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botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
, a tree is a
perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
plant with an elongated stem, or trunk, usually supporting branches and
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
. In some usages, the definition of a tree may be narrower, e.g., including only woody plants with secondary growth, only plants that are usable as
lumber Lumber is wood that has been processed into uniform and useful sizes (dimensional lumber), including beams and planks or boards. Lumber is mainly used for construction framing, as well as finishing (floors, wall panels, window frames). ...
, or only plants above a specified height. But wider definitions include taller palms, tree ferns,
bananas A banana is an elongated, edible fruit – berry (botany), botanically a berry – produced by several kinds of large treelike herbaceous flowering plants in the genus ''Musa (genus), Musa''. In some countries, cooking bananas are called pla ...
, and
bamboo Bamboos are a diverse group of mostly evergreen perennial plant, perennial flowering plants making up the subfamily (biology), subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family, in th ...
s. Trees are not 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
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
s or hardwoods; of the rest, many are gymnosperms or softwoods. Trees tend to be long-lived, some trees reaching several thousand years old. Trees evolved around 400 million years ago, and it 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 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 ...
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 the ground, the
roots A root is the part of a plant, generally underground, that anchors the plant body, and absorbs and stores water and nutrients. Root or roots may also refer to: Art, entertainment, and media * ''The Root'' (magazine), an online magazine focusin ...
branch and spread out widely; they serve to anchor the tree and extract moisture and nutrients from the
soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
. 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 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 ...
, 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
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 instead. Trees play a significant role in reducing
erosion Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust and then sediment transport, tran ...
and moderating the
climate Climate is the long-term weather pattern in a region, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteoro ...
. They remove
carbon dioxide Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
from the
atmosphere An atmosphere () is a layer of gases that envelop an astronomical object, held in place by the gravity of the object. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A stellar atmosph ...
and store large quantities of
carbon Carbon () is a chemical element; it has chemical symbol, symbol C and atomic number 6. It is nonmetallic and tetravalence, tetravalent—meaning that its atoms are able to form up to four covalent bonds due to its valence shell exhibiting 4 ...
in their tissues. Trees and
forest A forest is an ecosystem characterized by a dense ecological community, community of trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, ...
s provide a habitat for many species of animals and plants.
Tropical rainforest Tropical rainforests are dense and warm rainforests with high rainfall typically found between 10° north and south of the Equator. They are a subset of the tropical forest biome that occurs roughly within the 28° latitudes (in the torrid zo ...
s 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 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 Myth is a genre of folklore consisting primarily of narratives that play a fundamental role in a society. For scholars, this is very different from the vernacular usage of the term "myth" that refers to a belief that is not true. Instead, the ...
.


Definition

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, a tree is any plant with the general form of an elongated stem, or trunk, which supports the
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
leaves or branches at some distance above the ground. Trees are also typically defined by height, with smaller plants from being called
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s, 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 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 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.


Overview

The tree growth habit is an
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 ...
ary
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
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
shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...
s, which have 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 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
botanist Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
s, making tree diversity and ranges poorly known. The majority of tree species are
angiosperm Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
s or hardwoods. Of the rest, many are gymnosperms or softwood trees; these include
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, 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 the old lineages of flowering plants called basal angiosperms or paleodicots; these include ''
Amborella ''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 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 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 Capillary action (sometimes called capillarity, capillary motion, capillary rise, capillary effect, or wicking) is the process of a liquid flowing in a narrow space without the assistance of external forces like Gravitation, gravity. The effe ...
, 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 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 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 In botany, an evergreen is a plant which has Leaf, foliage that remains green and functional throughout the year. This contrasts with deciduous plants, which lose their foliage completely during the winter or dry season. Consisting of many diffe ...
, having foliage that persists and remains green throughout the year, or
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
, 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 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 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 , but these are composed not of wood but of
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s which grow vertically and are covered by numerous adventitious roots.


