|Common lady-fern, a non-seed-bearing plant|
|Lemon basil, a seed-bearing plant|
|Clade:|| Tracheophytes |
Sinnott, 1935 ex Cavalier-Smith, 1998
Vascular plants (from Latin vasculum 'duct'), also called tracheophytes ( // ) or collectively Tracheophyta (from Ancient Greek τραχεῖα ἀρτηρία (trakheîa artēría) 'windpipe',and φυτά (phutá) 'plants'), form a large group of land plants (c. 300,000 accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified tissue (the phloem) to conduct products of photosynthesis. Vascular plants include the clubmosses, horsetails, ferns, gymnosperms (including conifers), and angiosperms (flowering plants). Scientific names for the group include Tracheophyta, : 251 Tracheobionta and Equisetopsida sensu lato. Some early land plants (the rhyniophytes) 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 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.
Botanists define vascular plants by three primary characteristics:
Cavalier-Smith (1998) treated the Tracheophyta as a phylum 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). : 251
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 farther. 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.[ citation needed ]
A proposed phylogeny of the vascular plants after Kenrick and Crane 1997is 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 rhyniophytes 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 plants.
Water and nutrients in the form of inorganic solutes are drawn up from the soil by the roots and transported throughout the plant by the xylem. Organic compounds such as sucrose produced by photosynthesis in leaves are distributed by the phloem sieve-tube elements.
The xylem consists of vessels in flowering plants and of tracheids in other vascular plants. Xylem cells are dead hard-walled hollow cells arranged to form files of tubes that function in the transport of water. A tracheid cell-wall usually contains the polymer lignin.
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 ribosomes, but cells next to them, the companion cells, function to keep the sieve-tube members alive.
The most abundant compound in all plants, as in all cellular organisms, is water, which has an important structural role and a vital role in plant metabolism. Transpiration is the main process of water movement within plant tissues. Plants constantly transpire water through their stomata to the atmosphere and replace that water with soil moisture taken up by their roots. The movement of water out of the leaf stomata sets up a transpiration pull or tension in the water-column in the xylem vessels or tracheids. The pull is the result of water surface tension within the cell walls of the mesophyll cells, from the surfaces of which evaporation takes place when the stomata are open. Hydrogen bonds exist between water molecules, 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. 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.
Living root cells passively absorb water in the absence of transpiration pull via osmosis creating root pressure. It is possible for there to be no evapotranspiration and therefore no pull of water towards the shoots and leaves. This is usually due to high temperatures, high humidity, darkness or drought.[ citation needed ]
Xylem and phloem tissues each play a part in the conduction processes within plants. Sugars are conducted throughout the plant in the phloem; water and other nutrients 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 and transported to the growing shoots and roots (sinks) for use in growth, cellular respiration or storage. Minerals are absorbed in the roots (a source) and transported to the shoots to allow cell division and growth.
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 sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.
Plant cells are eukaryotic cells present in green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids with the capability to perform photosynthesis and store starch, a large vacuole that regulates turgor pressure, the absence of flagella or centrioles, except in the gametes, and a unique method of cell division involving the formation of a cell plate or phragmoplast that separates the new daughter cells.
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 Ancient Greek word ξύλον (xylon), meaning "wood"; the best-known xylem tissue is wood, though it is found throughout a plant. The term was introduced by Carl Nägeli in 1858.
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 called translocation. In trees, the phloem is the innermost layer of the bark, hence the name, derived from the Ancient Greek word φλοιός (phloiós), meaning "bark". The term was introduced by Carl Nägeli in 1858.
A fern is a member of a group of vascular plants that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except the lycopods, and differ from mosses and other bryophytes by being vascular, i.e., having specialized tissues that conduct water and nutrients and in having life cycles in which the branched sporophyte is the dominant phase. Ferns have complex leaves called megaphylls, that are more complex than the microphylls of clubmosses. Most ferns are leptosporangiate ferns. They produce coiled fiddleheads that uncoil and expand into fronds. The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the Polypodiopsida, comprising both the leptosporangiate (Polypodiidae) and eusporangiate ferns, the latter group including horsetails, whisk ferns, marattioid ferns, and ophioglossoid ferns.
In biology, tissue is a biological organizational level between cells and a complete organ. A tissue is an ensemble of similar cells and their extracellular matrix from the same origin that together carry out a specific function. Organs are then formed by the functional grouping together of multiple tissues.
Osmundaceae is a family of ferns containing four to six extant genera and 18–25 known species. It is the only living family of the order Osmundales in the class Polypodiopsida (ferns) or in some classifications the only order in the class Osmundopsida. This is an ancient and fairly isolated group that is often known as the "flowering ferns" because of the striking aspect of the ripe sporangia in Claytosmunda, Osmunda, Osmundastrum, and Plensium. In these genera the sporangia are borne naked on non-laminar pinnules, while Todea and Leptopteris bear sporangia naked on laminar pinnules. Ferns in this family are larger than most other ferns.
Root pressure is the transverse osmotic pressure within the cells of a root system that causes sap to rise through a plant stem to the leaves.
The Embryophyta, or land plants, are the most familiar group of green plants that comprise vegetation on Earth. Embryophytes have a common ancestor with green algae, having emerged within the Phragmoplastophyta clade of green algae as sister of the Zygnematophyceae. The Embryophyta consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, lycophytes, ferns, gymnosperms and flowering plants. The land plants have diplobiontic life cycles and it is accepted now that they emerged from freshwater, multi-celled algae.
