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Dendrocnide excelsa

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Xylem is one of the two types of transport 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 of the plants such as 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.

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135-402: Dendrocnide excelsa , commonly known as the giant stinging tree or fibrewood , is a rainforest tree in the nettle family ( Urticaceae ) which is endemic to eastern Australia . It occurs from Tathra, New South Wales to Imbil in southeastern Queensland , and is very common at Dorrigo National Park and other rainforest walks in eastern Australia. The habitat of the giant stinging tree

270-429: A monophyletic taxonomic group but consist of a wide variety of plant species that have independently evolved a trunk and branches as a way to tower above other plants to compete for sunlight. The majority of tree species are angiosperms or hardwoods; of the rest, many are gymnosperms or softwoods. Trees tend to be long-lived, some reaching several thousand years old. Trees evolved around 370 million years ago, and it

405-409: A symbiotic relationship with Frankia species, a filamentous bacterium that can fix nitrogen from the air, converting it into ammonia . They have actinorhizal root nodules on their roots in which the bacteria live. This process enables the tree to live in low nitrogen habitats where they would otherwise be unable to thrive. The plant hormones called cytokinins initiate root nodule formation, in

540-405: A tree is a perennial plant with an elongated stem , or trunk , usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth , plants that are usable as lumber or plants above a specified height. In wider definitions, the taller palms , tree ferns , bananas , and bamboos are also trees. Trees are not

675-416: A classic example of parallel evolution . With an estimated 60,000-100,000 species, the number of trees worldwide might total twenty-five per cent of all living plant species. The greatest number of these grow in tropical regions; many of these areas have not yet been fully surveyed by botanists , making tree diversity and ranges poorly known. The majority of tree species are angiosperms or hardwoods. Of

810-503: A drier savanna climate and insufficient rainfall to support dense forests, the canopy is not closed, and plenty of sunshine reaches the ground which is covered with grass and scrub. Acacia and baobab are well adapted to living in such areas. The roots of a tree serve to anchor it to the ground and gather water and nutrients to transfer to all parts of the tree. They are also used for reproduction, defence, survival, energy storage and many other purposes. The radicle or embryonic root

945-488: A fabric with small spaces. In small passages, such as that between the plant cell walls (or in tracheids), a column of water behaves like rubber – when molecules evaporate from one end, they pull the molecules behind them along the channels. Therefore, transpiration alone provided the driving force for water transport in early plants. However, without dedicated transport vessels, the cohesion-tension mechanism cannot transport water more than about 2 cm, severely limiting

1080-551: A film of surface moisture, enabling them to grow to much greater size. As a result of their independence from their surroundings, they lost their ability to survive desiccation – a costly trait to retain. During the Devonian, maximum xylem diameter increased with time, with the minimum diameter remaining pretty constant. By the middle Devonian, the tracheid diameter of some plant lineages ( Zosterophyllophytes ) had plateaued. Wider tracheids allow water to be transported faster, but

1215-423: A great deal of resistance on flow; vessel members have perforated end walls, and are arranged in series to operate as if they were one continuous vessel. The function of end walls, which were the default state in the Devonian, was probably to avoid embolisms . An embolism is where an air bubble is created in a tracheid. This may happen as a result of freezing, or by gases dissolving out of solution. Once an embolism

1350-418: A hundred litres are required to make one litre of birch syrup. Various parts of trees are used as spices. These include cinnamon , made from the bark of the cinnamon tree ( Cinnamomum zeylanicum ) and allspice , the dried small fruits of the pimento tree ( Pimenta dioica ). Nutmeg is a seed found in the fleshy fruit of the nutmeg tree ( Myristica fragrans ) and cloves are the unopened flower buds of

1485-400: A hundred meters from ground level to a tree 's highest branches. Transpirational pull requires that the vessels transporting the water be very small in diameter; otherwise, cavitation would break the water column. And as water evaporates from leaves, more is drawn up through the plant to replace it. When the water pressure within the xylem reaches extreme levels due to low water input from

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1620-455: A million times more conductive than the inter-cell method, giving the potential for transport over longer distances, and higher CO 2 diffusion rates. The earliest macrofossils to bear water-transport tubes are Silurian plants placed in the genus Cooksonia . The early Devonian pretracheophytes Aglaophyton and Horneophyton have structures very similar to the hydroids of modern mosses. Plants continued to innovate new ways of reducing

1755-426: A plant's vascular system based on the classical research of Dixon-Joly (1894), Eugen Askenasy (1845–1903) (1895), and Dixon (1914,1924). Water is a polar molecule . When two water molecules approach one another, the slightly negatively charged oxygen atom of one forms a hydrogen bond with a slightly positively charged hydrogen atom in the other. This attractive force, along with other intermolecular forces ,

1890-509: A process closely related to mycorrhizal association. It has been demonstrated that some trees are interconnected through their root system, forming a colony. The interconnections are made by the inosculation process, a kind of natural grafting or welding of vegetal tissues. The tests to demonstrate this networking are performed by injecting chemicals, sometimes radioactive , into a tree, and then checking for its presence in neighbouring trees. The roots are, generally, an underground part of

2025-404: A process known as cladoptosis . The crown is the spreading top of a tree including the branches and leaves, while the uppermost layer in a forest, formed by the crowns of the trees, is known as the canopy . A sapling is a young tree. Many tall palms are herbaceous monocots, which do not undergo secondary growth and never produce wood. In many tall palms, the terminal bud on the main stem

2160-422: A range of mechanisms to contain the damage. Small pits link adjacent conduits to allow fluid to flow between them, but not air – although these pits, which prevent the spread of embolism, are also a major cause of them. These pitted surfaces further reduce the flow of water through the xylem by as much as 30%. The diversification of xylem strand shapes with tracheid network topologies increasingly resistant to

2295-537: A robust internal structure that held long narrow channels for transporting water from the soil to all the different parts of the above-soil plant, especially to the parts where photosynthesis occurred. During the Silurian, CO 2 was readily available, so little water needed expending to acquire it. By the end of the Carboniferous, when CO 2 levels had lowered to something approaching today's, around 17 times more water

