and tropical forests. In temperate and boreal latitudes they are mostly deciduous, but in tropics and subtropics mostly evergreen. Hardwood (which comes from ...
" of
angiosperms 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 ...
from the "
softwood file:Pinus sylvestris wood ray section 1 beentree.jpg, Scots Pine, a typical and well-known softwood Softwood is wood from gymnosperm trees such as conifers. The term is opposed to hardwood, which is the wood from angiosperm trees. The main diff ...
" of
conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extan ...
.


Morphology

Xylem Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
is the tissue in
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They al ...
s that conducts water (and substances dissolved in it) upwards in a plant. There are two kinds of cell that are involved in the actual transport:
tracheid A tracheid is a long and tapered lignified cell in the xylem of vascular plants. It is a type of conductive cell called a tracheary element. Angiosperms use another type of tracheary element, called vessel elements, to transport water through the ...
s and vessel elements. Tracheids & vessel elements make the tracheary elements of xylem.Vessel elements are the building blocks of vessels, which constitute the major part of the water transporting system in those plants in which they occur. Vessels form an efficient system for transporting water (including necessary minerals) from the root to the leaves and other parts of the plant. In secondary xylem – the xylem that is produced as a stem thickens rather than when it first appears – a vessel element originates from the
vascular cambium The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xyle ...
. A long cell, oriented along the axis of the stem, called a "fusiform initial", divides along its length forming new vessel elements. The cell wall of a vessel element becomes strongly "lignified", i.e. it develops reinforcing material made of
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 rigidity ...
. The side walls of a vessel element have pits: more or less circular regions in contact with neighbouring cells. Tracheids also have pits, but only vessel elements have openings at both ends that connect individual vessel elements to form a continuous tubular vessel. These end openings are called perforations or perforation plates. They have a variety of shapes: the most common are the simple perforation (a simple opening) and the scalariform perforation (several elongated openings in a ladder-like design). Other types include the foraminate perforation plate (several round openings) and the reticulate perforation plate (a net-like pattern, with many openings). At maturity, the protoplast – the living material of the cell – dies and disappears, but the lignified cell walls persist. A vessel element is then a dead cell, but one that still has a function, and is still being protected by surrounding living cells.


Evolutionary significance

The presence of vessels in xylem has been considered to be one of the key innovations that led to the success of the flowering plants. It was once thought that vessel elements were an evolutionary innovation of flowering plants, but their absence from some basal
angiosperms 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 ...
and their presence in some members of the
Gnetales Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three reli ...
suggest that this hypothesis must be re-examined; vessel elements in Gnetales may not be homologous with those of angiosperms, or vessel elements that originated in a precursor to the angiosperms may have been subsequently lost in some basal lineages (e.g.,
Amborellaceae ''Amborella'' is a monotypic genus of understory shrubs or small trees endemic to the main island, Grande Terre, of New Caledonia in the southwest Pacific Ocean. The genus is the only member of the family Amborellaceae and the order Amborellale ...
,
Trochodendraceae Trochodendraceae is the only family of flowering plants in the order Trochodendrales. It comprises two extant genera, each with a single species along with up to five additional extinct genera and a number of extinct species. The living specie ...
, and
Winteraceae Winteraceae is a primitive family of tropical trees and shrubs including 93 species in five genera. It is of particular interest because it is such a primitive angiosperm family, distantly related to Magnoliaceae, though it has a much more s ...
), described by
Arthur Cronquist Arthur John Cronquist (March 19, 1919 – March 22, 1992) was an American biologist, botanist and a specialist on Compositae. He is considered one of the most influential botanists of the 20th century, largely due to his formulation of the Cr ...
as "primitively vesselless". Cronquist considered the vessels of ''
Gnetum ''Gnetum'' is a genus of gymnosperms, the sole genus in the family Gnetaceae within the Gnetophyta. They are tropical evergreen trees, shrubs and lianas. Unlike other gymnosperms, they possess vessel elements in the xylem. Some species have been ...
'' to be convergent with those of angiosperms. Vessel-like cells have also been found in the xylem of ''
Equisetum ''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass Eq ...
'' (horsetails), ''
Selaginella ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...
'' (spike-mosses), ''
Pteridium aquilinum ''Pteridium aquilinum'' (bracken, brake or common bracken), also known as eagle fern, is a species of fern occurring in temperate and subtropical regions in both hemispheres. Originally native to Eurasia and North America, the extreme lightness o ...
'' (bracken fern), ''
Marsilea ''Marsilea'' is a genus of approximately 65 species of aquatic ferns of the family Marsileaceae. The name honours Italian naturalist Luigi Ferdinando Marsili (1656–1730). These small plants are of unusual appearance and do not resemble common ...
'' and ''
Regnellidium Regnellidium is a monotypic genus of ferns of family Marsileaceae. The single living species, ''Regnellidium diphyllum'', the two-leaf water fern, is native to southeastern Brazil and adjacent regions of Argentina. It resembles its relatives fr ...
'' (
aquatic fern The order Salviniales (formerly known as the Hydropteridales and including the former Marsileales) is an order of ferns in the class Polypodiopsida. Description Salviniales are all aquatic and differ from all other ferns in being heterosporous, ...
s), and the enigmatic fossil group Gigantopteridales. In these cases, it is generally agreed that the vessels evolved independently. It is possible that vessels may have appeared more than once among the angiosperms as well.


See also

*
Tracheid A tracheid is a long and tapered lignified cell in the xylem of vascular plants. It is a type of conductive cell called a tracheary element. Angiosperms use another type of tracheary element, called vessel elements, to transport water through the ...


References


Further references

* Niklas, Karl J. (1997) ''The Evolutionary Biology of Plants''. Chicago and London: The University of Chicago Press. . * Schweingruber, F. H. (1990) ''Anatomie europäischer Hölzer - Anatomy of European woods''. Eidgenössische Forschungsanstalt für Wald, Schnee und Landscaft, Birmensdorf (Hrsg.) Haupt, Bern und Stuttgart. * Timonen, Tuuli (2002). ''Introduction to Microscopic Wood Identification''. Finnish Museum of Natural History, University of Helsinki. * {{Biological tissue Plant anatomy Wood