Distribution

The number of trees in the world, according to a 2015 estimate, is 3.04 trillion, of which 1.39 trillion (46%) are in 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 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 Queensland ( , commonly abbreviated as Qld) is a States and territories of Australia, state in northeastern Australia, and is the second-largest and third-most populous state in Australia. It is bordered by the Northern Territory, South Austr ...
, or 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 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 A biome () is a distinct geographical region with specific climate, vegetation, and animal life. It consists of a biological community that has formed in response to its physical environment and regional climate. In 1935, Tansley added the ...
, 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 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 is also found in the southern hemisphere, as for example in the Eastern Australia temperate forest, characterised by ''
Eucalyptus ''Eucalyptus'' () is a genus of more than 700 species of flowering plants in the family Myrtaceae. Most species of ''Eucalyptus'' are trees, often Mallee (habit), mallees, and a few are shrubs. Along with several other genera in the tribe Eucalyp ...
'' forest and open acacia woodland. In tropical regions with a
monsoon A monsoon () is traditionally a seasonal reversing wind accompanied by corresponding changes in precipitation but is now used to describe seasonal changes in Atmosphere of Earth, atmospheric circulation and precipitation associated with annu ...
or monsoon-like climate, where a drier part of the year alternates with a wet period as in the
Amazon rainforest The Amazon rainforest, also called the Amazon jungle or Amazonia, is a Tropical and subtropical moist broadleaf forests, moist broadleaf tropical rainforest in the Amazon biome that covers most of the Amazon basin of South America. This basin ...
, different species of broad-leaved trees dominate the forest, some of them being deciduous. In tropical regions with 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.


Parts


Roots

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 is the first part of a
seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embry ...
to emerge from the seed during the process of germination. This develops into a
taproot A taproot is a large, central, and dominant root from which other roots sprout laterally. Typically a taproot is somewhat straight and very thick, is tapering in shape, and grows directly downward. In some plants, such as the carrot, the 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 the soil particles and can absorb water and nutrients such as
potassium Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...
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 A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
of fungi. Many of these are known as mycorrhiza and form a mutualistic relationship with the tree roots. Some are specific to 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 Phosphorus is a chemical element; it has Chemical symbol, symbol P and atomic number 15. All elemental forms of phosphorus are highly Reactivity (chemistry), reactive and are therefore never found in nature. They can nevertheless be prepared ar ...
from the fungus, while the fungus obtains the
carbohydrate A carbohydrate () is a biomolecule composed of carbon (C), hydrogen (H), and oxygen (O) atoms. The typical hydrogen-to-oxygen atomic ratio is 2:1, analogous to that of water, and is represented by the empirical formula (where ''m'' and ''n'' ...
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 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 The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
, four hundred million years ago, when the first
vascular plant Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
s colonised dry land. Some trees such as
Alder Alders are trees of the genus ''Alnus'' in the birch family Betulaceae. The genus includes about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few species ex ...
(''Alnus'' species) have a symbiotic relationship with '' Frankia'' species, a filamentous bacterium that can fix nitrogen from the air, converting it into
ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...
. 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
cytokinin Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in Cell (biology), cell growth and cellular differentiation, differentiation, but also affect apical ...
s initiate root nodule formation, in 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 Grafting or graftage is a horticulture, horticultural technique whereby tissues of plants are joined so as to continue their growth together. The upper part of the combined plant is called the scion () while the lower part is called the roots ...
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 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 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 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.


Trunk

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 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 is continually replaced by a living layer of cells called the cork cambium or phellogen. The
London plane The London plane, or sometimes hybrid plane, ''Platanus'' × ''hispanica'', is a tree in the genus ''Platanus''. It is often known by the Synonym (taxonomy), synonym ''Platanus'' × ''acerifolia'', a later name. It is a Hybrid (biology ...
(''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 the
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...
(''Pinus'' species) the bark exudes sticky
resin A resin is a solid or highly viscous liquid that can be converted into a polymer. Resins may be biological or synthetic in origin, but are typically harvested from plants. Resins are mixtures of organic compounds, predominantly terpenes. Commo ...
which deters attackers whereas in rubber trees (''Hevea brasiliensis'') it is a milky
latex Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latices are found in nature, but synthetic latices are common as well. In nature, latex is found as a wikt:milky, milky fluid, which is present in 10% of all floweri ...
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 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 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 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 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 ...
on 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 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 Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
, a
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wat ...
, and most of the remainder is lignin, a complex
polymer A polymer () is a chemical substance, substance or material that consists of very large molecules, or macromolecules, that are constituted by many repeat unit, repeating subunits derived from one or more species of monomers. Due to their br ...
. 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 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.