2430-422: A second spurt of growth, a Lammas growth may occur which is believed to be a strategy to compensate for loss of early foliage to insect predators. Primary growth is the elongation of the stems and roots. Secondary growth consists of a progressive thickening and strengthening of the tissues as the outer layer of the epidermis is converted into bark and the cambium layer creates new phloem and xylem cells. The bark

2565-489: A significant role in reducing erosion and moderating the climate . They remove carbon dioxide from the atmosphere and store large quantities of carbon in their tissues. Trees and forests provide a habitat for many species of animals and plants. Tropical rainforests are among the most biodiverse habitats in the world. Trees provide shade and shelter , timber for construction, fuel for cooking and heating, and fruit for food as well as having many other uses. In much of

2700-424: A similar growth form, by usually growing larger and having a single main stem; but there is no consistent distinction between a tree and a shrub, made more confusing by the fact that trees may be reduced in size under harsher environmental conditions such as on mountains and subarctic areas. The tree form has evolved separately in unrelated classes of plants in response to similar environmental challenges, making it

2835-467: A single tree species, which will not flourish in the absence of its mycorrhizal associate. Others are generalists and associate with many species. The tree acquires minerals such as phosphorus from the fungus, while the fungus obtains the carbohydrate products of photosynthesis from the tree. The hyphae of the fungus can link different trees and a network is formed, transferring nutrients and signals from one place to another. The fungus promotes growth of

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2970-636: A slightly looser definition; while the Joshua tree , bamboos and palms do not have secondary growth and never produce true wood with growth rings, they may produce "pseudo-wood" by lignifying cells formed by primary growth. Tree species in the genus Dracaena , despite also being monocots, do have secondary growth caused by meristem in their trunk, but it is different from the thickening meristem found in dicotyledonous trees. Aside from structural definitions, trees are commonly defined by use; for instance, as those plants which yield lumber. The tree growth habit

3105-446: A stem or root is elongating. Later, 'metaxylem' develops in the strands of xylem. Metaxylem vessels and cells are usually larger; the cells have thickenings typically either in the form of ladderlike transverse bars (scalariform) or continuous sheets except for holes or pits (pitted). Functionally, metaxylem completes its development after elongation ceases when the cells no longer need to grow in size. There are four primary patterns to

3240-484: A tree is any plant with the general form of an elongated stem, or trunk, which supports the photosynthetic leaves or branches at some distance above the ground. Trees are also typically defined by height, with smaller plants from 0.5 to 10 m (1.6 to 32.8 ft) being called shrubs , so the minimum height of a tree is only loosely defined. Large herbaceous plants such as papaya and bananas are trees in this broad sense. A commonly applied narrower definition

3375-458: A vascular bundle will contain primary xylem only. The branching pattern exhibited by xylem follows Murray's law . Primary xylem is formed during primary growth from procambium . It includes protoxylem and metaxylem. Metaxylem develops after the protoxylem but before secondary xylem. Metaxylem has wider vessels and tracheids than protoxylem. Secondary xylem is formed during secondary growth from vascular cambium . Although secondary xylem

3510-436: A wide range of edible fruits are found in the tropics. Other commercially important fruit include dates, figs and olives. Palm oil is obtained from the fruits of the oil palm ( Elaeis guineensis ). The fruits of the cocoa tree ( Theobroma cacao ) are used to make cocoa and chocolate and the berries of coffee trees, Coffea arabica and Coffea canephora , are processed to extract the coffee beans. In many rural areas of

3645-416: A wide range of shapes and sizes, in response to environmental pressures including climate and predation. They can be broad or needle-like, simple or compound, lobed or entire, smooth or hairy, delicate or tough, deciduous or evergreen. The needles of coniferous trees are compact but are structurally similar to those of broad-leaved trees. They are adapted for life in environments where resources are low or water

3780-542: A widespread diverse group of which the only survivor is the maidenhair tree Ginkgo biloba . This is considered to be a living fossil because it is virtually unchanged from the fossilised specimens found in Triassic deposits. During the Mesozoic (245 to 66 million years ago) the conifers flourished and became adapted to live in all the major terrestrial habitats. Subsequently, the tree forms of flowering plants evolved during

3915-415: A young vascular plant grows, one or more strands of primary xylem form in its stems and roots. The first xylem to develop is called 'protoxylem'. In appearance, protoxylem is usually distinguished by narrower vessels formed of smaller cells. Some of these cells have walls that contain thickenings in the form of rings or helices. Functionally, protoxylem can extend: the cells can grow in size and develop while

4050-549: Is a flavouring obtained from distilling bark from the roots of the sassafras tree ( Sassafras albidum ). The leaves of trees are widely gathered as fodder for livestock and some can be eaten by humans but they tend to be high in tannins which makes them bitter. Leaves of the curry tree ( Murraya koenigii ) are eaten, those of kaffir lime ( Citrus × hystrix ) (in Thai food ) and Ailanthus (in Korean dishes such as bugak ) and those of

4185-437: Is a lot lighter, thus cheaper to make, as vessels need to be much more reinforced to avoid cavitation. Xylem development can be described by four terms: centrarch, exarch, endarch and mesarch . As it develops in young plants, its nature changes from protoxylem to metaxylem (i.e. from first xylem to after xylem ). The patterns in which protoxylem and metaxylem are arranged are essential in studying plant morphology. As

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4320-415: Is a purple or blackish nut, maturing from March to August. The flesh is edible, but the danger of stinging hairs precludes human consumption. The fruit is eaten by many rainforest birds, including the regent bowerbird and the green catbird . All aerial parts of the tree have stinging hairs, and can cause a severe reaction on contact with skin, so it is a hazard to livestock, travelers and campers. However,

4455-512: Is a rudimentary stalk and neatly folded miniature leaves, ready to expand when the next growing season arrives. Buds also form in the axils of the leaves ready to produce new side shoots. A few trees, such as the eucalyptus , have "naked buds" with no protective scales and some conifers, such as the Lawson's cypress , have no buds but instead have little pockets of meristem concealed among the scale-like leaves. When growing conditions improve, such as

4590-416: Is a sudden movement of sap at the end of the winter as trees prepare to burst into growth. In North America, the sap of the sugar maple ( Acer saccharum ) is used in the production of maple syrup . About 90% of the sap is water, the remaining 10% being a mixture of various sugars and certain minerals. The sap is harvested by drilling holes in the trunks of the trees and collecting the liquid that flows out of