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 xylem. The main functions of tracheid cells are to transport water and inorganic salts, and to provide structural support for trees. There are often pits on the cell walls of tracheids, which allows for water flow between cells. Tracheids are dead at functional maturity and do not have a protoplast. The wood (softwood) of gymnosperms such as pines and other conifers is mainly composed of tracheids. Tracheids are also the main conductive cells in the primary xylem of ferns.
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.
Equisetidae is one of the four subclasses of Polypodiopsida (ferns), a group of vascular plants with a fossil record going back to the Devonian. They are commonly known as horsetails. They typically grow in wet areas, with whorls of needle-like branches radiating at regular intervals from a single vertical stem.
A pteridophyte is a vascular plant that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden. Ferns, horsetails, and lycophytes are all pteridophytes. However, they do not form a monophyletic group because ferns are more closely related to seed plants than to lycophytes. "Pteridophyta" is thus no longer a widely accepted taxon, but the term pteridophyte remains in common parlance, as do pteridology and pteridologist as a science and its practitioner, respectively. Ferns and lycophytes share a life cycle and are often collectively treated or studied, for example by the International Association of Pteridologists and the Pteridophyte Phylogeny Group.
Psilotaceae is a family of ferns consisting of two genera, Psilotum and Tmesipteris with about a dozen species. It is the only family in the order Psilotales.
A vessel element or vessel member is one of the cell types found in xylem, the water conducting tissue of plants. Vessel elements are typically found in angiosperms but absent from most gymnosperms such as conifers. Vessel elements are the main feature distinguishing the "hardwood" of angiosperms from the "softwood" of conifers.
The Hymenophyllaceae, the filmy ferns and bristle ferns, are a family of two to nine genera and about 650 known species of ferns, with a subcosmopolitan distribution, but generally restricted to very damp places or to locations where they are wetted by spray from waterfalls or springs. A recent fossil find shows that ferns of Hymenophyllaceae have existed since at least the Upper Triassic.
The evolution of plants has resulted in a wide range of complexity, from the earliest algal mats, through multicellular marine and freshwater green algae, terrestrial bryophytes, lycopods and ferns, to the complex gymnosperms and angiosperms of today. While many of the earliest groups continue to thrive, as exemplified by red and green algae in marine environments, more recently derived groups have displaced previously ecologically dominant ones; for example, the ascendance of flowering plants over gymnosperms in terrestrial environments.
Lepidodendrales were primitive, vascular, arborescent (tree-like) plants related to the lycopsids. Members of Lepidodendrales are the best understood of the fossil lycopsids due to the vast diversity of Lepidodendrales specimens and the diversity in which they were preserved; the extensive distribution of Lepidodendrales specimens as well as their well-preservedness lends paleobotanists exceptionally detailed knowledge of the coal-swamp giants’ reproductive biology, vegetative development, and role in their paleoecosystem. The defining characteristics of the Lepidodendrales are their secondary xylem, extensive periderm development, three-zoned cortex, rootlike appendages known as stigmarian rootlets arranged in a spiralling pattern, and megasporangium each containing a single functional megaspore that germinates inside the sporangium. Many of these different plant organs have been assigned both generic and specific names as relatively few have been found organically attached to each other. Some specimens have been discovered which indicate heights of 40 and even 50 meters and diameters of over 2 meters at the base. The massive trunks of some species branched profusely, producing large crowns of leafy twigs; though some leaves were up to 1 meter long, most were much shorter, and when leaves dropped from branches their conspicuous leaf bases remained on the surface of branches. Strobili could be found at the tips of distal branches or in an area at the top of the main trunk. The underground organs of Lepidodendrales typically consisted of dichotomizing axes bearing helically arranged, lateral appendages serving an equivalent function to roots. Sometimes called "giant club mosses", they are believed to be more closely related to extant quillworts based on xylem, although fossil specimens of extinct Selaginellales from the Late Carboniferous also had secondary xylem.
A stem is one of two main structural axes of a vascular plant, the other being the root. It supports leaves, flowers and fruits, transports water and dissolved substances between the roots and the shoots in the xylem and phloem, stores nutrients, and produces new living tissue.
Transpiration is the process of water movement through a plant and its evaporation from aerial parts, such as leaves, stems and flowers. Water is necessary for plants but only a small amount of water taken up by the roots is used for growth and metabolism. The remaining 97–99.5% is lost by transpiration and guttation. Leaf surfaces are dotted with pores called stomata, and in most plants they are more numerous on the undersides of the foliage. The stomata are bordered by guard cells and their stomatal accessory cells that open and close the pore. Transpiration occurs through the stomatal apertures, and can be thought of as a necessary "cost" associated with the opening of the stomata to allow the diffusion of carbon dioxide gas from the air for photosynthesis. Transpiration also cools plants, changes osmotic pressure of cells, and enables mass flow of mineral nutrients and water from roots to shoots. Two major factors influence the rate of water flow from the soil to the roots: the hydraulic conductivity of the soil and the magnitude of the pressure gradient through the soil. Both of these factors influence the rate of bulk flow of water moving from the roots to the stomatal pores in the leaves via the xylem.