Buds and growth

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 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 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 ''Eucalyptus'' () is a genus of more than 700 species of flowering plants in the family Myrtaceae. Most species of ''Eucalyptus'' are trees, often Mallee (habit), mallees, and a few are shrubs. Along with several other genera in the tribe Eucalyp ...
, 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 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, 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 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. File:Illustration Quercus robur0.jpg, Buds, leaves, flowers and fruit of pedunculate oak, '' Quercus robur'' File:Illustration Abies alba0.jpg, Buds, leaves and reproductive structures of white fir, '' Abies alba'' File:Cycas circinalis(draw).jpg, Form, leaves and reproductive structures of queen sago, '' Cycas circinalis'' File:Buds_of_Fraxinus_excelsior_03.jpg, Dormant buds of ash, '' Fraxinus excelsior''


Leaves

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
phytolith Phytoliths (from Greek language, Greek, "plant stone") are rigid, microscopic mineral deposits found in some plant tissues, often persisting after the decay of the plant. Although some use "phytolith" to refer to all mineral secretions by plants, ...
s,
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 ...
s,
tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and Precipitation (chemistry), precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' is widel ...
s or
poison A poison is any chemical substance that is harmful or lethal to living organisms. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figurati ...
s to discourage herbivory. Trees have evolved leaves in 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 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 the temperature begins to decrease, the leaves no longer make new
chlorophyll Chlorophyll is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words (, "pale green") and (, "leaf"). Chlorophyll allows plants to absorb energy ...
and the red and yellow pigments already present in the blades become apparent. Synthesis in the leaf of a
plant hormone Plant hormones (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, including embryogenesis, the regulation of Organ (anat ...
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 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''.Page, Christopher N. (1990). "Phyllocladaceae" pp. 317–319. In: Klaus Kubitzki (general editor); Karl U. Kramer and Peter S. Green (volume editors) ''The Families and Genera of Vascular Plants'' volume I. Springer-Verlag: Berlin; Heidelberg, Germany.


Reproduction

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 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

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 the largest tree, ''
Sequoiadendron giganteum ''Sequoiadendron giganteum'' (also known as the giant sequoia, giant redwood, Sierra redwood or Wellingtonia) is a species of coniferous tree, classified in the family Cupressaceae in the subfamily Sequoioideae. Giant sequoia specimens are the la ...
'', 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 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 Alders are trees of the genus ''Alnus'' in the birch family Betulaceae. The genus includes about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few species ex ...
trees produce seeds that contain small droplets of oil that help disperse 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 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 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 the cone for years waiting for a trigger event to liberate it. Fire stimulates release and germination of seeds of the
jack pine Jack pine (''Pinus banksiana''), also known as grey pine or scrub pine, is a North American pine. Distribution and habitat Its native range in Canada is east of the Rocky Mountains from the Mackenzie River in the Northwest Territories t ...
, 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 Ginkgoopsida is a proposed class of gymnosperms defined by Sergei V. Meyen in 1984 to encompass Ginkgoales (which contains the living ''Ginkgo'') alongside a number of extinct seed plant groups, which he considered to be closely related based ...
('' Ginkgo biloba'') has fleshy seeds produced at 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.


Evolutionary history

The earliest trees were tree ferns, horsetails and lycophytes, which grew in forests in the
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period. The first tree may have been '' Wattieza'', fossils of which were found in
New York state New York, also called New York State, is a state in the northeastern United States. Bordered by New England to the east, Canada to the north, and Pennsylvania and New Jersey to the south, its territory extends into both the Atlantic Ocean and ...
in 2007 dating back to the Middle Devonian (about 385 million years ago). Prior to this discovery, '' Archaeopteris'' 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 The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
period. The gymnosperms include conifers, cycads, gnetales and
ginkgo ''Ginkgo'' is a genus of non-flowering seed plants, assigned to the gymnosperms. The scientific name is also used as the English common name. The order to which the genus belongs, Ginkgoales, first appeared in the Permian, , and ''Ginkgo'' is n ...
s and these may have appeared as a result of a whole genome duplication event which took place about 319 million years ago.
Ginkgophyta Ginkgoopsida is a proposed class of gymnosperms defined by Sergei V. Meyen in 1984 to encompass Ginkgoales (which contains the living ''Ginkgo'') alongside a number of extinct seed plant groups, which he considered to be closely related based ...
was once a widespread diverse group of which the only survivor is the maidenhair tree '' Ginkgo biloba''. This is considered to be a
living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...
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 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 evolved during the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
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 the next glacial period.


Ecology

Trees are an important part of the terrestrial
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
, providing essential habitats including many kinds of forest for communities of organisms.
Epiphytic An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
plants such as
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, some mosses, liverworts,
orchid Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant. Orchids are cosmopolitan plants that are found in almost every habitat on Eart ...
s 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 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 Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
and ecosystem balance. Many species of tree support their own specialised
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s. In their natural habitats, 284 different species of insect have been found on 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 ''Acer pseudoplatanus'', known as the sycamore in the British Isles and as the sycamore maple in the United States, is a species of maple native to Central Europe and Western Asia. It is a large deciduous, broad-leaved tree, tolerant of wind an ...
''), 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 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 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.