4725-493: Is also found in members of the gymnosperm groups Gnetophyta and Ginkgophyta and to a lesser extent in members of the Cycadophyta , the two main groups in which secondary xylem can be found are: The xylem, vessels and tracheids of the roots, stems and leaves are interconnected to form a continuous system of water-conducting channels reaching all parts of the plants. The system transports water and soluble mineral nutrients from

4860-545: Is also found in the southern hemisphere, as for example in the Eastern Australia temperate forest, characterised by Eucalyptus forest and open acacia woodland. In tropical regions with a monsoon or monsoon-like climate, where a drier part of the year alternates with a wet period as in the Amazon rainforest , different species of broad-leaved trees dominate the forest, some of them being deciduous. In tropical regions with

4995-528: Is an evolutionary adaptation found in different groups of plants: by growing taller, trees are able to compete better for sunlight. Trees tend to be tall and long-lived, some reaching several thousand years old. Several trees are among the oldest organisms now living. Trees have modified structures such as thicker stems composed of specialised cells that add structural strength and durability, allowing them to grow taller than many other plants and to spread out their foliage. They differ from shrubs , which have

5130-418: Is continually replaced by a living layer of cells called the cork cambium or phellogen. The London plane ( Platanus × hispanica ) periodically sheds its bark in large flakes. Similarly, the bark of the silver birch ( Betula pendula ) peels off in strips. As the tree's girth expands, newer layers of bark are larger in circumference, and the older layers develop fissures in many species. In some trees such as

5265-499: Is developed by the Indian banyan . Many large trees have buttress roots which flare out from the lower part of the trunk. These brace the tree rather like angle brackets and provide stability, reducing sway in high winds. They are particularly prevalent in tropical rainforests where the soil is poor and the roots are close to the surface. Some tree species have developed root extensions that pop out of soil, in order to get oxygen, when it

5400-414: Is estimated that there are around three trillion mature trees in the world currently. A tree typically has many secondary branches supported clear of the ground by the trunk, which typically contains woody tissue for strength, and vascular tissue to carry materials from one part of the tree to another. For most trees the trunk is surrounded by a layer of bark which serves as a protective barrier. Below

5535-474: Is formed, it usually cannot be removed (but see later); the affected cell cannot pull water up, and is rendered useless. End walls excluded, the tracheids of prevascular plants were able to operate under the same hydraulic conductivity as those of the first vascular plant, Cooksonia . The size of tracheids is limited as they comprise a single cell; this limits their length, which in turn limits their maximum useful diameter to 80 μm. Conductivity grows with

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5670-472: Is grey and smooth, with minor corky markings. The trunk and buttresses are shaped in even curves. The leaves are alternate and toothed, heart-shaped and very large in positions of shade, exceeding 30 cm in length and a similar width. The sun leaves are smaller. The leaves, which are covered in stinging hairs, are eaten by various insects such as the chrysomelid beetle, and mammals. Flowers appear from November to April, forming in short panicles . The fruit

5805-486: Is inelastic. Eventually the growth of a tree slows down and stops and it gets no taller. If damage occurs the tree may in time become hollow. Leaves are structures specialised for photosynthesis and are arranged on the tree in such a way as to maximise their exposure to light without shading each other. They are an important investment by the tree and may be thorny or contain phytoliths , lignins , tannins or poisons to discourage herbivory. Trees have evolved leaves in

5940-597: Is lost much faster than CO 2 is absorbed, so plants need to replace it, and have developed systems to transport water from the moist soil to the site of photosynthesis. Early plants sucked water between the walls of their cells, then evolved the ability to control water loss (and CO 2 acquisition) through the use of stomata. Specialized water transport tissues soon evolved in the form of hydroids, tracheids, then secondary xylem, followed by an endodermis and ultimately vessels. The high CO 2 levels of Silurian-Devonian times, when plants were first colonizing land, meant that

6075-424: Is not available in the soil because of excess water. These root extensions are called pneumatophores , and are present, among others, in black mangrove and pond cypress. The main purpose of the trunk is to raise the leaves above the ground, enabling the tree to overtop other plants and outcompete them for light. It also transports water and nutrients from the roots to the aerial parts of the tree, and distributes

6210-435: Is one of the principal factors responsible for the occurrence of surface tension in liquid water. It also allows plants to draw water from the root through the xylem to the leaf. Water is constantly lost through transpiration from the leaf. When one water molecule is lost another is pulled along by the processes of cohesion and tension. Transpiration pull, utilizing capillary action and the inherent surface tension of water,

6345-458: Is ready to eat when the process is complete, having been tenderised and flavoured by the smoke it has absorbed. In the cold process, the temperature is not allowed to rise above 100 °F (38 °C). The flavour of the food is enhanced but raw food requires further cooking. If it is to be preserved, meat should be cured before cold smoking. Wood has traditionally been used for fuel, especially in rural areas. In less developed nations it may be

6480-428: Is related to climatic conditions; growth normally ceases when conditions are either too cold or too dry. In readiness for the inactive period, trees form buds to protect the meristem , the zone of active growth. Before the period of dormancy, the last few leaves produced at the tip of a twig form scales. These are thick, small and closely wrapped and enclose the growing point in a waterproof sheath. Inside this bud there

6615-407: Is scarce. Frozen ground may limit water availability and conifers are often found in colder places at higher altitudes and higher latitudes than broad leaved trees. In conifers such as fir trees, the branches hang down at an angle to the trunk, enabling them to shed snow. In contrast, broad leaved trees in temperate regions deal with winter weather by shedding their leaves. When the days get shorter and

6750-450: Is subtropical, warm temperate or littoral rainforest, particularly in disturbed areas, previously flattened by storms or cyclones. It is well known for its extremely painful, long-lasting sting. Dendrocnide excelsa is a medium to large-sized tree with a buttressed base, sometimes over 40 metres tall and in excess of 6 metres wide at the base. The DBH is up to 4.3 m (14.1 ft). The trunk can be fluted or flanged. The outer bark

6885-402: Is that a tree has a woody trunk formed by secondary growth , meaning that the trunk thickens each year by growing outwards, in addition to the primary upwards growth from the growing tip . Under such a definition, herbaceous plants such as palms , bananas and papayas are not considered trees regardless of their height, growth form or stem girth. Certain monocots may be considered trees under

