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botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
, a tree is a
perennial plant A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
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 In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of ...
, plants that are usable as
lumber Lumber is wood that has been processed into dimensional lumber, including beams and planks or boards, a stage in the process of wood production. Lumber is mainly used for construction framing, as well as finishing (floors, wall panels, wi ...
or plants above a specified height. In wider definitions, the taller palms,
tree ferns The tree ferns are arborescent (tree-like) ferns that grow with a trunk elevating the fronds above ground level, making them trees. Many extant tree ferns are members of the order Cyatheales, to which belong the families Cyatheaceae (scaly tre ...
,
bananas A banana is an elongated, edible fruit – botanically a berry – produced by several kinds of large herbaceous flowering plants in the genus ''Musa''. In some countries, bananas used for cooking may be called "plantains", distinguis ...
, and
bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, bu ...
s are also trees. Trees are not a
taxonomic group In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
but include a 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 (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s or hardwoods; of the rest, many are
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s or softwoods. Trees tend to be long-lived, some reaching several thousand years old. Trees have been in existence for 370 million years. It is estimated that there are some three trillion mature trees in the world. A tree typically has many secondary branches supported clear of the ground by the trunk. This trunk typically contains
woody tissue Xylem is one of the two types of transport tissue in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
for strength, and
vascular tissue Vascular tissue is a complex conducting 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. The ...
to carry materials from one part of the tree to another. For most trees it 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 focusing ...
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 Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
, providing the food for the tree's growth and development. Trees usually reproduce using seeds. Flowers and fruit may be present, but some trees, such as conifers, instead have pollen cones and seed cones. Palms, bananas, and bamboos also produce seeds, but tree ferns produce
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s instead. Trees play a significant role in reducing
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is distin ...
and moderating the
climate Climate is the long-term weather pattern in an area, 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 meteorologi ...
. They remove
carbon dioxide Carbon dioxide (chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is transpar ...
from the
atmosphere An atmosphere () is a layer of gas or layers of gases that envelop a planet, and is held in place by the gravity of the planetary body. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A s ...
and store large quantities of
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent In chemistry, the valence (US spelling) or valency (British spelling) of an element is the measure of its combining capacity with o ...
in their tissues. Trees and forests provide a habitat for many species of animals and plants.
Tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equatori ...
s are among the most
biodiverse Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
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 parts 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 grove Sacred groves or sacred woods are groves of trees and have special religious importance within a particular culture. Sacred groves feature in various cultures throughout the world. They were important features of the mythological landscape and ...
s in various cultures, and they play a role in many of the world's
mythologies Myth is a folklore genre consisting of narratives that play a fundamental role in a society, such as foundational tales or origin myths. Since "myth" is widely used to imply that a story is not objectively true, the identification of a narrati ...
.