Uses


Food

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 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 ''Theobroma cacao'' (cacao tree or cocoa tree) is a small ( tall) evergreen tree in the Malvaceae family. Its seedscocoa beansare used to make chocolate liquor, cocoa solids, cocoa butter and chocolate. Although the tree is native to the tropi ...
'') are used to make cocoa and chocolate and the berries of coffee trees, ''
Coffea arabica ''Coffea arabica'' (), also known as the Arabica coffee, is a species of flowering plant in the coffee and madder family Rubiaceae. It is believed to be the first species of coffee to have been cultivated and is the dominant cultivar, represe ...
'' and ''
Coffea canephora ''Coffea canephora'' (especially ''C. canephora var. robusta'', so predominantly cultivated that it is often simply termed ''Coffea robusta'', or commonly robusta coffee) is a species of coffee plant that has its origins in central and weste ...
'', are processed to extract the coffee beans. In many rural areas of 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
coconut The coconut tree (''Cocos nucifera'') is a member of the palm tree family (biology), family (Arecaceae) and the only living species of the genus ''Cocos''. The term "coconut" (or the archaic "cocoanut") can refer to the whole coconut palm, ...
s (''Cocos nucifera''), Brazil nuts (''Bertholletia excelsa''), pecans (''Carya illinoinensis''), hazel nuts (''Corylus''),
almond The almond (''Prunus amygdalus'', Synonym (taxonomy)#Botany, syn. ''Prunus dulcis'') is a species of tree from the genus ''Prunus''. Along with the peach, it is classified in the subgenus ''Amygdalus'', distinguished from the other subgenera ...
s (''Prunus dulcis''), walnuts (''Juglans regia''),
pistachio The pistachio (, ; ''Pistacia vera'') is a small to medium-sized tree of the Anacardiaceae, cashew family, originating in Iran. The tree produces nut (fruit)#Culinary definition and uses, seeds that are widely consumed as food. In 2022, world ...
s (''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 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 ''Acer saccharum'', the sugar maple, is a species of flowering plant in the soapberry and lychee family Sapindaceae. It is native to the hardwood forests of eastern Canada and the eastern United States. Sugar maple is best known for being the p ...
'') is used in the production of
maple syrup Maple syrup is a sweet syrup made from the sap of maple trees. In cold climates, these trees store starch in their trunks and roots before winter; the starch is then converted to sugar that rises in the sap in late winter and early spring. Ma ...
. 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 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; a hundred litres are required to make one litre of birch syrup. Various parts of trees are used as spices. These include
cinnamon Cinnamon is a spice obtained from the inner bark of several tree species from the genus ''Cinnamomum''. Cinnamon is used mainly as an aromatic condiment and flavouring additive in a wide variety of cuisines, sweet and savoury dishes, biscuits, b ...
, 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 the clove tree ('' Syzygium aromaticum''). Many trees have flowers rich in
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
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 ''Prunus'' is a genus of flowering plant, flowering trees and shrubs from the family (biology), family Rosaceae. The genus includes plums, cherries, peaches, nectarines, apricots and almonds (collectively Drupe, stonefruit). The genus has a cosm ...
spp.'') can be candied. Sassafras oil 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 ''Ailanthus'' (; derived from ''ailanto,'' an Ambonese Malay, Ambonese word probably meaning "tree of the gods" or "tree of heaven") is a genus of trees belonging to the family Simaroubaceae, in the order Sapindales (formerly Rutales or Geranial ...
'' (in Korean dishes such as bugak) and those of 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 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 . 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.


Fuel

Wood has traditionally been used for fuel, especially in rural areas. In less developed nations it may be 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. Charcoal can be made by slow
pyrolysis Pyrolysis is a process involving the Bond cleavage, separation of covalent bonds in organic matter by thermal decomposition within an Chemically inert, inert environment without oxygen. Etymology The word ''pyrolysis'' is coined from the Gree ...
of wood by heating it in the absence of air in a
kiln A kiln is a thermally insulated chamber, a type of oven, that produces temperatures sufficient to complete some process, such as hardening, drying, or Chemical Changes, chemical changes. Kilns have been used for millennia to turn objects m ...
. The carefully stacked branches, often oak, are burned with a very limited amount of air. The process of converting them into charcoal takes about fifteen hours. Charcoal is used as a fuel in barbecues and by
blacksmith A blacksmith is a metalsmith who creates objects primarily from wrought iron or steel, but sometimes from #Other metals, other metals, by forging the metal, using tools to hammer, bend, and cut (cf. tinsmith). Blacksmiths produce objects such ...
s and has many industrial and other uses.