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7020-471: Is the first part of a seedling to emerge from the seed during the process of germination . This develops into a taproot which goes straight downwards. Within a few weeks lateral roots branch out of the side of this and grow horizontally through the upper layers of the soil. In most trees, the taproot eventually withers away and the wide-spreading laterals remain. Near the tip of the finer roots are single cell root hairs . These are in immediate contact with

7155-452: Is the only one to develop, so they have unbranched trunks with large spirally arranged leaves. Some of the tree ferns, order Cyatheales , have tall straight trunks, growing up to 20 metres (66 ft), but these are composed not of wood but of rhizomes which grow vertically and are covered by numerous adventitious roots . The number of trees in the world, according to a 2015 estimate, is 3.04 trillion, of which 1.39 trillion (46%) are in

7290-497: Is the primary mechanism of water movement in plants. However, it is not the only mechanism involved. Any use of water in leaves forces water to move into them. Transpiration in leaves creates tension (differential pressure) in the cell walls of mesophyll cells. Because of this tension, water is being pulled up from the roots into the leaves, helped by cohesion (the pull between individual water molecules, due to hydrogen bonds) and adhesion (the stickiness between water molecules and

7425-428: Is well worth plants' while to avoid cavitation occurring. For this reason, pits in tracheid walls have very small diameters, to prevent air entering and allowing bubbles to nucleate. Freeze-thaw cycles are a major cause of cavitation. Damage to a tracheid's wall almost inevitably leads to air leaking in and cavitation, hence the importance of many tracheids working in parallel. Once cavitation has occurred, plants have

7560-504: The Cretaceous period. These began to displace the conifers during the Tertiary era (66 to 2 million years ago) when forests covered the globe. When the climate cooled 1.5 million years ago and the first of four glacial periods occurred, the forests retreated as the ice advanced. In the interglacials , trees recolonised the land that had been covered by ice, only to be driven back again in

7695-705: The Silurian (more than 400 million years ago), and trace fossils resembling individual xylem cells may be found in earlier Ordovician rocks. The earliest true and recognizable xylem consists of tracheids with a helical-annular reinforcing layer added to the cell wall . This is the only type of xylem found in the earliest vascular plants, and this type of cell continues to be found in the protoxylem (first-formed xylem) of all living groups of vascular plants. Several groups of plants later developed pitted tracheid cells independently through convergent evolution . In living plants, pitted tracheids do not appear in development until

7830-414: The annual growth rings There may also be rays running at right angles to growth rings. These are vascular rays which are thin sheets of living tissue permeating the wood. Many older trees may become hollow but may still stand upright for many years. Trees do not usually grow continuously throughout the year but mostly have spurts of active expansion followed by periods of rest. This pattern of growth

7965-603: The hydrophilic cell walls of plants). This mechanism of water flow works because of water potential (water flows from high to low potential), and the rules of simple diffusion . Over the past century, there has been a great deal of research regarding the mechanism of xylem sap transport; today, most plant scientists continue to agree that the cohesion-tension theory best explains this process, but multiforce theories that hypothesize several alternative mechanisms have been suggested, including longitudinal cellular and xylem osmotic pressure gradients , axial potential gradients in

8100-546: The pine ( Pinus species) the bark exudes sticky resin which deters attackers whereas in rubber trees ( Hevea brasiliensis ) it is a milky latex that oozes out. The quinine bark tree ( Cinchona officinalis ) contains bitter substances to make the bark unpalatable. Large tree-like plants with lignified trunks in the Pteridophyta , Arecales , Cycadophyta and Poales such as the tree ferns, palms, cycads and bamboos have different structures and outer coverings. Although

8235-402: The "next generation" of transport cell design, have a more rigid structure than hydroids, allowing them to cope with higher levels of water pressure. Tracheids may have a single evolutionary origin, possibly within the hornworts, uniting all tracheophytes (but they may have evolved more than once). Water transport requires regulation, and dynamic control is provided by stomata . By adjusting

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8370-525: The 12,000 years since the start of human agriculture, the number of trees worldwide has decreased by 46%. There are approximately 64,100 known tree species in the world. With 43% of all tree species, South America has the highest biodiversity, followed by Eurasia (22%), Africa (16%), North America (15%), and Oceania (11%). In suitable environments, such as the Daintree Rainforest in Queensland , or

8505-603: The English oak ( Quercus robur ) and 306 species of invertebrate on the Tasmanian oak ( Eucalyptus obliqua ). Non-native tree species provide a less biodiverse community, for example in the United Kingdom the sycamore ( Acer pseudoplatanus ), which originates from southern Europe, has few associated invertebrate species, though its bark supports a wide range of lichens, bryophytes and other epiphytes. Trees differ ecologically in

8640-535: The European bay tree ( Laurus nobilis ) and the California bay tree ( Umbellularia californica ) are used for flavouring food. Camellia sinensis , the source of tea, is a small tree but seldom reaches its full height, being heavily pruned to make picking the leaves easier. Wood smoke can be used to preserve food. In the hot smoking process the food is exposed to smoke and heat in a controlled environment. The food

8775-466: The amount of gas exchange, they can restrict the amount of water lost through transpiration. This is an important role where water supply is not constant, and indeed stomata appear to have evolved before tracheids, being present in the non-vascular hornworts. An endodermis probably evolved during the Silu-Devonian, but the first fossil evidence for such a structure is Carboniferous. This structure in

8910-497: The arrangement of protoxylem and metaxylem in stems and roots. The other three terms are used where there is more than one strand of primary xylem. In his book De plantis libri XVI (On Plants, in 16 books) (1583), the Italian physician and botanist Andrea Cesalpino proposed that plants draw water from soil not by magnetism ( ut magnes ferrum trahit , as magnetic iron attracts) nor by suction ( vacuum ), but by absorption, as occurs in

9045-496: The arrival of warmer weather and the longer days associated with spring in temperate regions, growth starts again. The expanding shoot pushes its way out, shedding the scales in the process. These leave behind scars on the surface of the twig. The whole year's growth may take place in just a few weeks. The new stem is unlignified at first and may be green and downy. The Arecaceae (palms) have their leaves spirally arranged on an unbranched trunk. In some tree species in temperate climates,

9180-437: The bark functions as a protective barrier, it is itself attacked by boring insects such as beetles. These lay their eggs in crevices and the larvae chew their way through the cellulose tissues leaving a gallery of tunnels. This may allow fungal spores to gain admittance and attack the tree. Dutch elm disease is caused by a fungus ( Ophiostoma species) carried from one elm tree to another by various beetles. The tree reacts to

9315-458: The case of linen, sponges, or powders. The Italian biologist Marcello Malpighi was the first person to describe and illustrate xylem vessels, which he did in his book Anatome plantarum ... (1675). Although Malpighi believed that xylem contained only air, the British physician and botanist Nehemiah Grew , who was Malpighi's contemporary, believed that sap ascended both through the bark and through

9450-440: The clove tree ( Syzygium aromaticum ). Many trees have flowers rich in nectar which are attractive to bees. The production of forest honey is an important industry in rural areas of the developing world where it is undertaken by small-scale beekeepers using traditional methods. The flowers of the elder ( Sambucus ) are used to make elderflower cordial and petals of the plum ( Prunus spp. ) can be candied. Sassafras oil

9585-465: The cone for years waiting for a trigger event to liberate it. Fire stimulates release and germination of seeds of the jack pine , and also enriches the forest floor with wood ash and removes competing vegetation. Similarly, a number of angiosperms including Acacia cyclops and Acacia mangium have seeds that germinate better after exposure to high temperatures. The single extant species of Ginkgophyta ( Ginkgo biloba ) has fleshy seeds produced at

9720-405: The disc is sucked into the pore on that side, and blocks further flow. Other plants simply tolerate cavitation. For instance, oaks grow a ring of wide vessels at the start of each spring, none of which survive the winter frosts. Maples use root pressure each spring to force sap upwards from the roots, squeezing out any air bubbles. Growing to height also employed another trait of tracheids –

9855-424: The ease with which they can be found by herbivores. Tree apparency varies with a tree's size and semiochemical content, and with the extent to which it is concealed by nonhost neighbours from its insect pests . In ecosystems such as mangrove swamps, trees play a role in developing the habitat, since the roots of the mangrove trees reduce the speed of flow of tidal currents and trap water-borne sediment, reducing

9990-645: The ends of short branches on female trees, and Gnetum , a tropical and subtropical group of gymnosperms produce seeds at the tip of a shoot axis. The earliest trees were tree ferns , horsetails and lycophytes , which grew in forests in the Carboniferous period. The first tree may have been Wattieza , fossils of which were found in New York state in 2007 dating back to the Middle Devonian (about 385 million years ago). Prior to this discovery, Archaeopteris

10125-461: The far north of the northern hemisphere is moist taiga or northern coniferous forest (also called boreal forest). Taiga is the world's largest land biome , forming 29% of the world's forest cover. The long cold winter of the far north is unsuitable for plant growth and trees must grow rapidly in the short summer season when the temperature rises and the days are long. Light is very limited under their dense cover and there may be little plant life on

10260-480: The food produced by the leaves to all other parts, including the roots. In the case of angiosperms and gymnosperms, the outermost layer of the trunk is the bark , mostly composed of dead cells of phellem (cork). It provides a thick, waterproof covering to the living inner tissue. It protects the trunk against the elements, disease, animal attack and fire. It is perforated by a large number of fine breathing pores called lenticels , through which oxygen diffuses. Bark

10395-401: The forest floor, although fungi may abound. Similar woodland is found on mountains where the altitude causes the average temperature to be lower thus reducing the length of the growing season. Where rainfall is relatively evenly spread across the seasons in temperate regions, temperate broadleaf and mixed forest typified by species like oak, beech, birch and maple is found. Temperate forest

10530-435: The fourth power of diameter, so increased diameter has huge rewards; vessel elements , consisting of a number of cells, joined at their ends, overcame this limit and allowed larger tubes to form, reaching diameters of up to 500 μm, and lengths of up to 10 m. Vessels first evolved during the dry, low CO 2 periods of the late Permian, in the horsetails, ferns and Selaginellales independently, and later appeared in

10665-425: The fruits and either discard the seeds, or swallow them so they pass through the gut to be deposited in the animal's droppings well away from the parent tree. The germination of some seeds is improved when they are processed in this way. Nuts may be gathered by animals such as squirrels that cache any not immediately consumed. Many of these caches are never revisited; the nut-casing softens with rain and frost, and

10800-480: The ground underneath trees there is shade, and often there is undergrowth, leaf litter , and decaying wood that provide other habitat. Trees stabilise the soil, prevent rapid run-off of rain water, help prevent desertification, have a role in climate control and help in the maintenance of biodiversity and ecosystem balance. Many species of tree support their own specialised invertebrates . In their natural habitats, 284 different species of insect have been found on

10935-557: The ground, the roots branch and spread out widely; they serve to anchor the tree and extract moisture and nutrients from the soil . Above ground, the branches divide into smaller branches and shoots. The shoots typically bear leaves, which capture light energy and convert it into sugars by photosynthesis , providing the food for the tree's growth and development. Trees usually reproduce using seeds . Flowering plants have their seeds inside fruits, while conifers carry their seeds in cones, and tree ferns produce spores instead. Trees play

11070-421: The ground. Trees use a variety of methods of seed dispersal . Some rely on wind, with winged or plumed seeds. Others rely on animals, for example with edible fruits. Others again eject their seeds (ballistic dispersal), or use gravity so that seeds fall and sometimes roll. Seeds are the primary way that trees reproduce and their seeds vary greatly in size and shape. Some of the largest seeds come from trees, but

11205-413: The growth of the fungus by blocking off the xylem tissue carrying sap upwards and the branch above, and eventually the whole tree, is deprived of nourishment and dies. In Britain in the 1990s, 25 million elm trees were killed by this disease. The innermost layer of bark is known as the phloem and this is involved in the transport of the sap containing the sugars made by photosynthesis to other parts of

11340-447: The heartwood are blocked in some species. Heartwood is usually darker in colour than the sapwood. It is the dense central core of the trunk giving it rigidity. Three quarters of the dry mass of the xylem is cellulose , a polysaccharide , and most of the remainder is lignin, a complex polymer . A transverse section through a tree trunk or a horizontal core will show concentric circles of lighter or darker wood – tree rings. These rings are