Timber

Timber, "trees that are grown in order to produce wood" is cut into lumber (sawn wood) for use in construction. Wood has been an important, easily available material for construction since humans started building shelters. Engineered wood products are available which bind the particles, fibres or veneers of wood together with adhesives to form
composite material A composite or composite material (also composition material) is a material which is produced from two or more constituent materials. These constituent materials have notably dissimilar chemical or physical properties and are merged to create a ...
s. Plastics have taken over from wood for some traditional uses. Wood is used in the construction of buildings, bridges, trackways, piles, poles for power lines, masts for boats, pit props, railway sleepers, fencing, hurdles, shuttering for concrete, pipes, scaffolding and pallets. In housebuilding it is used in joinery, for making joists, roof trusses, roofing shingles, thatching, staircases, doors, window frames, floor boards, parquet flooring, panelling and cladding. Wood is used to construct carts, farm implements, boats, dugout canoes and in shipbuilding. It is used for making furniture, tool handles, boxes, ladders, musical instruments, bows, weapons, matches, clothes pegs, brooms, shoes, baskets, turnery, carving, toys, pencils, rollers, cogs, wooden screws, barrels, coffins, skittles, veneers, artificial limbs, oars, skis, wooden spoons, sports equipment and wooden balls. Wood is pulped for paper and used in the manufacture of cardboard and made into engineered wood products for use in construction such as fibreboard, hardboard, chipboard and plywood. The wood of gymnosperms is known as softwood while that of angiosperms is known as
hardwood Hardwood is wood from Flowering plant, angiosperm trees. These are usually found in broad-leaved temperate and tropical forests. In temperate and boreal ecosystem, boreal latitudes they are mostly deciduous, but in tropics and subtropics mostl ...
.


Art

Besides inspiring artists down the centuries, trees have been used to create art. Living trees have been used in
bonsai Bonsai (; , ) is the Japanese art of Horticulture, growing and shaping miniature trees in containers, with a long documented history of influences and native Japanese development over a thousand years, and with unique aesthetics, cultural hist ...
and in tree shaping, and both living and dead specimens have been sculpted into sometimes fantastic shapes.


Bonsai

is the practice of growing and shaping small trees, originating in China as penjing and spreading to Japan more than a thousand years ago, there are also similar practices in other cultures like the living miniature landscapes of Vietnam ''hòn non bộ''. The word ''bonsai'' is often used in English as an
umbrella term Hypernymy and hyponymy are the wikt:Wiktionary:Semantic relations, semantic relations between a generic term (''hypernym'') and a more specific term (''hyponym''). The hypernym is also called a ''supertype'', ''umbrella term'', or ''blanket term ...
for all miniature trees in containers or pots. The purposes of bonsai are primarily contemplation (for the viewer) and the pleasant exercise of effort and ingenuity (for the grower). Bonsai practice focuses on long-term cultivation and shaping of one or more small trees growing in a container, beginning with a cutting, seedling, or small tree of a species suitable for bonsai development. Bonsai can be created from nearly any
perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
woody-stemmed tree or shrub species that produces true branches and can be cultivated to remain small through pot confinement with crown and root pruning. Some
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
are popular as bonsai material because they have characteristics, such as small leaves or needles, that make them appropriate for the compact visual scope of bonsai and a miniature deciduous forest can even be created using such species as Japanese maple, Japanese zelkova or hornbeam.


Tree shaping

Tree shaping is the practice of changing living trees and other woody plants into man made shapes for art and useful structures. There are a few different methods of shaping a tree. There is a gradual method and there is an instant method. The gradual method slowly guides the growing tip along predetermined pathways over time whereas the instant method bends and weaves saplings long into a shape that becomes more rigid as they thicken up. Most artists use grafting of living trunks, branches, and roots, for art or functional structures and there are plans to grow "living houses" with the branches of trees knitting together to give a solid, weatherproof exterior combined with an interior application of straw and clay to provide a
stucco Stucco or render is a construction material made of aggregates, a binder, and water. Stucco is applied wet and hardens to a very dense solid. It is used as a decorative coating for walls and ceilings, exterior walls, and as a sculptural and ...
-like inner surface. Tree shaping has been practised for at least several hundred years, the oldest known examples being the living root bridges built and maintained by the Khasi people of Meghalaya, India using the roots of the rubber tree (''Ficus elastica'').