11475-470: The height of a plant increases and upwards transport of water by xylem is considered to limit the maximum height of trees. Three phenomena cause xylem sap to flow: The primary force that creates the capillary action movement of water upwards in plants is the adhesion between the water and the surface of the xylem conduits. Capillary action provides the force that establishes an equilibrium configuration, balancing gravity. When transpiration removes water at

11610-411: The inserted spigots; the sap is then heated to concentrate the flavour. Similarly in northern Europe the spring rise in the sap of the silver birch ( Betula pendula ) is tapped and collected, either to be drunk fresh or fermented into an alcoholic drink. In Alaska, the sap of the sweet birch ( Betula lenta ) is made into a syrup with a sugar content of 67%. Sweet birch sap is more dilute than maple sap;

11745-400: The inside. The newly created xylem is the sapwood . It is composed of water-conducting cells and associated cells which are often living, and is usually pale in colour. It transports water and minerals from the roots to the upper parts of the tree. The oldest, inner part of the sapwood is progressively converted into heartwood as new sapwood is formed at the cambium. The conductive cells of

11880-413: The key innovations that led to the success of the angiosperms . However, the occurrence of vessel elements is not restricted to angiosperms, and they are absent in some archaic or "basal" lineages of the angiosperms: (e.g., Amborellaceae , Tetracentraceae , Trochodendraceae , and Winteraceae ), and their secondary xylem is described by Arthur Cronquist as "primitively vesselless". Cronquist considered

12015-528: The largest tree, Sequoiadendron giganteum , produces one of the smallest tree seeds. The great diversity in tree fruits and seeds reflects the many different ways that tree species have evolved to disperse their offspring. For a tree seedling to grow into an adult tree it needs light. If seeds only fell straight to the ground, competition among the concentrated saplings and the shade of the parent would likely prevent it from flourishing. Many seeds such as birch are small and have papery wings to aid dispersal by

12150-405: The likelihood of cavitation. Cavitation occurs when a bubble of air forms within a vessel, breaking the bonds between chains of water molecules and preventing them from pulling more water up with their cohesive tension. A tracheid, once cavitated, cannot have its embolism removed and return to service (except in a few advanced angiosperms which have developed a mechanism of doing so). Therefore, it

12285-444: The long tracheary elements that transport water. Tracheids and vessel elements are distinguished by their shape; vessel elements are shorter, and are connected together into long tubes that are called vessels . Xylem also contains two other type of cells: parenchyma and fibers . Xylem can be found: In transitional stages of plants with secondary growth , the first two categories are not mutually exclusive, although usually

12420-408: The maturation of the metaxylem (following the protoxylem ). In most plants, pitted tracheids function as the primary transport cells. The other type of vascular element, found in angiosperms, is the vessel element . Vessel elements are joined end to end to form vessels in which water flows unimpeded, as in a pipe. The presence of xylem vessels (also called trachea ) is considered to be one of

12555-403: The mid Cretaceous in angiosperms and gnetophytes. Vessels allow the same cross-sectional area of wood to transport around a hundred times more water than tracheids! This allowed plants to fill more of their stems with structural fibers, and also opened a new niche to vines , which could transport water without being as thick as the tree they grew on. Despite these advantages, tracheid-based wood

12690-409: The mixed podocarp and broadleaf forest of Ulva Island, New Zealand , forest is the more-or-less stable climatic climax community at the end of a plant succession, where open areas such as grassland are colonised by taller plants, which in turn give way to trees that eventually form a forest canopy. In cool temperate regions, conifers often predominate; a widely distributed climax community in

12825-433: The need for water was relatively low. As CO 2 was withdrawn from the atmosphere by plants, more water was lost in its capture, and more elegant transport mechanisms evolved. As water transport mechanisms, and waterproof cuticles, evolved, plants could survive without being continually covered by a film of water. This transition from poikilohydry to homoiohydry opened up new potential for colonization. Plants then needed

12960-519: The next glacial period. Trees are an important part of the terrestrial ecosystem , providing essential habitats including many kinds of forest for communities of organisms. Epiphytic plants such as ferns , some mosses, liverworts, orchids and some species of parasitic plants (e.g., mistletoe ) hang from branches; these along with arboreal lichens, algae, and fungi provide micro-habitats for themselves and for other organisms, including animals. Leaves, flowers and fruits are seasonally available. On

13095-420: The old lineages of flowering plants called basal angiosperms or paleodicots ; these include Amborella , Magnolia , nutmeg and avocado , while trees such as bamboo, palms and bananas are monocots . Wood gives structural strength to the trunk of most types of tree; this supports the plant as it grows larger. The vascular system of trees allows water, nutrients and other chemicals to be distributed around

13230-502: The only fuel available and collecting firewood is often a time-consuming task as it becomes necessary to travel further and further afield in the search for fuel. It is often burned inefficiently on an open fire. In more developed countries other fuels are available and burning wood is a choice rather than a necessity. Modern wood-burning stoves are very fuel efficient and new products such as wood pellets are available to burn. Woody tissue The most distinctive xylem cells are

13365-480: The overall transport rate depends also on the overall cross-sectional area of the xylem bundle itself. The increase in vascular bundle thickness further seems to correlate with the width of plant axes, and plant height; it is also closely related to the appearance of leaves and increased stomatal density, both of which would increase the demand for water. While wider tracheids with robust walls make it possible to achieve higher water transport tensions, this increases

13500-585: The plant and reduce water loss. Both the production of wood and the production of cork are forms of secondary growth. Trees are either evergreen , having foliage that persists and remains green throughout the year, or deciduous , shedding their leaves at the end of the growing season and then having a dormant period without foliage. Most conifers are evergreens, but larches ( Larix and Pseudolarix ) are deciduous, dropping their needles each autumn, and some species of cypress ( Glyptostrobus , Metasequoia and Taxodium ) shed small leafy shoots annually in

13635-409: The plant, and without it trees would not be able to grow as large as they do. Trees need to draw water high up the stem through the xylem from the roots by capillary action , as water continually evaporates from the leaves in the process of transpiration . If insufficient water is available the leaves will die. The three main parts of trees include the root, stem, and leaves; they are integral parts of