Bark

Cork is produced from the thick bark of the cork oak ('' Quercus suber''). It is harvested from the living trees about once every ten years in an environmentally sustainable industry. More than half the world's cork comes from Portugal and is largely used to make stoppers for wine bottles. Other uses include floor tiles, bulletin boards, balls, footwear, cigarette tips, packaging, insulation and joints in woodwind instruments. The bark of other varieties of oak has traditionally been used in Europe for the tanning of hides though bark from other species of tree has been used elsewhere. The active ingredient,
tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and Precipitation (chemistry), precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' is widel ...
, is extracted and after various preliminary treatments, the skins are immersed in a series of vats containing solutions in increasing concentrations. The tannin causes the hide to become supple, less affected by water and more resistant to bacterial attack. At least 120 drugs come from plant sources, many of them from the bark of trees.
Quinine Quinine is a medication used to treat malaria and babesiosis. This includes the treatment of malaria due to ''Plasmodium falciparum'' that is resistant to chloroquine when artesunate is not available. While sometimes used for nocturnal leg ...
originates from the cinchona tree (''
Cinchona ''Cinchona'' (pronounced or ) is a genus of flowering plants in the family Rubiaceae containing at least 23 species of trees and shrubs. All are native to the Tropical Andes, tropical Andean forests of western South America. A few species are ...
'') and was for a long time the remedy of choice for the treatment of
malaria Malaria is a Mosquito-borne disease, mosquito-borne infectious disease that affects vertebrates and ''Anopheles'' mosquitoes. Human malaria causes Signs and symptoms, symptoms that typically include fever, Fatigue (medical), fatigue, vomitin ...
. Aspirin was synthesised to replace the sodium salicylate derived from the bark of willow trees ('' Salix'') which had unpleasant side effects. The anti-cancer drug Paclitaxel is derived from taxol, a substance found in the bark of the Pacific yew ('' Taxus brevifolia''). Other tree based drugs come from the paw-paw ('' Carica papaya''), the cassia ( ''Cassia'' spp.), the cocoa tree (''Theobroma cacao''), the tree of life ('' Camptotheca acuminata'') and the downy birch ('' Betula pubescens''). The papery bark of the paper birch ('' Betula papyrifera'') tree was used extensively by Native Americans. Wigwams were covered by it and
canoe A canoe is a lightweight, narrow watercraft, water vessel, typically pointed at both ends and open on top, propelled by one or more seated or kneeling paddlers facing the direction of travel and using paddles. In British English, the term ' ...
s were constructed from it. Other uses included food containers, hunting and fishing equipment, musical instruments, toys and sledges. Nowadays, bark chips, a by-product of the timber industry, are used as a mulch and as a growing medium for epiphytic plants that need a soil-free compost.


Ornamental trees

Trees create a visual impact in the same way as do other landscape features and give a sense of maturity and permanence to park and garden. They are grown for the beauty of their forms, their foliage, flowers, fruit and bark and their siting is of major importance in creating a landscape. They can be grouped informally, often surrounded by plantings of bulbs, laid out in stately avenues or used as specimen trees. As living things, their appearance changes with the season and from year to year. Trees are often planted in town environments where they are known as street trees or amenity trees. They can provide shade and cooling through
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 ...
, absorb greenhouse gases and pollutants, intercept rainfall, and reduce the risk of flooding. Scientific studies show that street trees help cities be more sustainable, and improve the physical and mental wellbeing of the citizens. It has been shown that they are beneficial to humans in creating a sense of well-being and reducing stress. Many towns have initiated tree-planting programmes. In London for example, there is an initiative to plant 20,000 new street trees and to have an increase in tree cover of 5% by 2025, equivalent to one tree for every resident.


Other uses

Latex is a sticky defensive secretion that protects plants against
herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...
s. Many trees produce it when injured but the main source of the latex used to make
natural rubber Rubber, also called India rubber, latex, Amazonian rubber, ''caucho'', or ''caoutchouc'', as initially produced, consists of polymers of the organic compound isoprene, with minor impurities of other organic compounds. Types of polyisoprene ...
is the Pará rubber tree (''Hevea brasiliensis''). Originally used to create bouncy balls and for the waterproofing of cloth, natural rubber is now mainly used in tyres for which synthetic materials have proved less durable. The latex exuded by the balatá tree ('' Manilkara bidentata'') is used to make golf balls and is similar to gutta-percha, made from the latex of the "getah perca" tree '' Palaquium''. This is also used as an insulator, particularly of undersea cables, and in dentistry, walking sticks and gun butts. It has now largely been replaced by synthetic materials. Resin is another plant exudate that may have a defensive purpose. It is a viscous liquid composed mainly of volatile
terpene Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n ≥ 2. Terpenes are major biosynthetic building blocks. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predomi ...
s and is produced mostly by coniferous trees. It is used in varnishes, for making small castings and in
ten-pin bowling Tenpin bowling is a type of bowling in which a bowler bowling form, rolls a bowling ball down a wood or synthetic lane toward ten pins positioned Tetractys, evenly in four rows in an equilateral triangle. The goal is to knock down all ten Bowlin ...
balls. When heated, the terpenes are driven off and the remaining product is called "rosin" and is used by stringed instrumentalists on their bows. Some resins contain
essential oil An essential oil is a concentrated hydrophobic liquid containing volatile (easily evaporated at normal temperatures) chemical compounds from plants. Essential oils are also known as volatile oils, ethereal oils, aetheroleum, or simply as the ...
s and are used in
incense Incense is an aromatic biotic material that releases fragrant smoke when burnt. The term is used for either the material or the aroma. Incense is used for aesthetic reasons, religious worship, aromatherapy, meditation, and ceremonial reasons. It ...
and aromatherapy. Fossilised resin is known as amber and was mostly formed in the Cretaceous (145 to 66 million years ago) or more recently. The resin that oozed out of trees sometimes trapped insects or spiders and these are still visible in the interior of the amber. The camphor tree ('' Cinnamomum camphora'') produces an essential oil and the eucalyptus tree (''
Eucalyptus globulus ''Eucalyptus globulus'', commonly known as southern blue gum or blue gum, is a species of flowering plant in the family Myrtaceae. It is a tall, evergreen tree Endemism, endemic to southeastern Australia. This ''Eucalyptus'' species has most ...
'') is the main source of eucalyptus oil which is used in medicine, as a fragrance and in industry.


Threats


Individual trees

Dead trees pose a safety risk, especially during high winds and severe storms, and removing dead trees involves a financial burden, whereas the presence of healthy trees can clean the air, increase property values, and reduce the temperature of the built environment and thereby reduce building cooling costs. During times of drought, trees can fall into water stress, which may cause a tree to become more susceptible to disease and insect problems, and ultimately may lead to a tree's death. Irrigating trees during dry periods can reduce the risk of water stress and death.


Conservation

About a third of all tree species, some twenty thousand, are included in the
IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
Red List of Threatened Species. Of those, over eight thousand are globally threatened, including at least 1400 which are classed as "critically endangered".