13770-415: The resistance to flow within their cells, thereby increasing the efficiency of their water transport. Bands on the walls of tubes, in fact apparent from the early Silurian onwards, are an early improvisation to aid the easy flow of water. Banded tubes, as well as tubes with pitted ornamentation on their walls, were lignified and, when they form single celled conduits, are considered to be tracheids . These,

13905-437: The rest, many are gymnosperms or softwood trees; these include conifers , cycads , ginkgophytes and gnetales , which produce seeds which are not enclosed in fruits, but in open structures such as pine cones , and many have tough waxy leaves, such as pine needles. Most angiosperm trees are eudicots , the "true dicotyledons", so named because the seeds contain two cotyledons or seed leaves. There are also some trees among

14040-416: The roots (if, for example, the soil is dry), then the gases come out of solution and form a bubble – an embolism forms, which will spread quickly to other adjacent cells, unless bordered pits are present (these have a plug-like structure called a torus, that seals off the opening between adjacent cells and stops the embolism from spreading). Even after an embolism has occurred, plants are able to refill

14175-428: The roots and helps protect the trees against predators and pathogens. It can also limit damage done to a tree by pollution as the fungus accumulate heavy metals within its tissues. Fossil evidence shows that roots have been associated with mycorrhizal fungi since the early Paleozoic , four hundred million years ago, when the first vascular plants colonised dry land. Some trees such as Alder ( Alnus species) have

14310-472: The roots covers the water transport tissue and regulates ion exchange (and prevents unwanted pathogens etc. from entering the water transport system). The endodermis can also provide an upwards pressure, forcing water out of the roots when transpiration is not enough of a driver. Once plants had evolved this level of controlled water transport, they were truly homoiohydric, able to extract water from their environment through root-like organs rather than relying on

14445-445: The roots throughout the plant. It is also used to replace water lost during transpiration and photosynthesis. Xylem sap consists mainly of water and inorganic ions, although it can also contain a number of organic chemicals as well. The transport is passive, not powered by energy spent by the tracheary elements themselves, which are dead by maturity and no longer have living contents. Transporting sap upwards becomes more difficult as

14580-464: The seeds on the surface of water. Mangroves often grow in water and some species have buoyant fruits with seeds that start germinating before they detach from the parent tree. These float on the water and may become lodged on emerging mudbanks and successfully take root. Other seeds, such as apple pips and plum stones, have fleshy receptacles and smaller fruits like hawthorns have seeds enclosed in edible tissue; animals including mammals and birds eat

14715-470: The size of the earliest plants. This process demands a steady supply of water from one end, to maintain the chains; to avoid exhausting it, plants developed a waterproof cuticle . Early cuticle may not have had pores but did not cover the entire plant surface, so that gas exchange could continue. However, dehydration at times was inevitable; early plants cope with this by having a lot of water stored between their cell walls, and when it comes to it sticking out

14850-434: The soil particles and can absorb water and nutrients such as potassium in solution. The roots require oxygen to respire and only a few species such as mangroves and the pond cypress ( Taxodium ascendens ) can live in permanently waterlogged soil. In the soil, the roots encounter the hyphae of fungi. Many of these are known as mycorrhiza and form a mutualistic relationship with the tree roots. Some are specific to

14985-526: The spread of embolism likely facilitated increases in plant size and the colonization of drier habitats during the Devonian radiation . Conifers, by the Jurassic, developed bordered pits had valve-like structures to isolate cavitated elements. These torus-margo structures have an impermeable disc (torus) suspended by a permeable membrane (margo) between two adjacent pores. When a tracheid on one side depressurizes,

15120-471: The start of the dry season. Many deciduous trees flower before the new leaves emerge. A few trees do not have true leaves but instead have structures with similar external appearance such as Phylloclades – modified stem structures – as seen in the genus Phyllocladus . Trees can be pollinated either by wind or by animals, mostly insects. Many angiosperm trees are insect pollinated. Wind pollination may take advantage of increased wind speeds high above

15255-399: The sting is to apply wax hair-removal strips and then yank them off to remove the trees' hairs. Dendrocnide stings have been known to kill dogs and horses that have brushed against them. It was found that even heavy welding gloves did not protect researchers from stings while studying the tree. Indigenous Australians used the fibres to make nets and lines. Tree In botany ,

15390-462: The support offered by their lignified walls. Defunct tracheids were retained to form a strong, woody stem, produced in most instances by a secondary xylem. However, in early plants, tracheids were too mechanically vulnerable, and retained a central position, with a layer of tough sclerenchyma on the outer rim of the stems. Even when tracheids do take a structural role, they are supported by sclerenchymatic tissue. Tracheids end with walls, which impose

15525-413: The surviving seeds germinate in the spring. Pine cones may similarly be hoarded by red squirrels , and grizzly bears may help to disperse the seed by raiding squirrel caches. The seeds of conifers, the largest group of gymnosperms, are enclosed in a cone and most species have seeds that are light and papery that can be blown considerable distances once free from the cone. Sometimes the seed remains in

15660-661: The temperature begins to decrease, the leaves no longer make new chlorophyll and the red and yellow pigments already present in the blades become apparent. Synthesis in the leaf of a plant hormone called auxin also ceases. This causes the cells at the junction of the petiole and the twig to weaken until the joint breaks and the leaf floats to the ground. In tropical and subtropical regions, many trees keep their leaves all year round. Individual leaves may fall intermittently and be replaced by new growth but most leaves remain intact for some time. Other tropical species and those in arid regions may shed all their leaves annually, such as at

15795-450: The top, the flow is needed to return to the equilibrium. Transpirational pull results from the evaporation of water from the surfaces of cells in the leaves . This evaporation causes the surface of the water to recess into the pores of the cell wall . By capillary action , the water forms concave menisci inside the pores. The high surface tension of water pulls the concavity outwards, generating enough force to lift water as high as

15930-543: The tough times by putting life "on hold" until more water is supplied. To be free from the constraints of small size and constant moisture that the parenchymatic transport system inflicted, plants needed a more efficient water transport system. During the early Silurian , they developed specialized cells, which were lignified (or bore similar chemical compounds) to avoid implosion; this process coincided with cell death, allowing their innards to be emptied and water to be passed through them. These wider, dead, empty cells were