Mythology

Trees have been venerated since time immemorial. To the ancient
Celts The Celts ( , see Names of the Celts#Pronunciation, pronunciation for different usages) or Celtic peoples ( ) were a collection of Indo-European languages, Indo-European peoples. "The Celts, an ancient Indo-European people, reached the apoge ...
, certain trees, especially the oak, ash and thorn, held special significance as providing fuel, building materials, ornamental objects and weaponry. Other cultures have similarly revered trees, often linking the lives and fortunes of individuals to them or using them as oracles. In
Greek mythology Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...
, dryads were believed to be shy nymphs who inhabited trees. The Oubangui people of west Africa plant a tree when a child is born. As the tree flourishes, so does the child but if the tree fails to thrive, the health of the child is considered at risk. When it flowers it is time for marriage. Gifts are left at the tree periodically and when the individual dies, their spirit is believed to live on in the tree. Trees have their roots in the ground and their trunk and branches extended towards the sky. This concept is found in many of the world's religions as a tree which links the underworld and the earth and holds up the heavens. In
Norse mythology Norse, Nordic, or Scandinavian mythology, is the body of myths belonging to the North Germanic peoples, stemming from Old Norse religion and continuing after the Christianization of Scandinavia as the Nordic folklore of the modern period. The ...
, Yggdrasil is a central cosmic tree whose roots and branches extend to various worlds. Various creatures live on it.Lindow, John (2001). ''Norse Mythology: A Guide to the Gods, Heroes, Rituals, and Beliefs'', pp. 319–322. Oxford University Press. In India, Kalpavriksha is a wish-fulfilling tree, one of the nine jewels that emerged from the primitive ocean. Icons are placed beneath it to be worshipped, tree nymphs inhabit the branches and it grants favours to the devout who tie threads round the trunk. Democracy started in North America when the Great Peacemaker formed the Iroquois Confederacy, inspiring the warriors of the original five American nations to bury their weapons under the Tree of Peace, an eastern white pine ('' Pinus strobus''). In the creation story in the Bible, the tree of life and the knowledge of good and evil was planted by God in the
Garden of Eden In Abrahamic religions, the Garden of Eden (; ; ) or Garden of God ( and ), also called the Terrestrial Paradise, is the biblical paradise described in Genesis 2–3 and Ezekiel 28 and 31.. The location of Eden is described in the Book of Ge ...
. In the cosmology of the Meso-American Maya culture, the
ceiba ''Ceiba'' is a genus of trees in the family Malvaceae, native to Tropics, tropical and Subtropics, subtropical areas of the Americas (from Mexico and the Caribbean to northern Argentina) and tropical West Africa. Some species can grow to tall ...
tree connects the planes of both the sky and the underworld, known as Xibalba, with the terrestrial world. As such, the ceiba tree is a national symbol in
Guatemala Guatemala, officially the Republic of Guatemala, is a country in Central America. It is bordered to the north and west by Mexico, to the northeast by Belize, to the east by Honduras, and to the southeast by El Salvador. It is hydrologically b ...
. Sacred groves exist in China, India, Africa and elsewhere. They are places where the deities live and where all the living things are either sacred or are companions of the gods. Folklore lays down the supernatural penalties that will result if desecration takes place for example by the felling of trees. Because of their protected status, sacred groves may be the only relicts of ancient forest and have a biodiversity much greater than the surrounding area. Some Ancient Indian
tree deities A tree deity or tree spirit is a nature deity related to a tree. Such deities are present in many cultures. They are usually represented as a young woman, often connected to ancient fertility and tree worship lore.Heinrich Zimmer, ''Myths and Sym ...
, such as Puliyidaivalaiyamman, the Tamil deity of the tamarind tree, or Kadambariyamman, associated with the cadamba tree, were seen as manifestations of a goddess who offers her blessings by giving fruits in abundance.


Superlative trees

Trees have a theoretical maximum height of about 120 to 138 meters (393.7 to 426.5 ft). The primary reason for this is their limited ability to siphon water up the trunk, meaning if a tree is too tall, it will die due to
desiccation Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
. The tallest known tree on earth is believed to be a coast redwood (''
Sequoia sempervirens ''Sequoia sempervirens'' ()''Sunset Western Garden Book,'' 1995: 606–607 is the sole living species of the genus ''Sequoia (genus), Sequoia'' in the cypress family Cupressaceae (formerly treated in Taxodiaceae). Common names include coast ...
'') at Redwood National Park, California. It has been named Hyperion and is tall. In 2006, it was reported to be tall. The tallest known broad-leaved tree is a mountain ash ('' Eucalyptus regnans'') growing in Tasmania with a height of . The largest tree by volume is believed to be a giant sequoia (''
Sequoiadendron giganteum ''Sequoiadendron giganteum'' (also known as the giant sequoia, giant redwood, Sierra redwood or Wellingtonia) is a species of coniferous tree, classified in the family Cupressaceae in the subfamily Sequoioideae. Giant sequoia specimens are the la ...
'') known as the General Sherman Tree in the Sequoia National Park in Tulare County, California. Only the trunk is used in the calculation and the volume is estimated to be . The oldest living tree with a verified age is also in California. It is a Great Basin bristlecone pine (''
Pinus longaeva ''Pinus longaeva'' (commonly referred to as the Great Basin bristlecone pine, intermountain bristlecone pine, or western bristlecone pine) is a long-living species of bristlecone pine tree found in the higher mountains of California, Nevada, and ...
'') growing in the White Mountains. It has been dated by drilling a core sample and counting the annual rings. It is estimated to currently be years old. A little farther south, at Santa Maria del Tule,
Oaxaca Oaxaca, officially the Free and Sovereign State of Oaxaca, is one of the 32 states that compose the political divisions of Mexico, Federative Entities of the Mexico, United Mexican States. It is divided into municipalities of Oaxaca, 570 munici ...
, Mexico, is the tree with the broadest trunk. It is a Montezuma cypress ('' Taxodium mucronatum'') known as Árbol del Tule and its diameter at breast height is giving it a girth of . The tree's trunk is far from round and the exact dimensions may be misleading as the circumference includes much empty space between the large buttress roots.


See also


Notes


References


Sources

* {{Authority control Forest ecology Plant morphology Plant life-forms Plants by habit