16065-416: The tree is an important member of the ecosystems of eastern Australian forests. The sting is considered more severe than Dendrocnide photiniphylla (shining-leaved stinging tree), but not as severe as Dendrocnide moroides (gympie stinger). The tree delivers neurotoxin proteins which researchers named gympietides , similar to toxins found in spiders and cone snails, and not previously seen in plants. It

16200-408: The tree, but some tree species have evolved roots that are aerial . The common purposes for aerial roots may be of two kinds, to contribute to the mechanical stability of the tree, and to obtain oxygen from air. An instance of mechanical stability enhancement is the red mangrove that develops prop roots that loop out of the trunk and branches and descend vertically into the mud. A similar structure

16335-432: The tree. It is a soft spongy layer of living cells, some of which are arranged end to end to form tubes. These are supported by parenchyma cells which provide padding and include fibres for strengthening the tissue. Inside the phloem is a layer of undifferentiated cells one cell thick called the vascular cambium layer. The cells are continually dividing, creating phloem cells on the outside and wood cells known as xylem on

16470-492: The tropics or sub-tropics , 0.61 trillion (20%) in the temperate zones , and 0.74 trillion (24%) in the coniferous boreal forests . The estimate is about eight times higher than previous estimates, and is based on tree densities measured on over 400,000 plots. It remains subject to a wide margin of error, not least because the samples are mainly from Europe and North America. The estimate suggests that about 15 billion trees are cut down annually and about 5 billion are planted. In

16605-403: The vascular system which interconnects all the living cells. In trees and other plants that develop wood, the vascular cambium allows the expansion of vascular tissue that produces woody growth. Because this growth ruptures the epidermis of the stem, woody plants also have a cork cambium that develops among the phloem. The cork cambium gives rise to thickened cork cells to protect the surface of

16740-471: The vessels of Gnetum to be convergent with those of angiosperms. Whether the absence of vessels in basal angiosperms is a primitive condition is contested, the alternative hypothesis states that vessel elements originated in a precursor to the angiosperms and were subsequently lost. To photosynthesize, plants must absorb CO 2 from the atmosphere. However, this comes at a price: while stomata are open to allow CO 2 to enter, water can evaporate. Water

16875-408: The vessels, and gel- and gas-bubble-supported interfacial gradients. Until recently, the differential pressure (suction) of transpirational pull could only be measured indirectly, by applying external pressure with a pressure bomb to counteract it. When the technology to perform direct measurements with a pressure probe was developed, there was initially some doubt about whether the classic theory

17010-429: The water depth and creating suitable conditions for further mangrove colonisation. Thus mangrove swamps tend to extend seawards in suitable locations. Mangrove swamps also provide an effective buffer against the more damaging effects of cyclones and tsunamis. Trees are the source of many of the world's best known fleshy fruits. Apples, pears, plums, cherries and citrus are all grown commercially in temperate climates and

17145-437: The wind. Ash trees and maples have larger seeds with blade shaped wings which spiral down to the ground when released. The kapok tree has cottony threads to catch the breeze. The flame tree Delonix regia shoots its seeds through the air when the two sides of its long pods crack apart explosively on drying. The miniature cone-like catkins of alder trees produce seeds that contain small droplets of oil that help disperse

17280-465: The world, forests are shrinking as trees are cleared to increase the amount of land available for agriculture. Because of their longevity and usefulness, trees have always been revered, with sacred groves in various cultures, and they play a role in many of the world's mythologies . Although "tree" is a common word, there is no universally recognised precise definition of what a tree is, either botanically or in common language. In its broadest sense,

17415-769: The world, fruit is gathered from forest trees for consumption. Many trees bear edible nuts which can loosely be described as being large, oily kernels found inside a hard shell. These include coconuts ( Cocos nucifera ), Brazil nuts ( Bertholletia excelsa ), pecans ( Carya illinoinensis ), hazel nuts ( Corylus ), almonds ( Prunus dulcis ), walnuts ( Juglans regia ), pistachios ( Pistacia vera ) and many others. They are high in nutritive value and contain high-quality protein, vitamins and minerals as well as dietary fibre. A variety of nut oils are extracted by pressing for culinary use; some such as walnut, pistachio and hazelnut oils are prized for their distinctive flavours, but they tend to spoil quickly. In temperate climates there

17550-400: The xylem and restore the functionality. The cohesion-tension theory is a theory of intermolecular attraction that explains the process of water flow upwards (against the force of gravity ) through the xylem of plants. It was proposed in 1894 by John Joly and Henry Horatio Dixon . Despite numerous objections, this is the most widely accepted theory for the transport of water through

17685-406: The xylem. However, according to Grew, capillary action in the xylem would raise the sap by only a few inches; to raise the sap to the top of a tree, Grew proposed that the parenchymal cells become turgid and thereby not only squeeze the sap in the tracheids but force some sap from the parenchyma into the tracheids. In 1727, English clergyman and botanist Stephen Hales showed that transpiration by

17820-468: Was correct, because some workers were unable to demonstrate negative pressures. More recent measurements do tend to validate the classic theory, for the most part. Xylem transport is driven by a combination of transpirational pull from above and root pressure from below, which makes the interpretation of measurements more complicated. Xylem appeared early in the history of terrestrial plant life. Fossil plants with anatomically preserved xylem are known from

17955-412: Was lost per unit of CO 2 uptake. However, even in these "easy" early days, water was at a premium, and had to be transported to parts of the plant from the wet soil to avoid desiccation . This early water transport took advantage of the cohesion-tension mechanism inherent in water. Water has a tendency to diffuse to areas that are drier, and this process is accelerated when water can be wicked along

18090-413: Was postulated that the long-term pain from the gympietides may be due to permanent changes in the sodium channels in sensory neurons. The venom binds to and fires pain receptors in the nervous system; there are hopes that understanding how these proteins work could lead to the creation of new painkillers. Minor stings can last for an hour or two. However, severe stinging can last for months. First aid for

18225-450: Was the earliest known tree. Both of these reproduced by spores rather than seeds and are considered to be links between ferns and the gymnosperms which evolved in the Triassic period. The gymnosperms include conifers, cycads, gnetales and ginkgos and these may have appeared as a result of a whole genome duplication event which took place about 319 million years ago. Ginkgophyta was once

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