Definition

Although "tree" is a term of common parlance, there is no universally recognised precise definition of what a tree is, either
botanically Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...
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 leaves or branches at some distance above the ground. Trees are also typically defined by height, with smaller plants from being called shrubs, so the minimum height of a tree is only loosely defined. Large
herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition of t ...
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 In botany, secondary growth is the growth that results from cell division in the cambia or lateral meristems and that causes the stems and roots to thicken, while primary growth is growth that occurs as a result of cell division at the tips of ...
, 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 Monocotyledons (), commonly referred to as monocots, ( Lilianae '' sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
may be considered trees under a slightly looser definition; while the
Joshua tree ''Yucca brevifolia'' is a plant species belonging to the genus ''Yucca''. It is tree-like in habit, which is reflected in its common names: Joshua tree, yucca palm, tree yucca, and palm tree yucca. This monocotyledonous tree is native to the ar ...
, 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 change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
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 po ...
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 (often also called a 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 ...
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 The subarctic zone is a region in the Northern Hemisphere immediately south of the true Arctic, north of humid continental regions and covering much of Alaska, Canada, Iceland, the north of Scandinavia, Siberia, and the Cairngorms. Generally, ...
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 Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and paral ...
. 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 biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek wo ...
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 (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s or hardwoods. Of the rest, many are
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό ...
s or softwood trees; these include conifers,
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
s, ginkgophytes and
gnetales Gnetophyta () is a division of plants (alternatively considered the subclass Gnetidae or order Gnetales), grouped within the gymnosperms (which also includes conifers, cycads, and ginkgos), that consists of some 70 species across the three reli ...
, which produce seeds which are not enclosed in fruits, but in open structures such as pine
cones A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines conn ...
, and many have tough waxy leaves, such as pine needles. Most angiosperm trees are
eudicots The eudicots, Eudicotidae, or eudicotyledons are a clade of flowering plants mainly characterized by having two seed leaves upon germination. The term derives from Dicotyledons. Traditionally they were called tricolpates or non-magnoliid dico ...
, the "true dicotyledons", so named because the seeds contain two
cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The num ...
s or seed leaves. There are also some trees among the old lineages of flowering plants called basal angiosperms or paleodicots; these include '' Amborella'', ''
Magnolia ''Magnolia'' is a large genus of about 210 to 340The number of species in the genus ''Magnolia'' depends on the taxonomic view that one takes up. Recent molecular and morphological research shows that former genera ''Talauma'', ''Dugandiodendro ...
'',
nutmeg Nutmeg is the seed or ground spice of several species of the genus ''Myristica''. ''Myristica fragrans'' (fragrant nutmeg or true nutmeg) is a dark-leaved evergreen tree cultivated for two spices derived from its fruit: nutmeg, from its seed, an ...
and
avocado The avocado (''Persea americana'') is a medium-sized, evergreen tree in the laurel family ( Lauraceae). It is native to the Americas and was first domesticated by Mesoamerican tribes more than 5,000 years ago. Then as now it was prized for ...
, while trees such as bamboo, palms and bananas are
monocots Monocotyledons (), commonly referred to as monocots, ( Lilianae '' sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
. 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, as relatively tall plants, need to draw water up the stem through the xylem from the roots by the suction produced as water evaporates from the leaves. 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 The vascular cambium is the main growth tissue in the stems and roots of many plants, specifically in dicots such as buttercups and oak trees, gymnosperms such as pine trees, as well as in certain other vascular plants. It produces secondary xy ...
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 Cork cambium (pl. cambia or cambiums) is a tissue found in many vascular plants as a part of the epidermis. It is one of the many layers of bark, between the cork and primary phloem. The cork cambium is a lateral meristem and is responsible for ...
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 foliage that remains green and functional through more than one growing season. This also pertains to plants that retain their foliage only in warm climates, and contrasts with deciduous plants, whic ...
, 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 leaves, usually in the autumn; to the shedding of petals, ...
, 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 Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains furt ...
'' and ''
Pseudolarix ''Pseudolarix'' is a genus of coniferous trees in the pine family Pinaceae containing three species, the extant '' Pseudolarix amabilis'' and the extinct species '' Pseudolarix japonica'' and '' Pseudolarix wehrii''. ''Pseudolarix'' species are ...
'') are deciduous, dropping their needles each autumn, and some species of cypress (''
Glyptostrobus ''Glyptostrobus'' is a small genus of conifers in the family Cupressaceae (formerly in the family Taxodiaceae). The sole living species, '' Glyptostrobus pensilis'', is native to subtropical southeastern China, from Fujian west to southeast Yu ...
'', ''
Metasequoia ''Metasequoia'', or dawn redwoods, is a genus of fast-growing deciduous trees, one of three species of conifers known as redwoods. The living species '' Metasequoia glyptostroboides'' is native to Lichuan county in Hubei province, China. Althou ...
'' and ''
Taxodium ''Taxodium'' is a genus of one to three species (depending on taxonomic opinion) of extremely flood-tolerant conifers in the cypress family, Cupressaceae. The generic name is derived from the Latin word ''taxus'', meaning "yew", and the Greek ...
'') shed small leafy shoots annually in a process known as
cladoptosis Cladoptosis (Ancient Greek "branch", "falling" oun sometimes pronounced with the p silent) is the regular shedding of branches. It is the counterpart for branches of the familiar process of regular leaf shedding by deciduous trees. As in leaf ...
. The
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, partic ...
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 Canopy may refer to: Plants * Canopy (biology), aboveground portion of plant community or crop (including forests) * Canopy (grape), aboveground portion of grapes Religion and ceremonies * Baldachin or canopy of state, typically placed over an ...
. A sapling is a young tree. Many tall palms are herbaceous monocots; these 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 The order Cyatheales, which includes most tree ferns, is a taxonomic order of the fern class, Polypodiopsida. No clear morphological features characterize all of the Cyatheales, but DNA sequence data indicate the order is monophyletic. Some sp ...
, have tall straight trunks, growing up to , but these are composed not of wood but of rhizomes which grow vertically and are covered by numerous
adventitious roots Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living pla ...
.


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 In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
, and 0.74 trillion (24%) in the coniferous
boreal forests Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruces, ...
. 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%. In suitable environments, such as the
Daintree Rainforest The Daintree Rainforest is a region on the northeast coast of Queensland, Australia, north of Mossman and Cairns. At around , the Daintree is a part of the largest continuous area of tropical rainforest on the Australian continent. The Daintr ...
in
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
, or the mixed
podocarp Podocarpaceae is a large family of mainly Southern Hemisphere conifers, known in English as podocarps, comprising about 156 species of evergreen trees and shrubs.James E. Eckenwalder. 2009. ''Conifers of the World''. Portland, Oregon: Timber Pr ...
and
broadleaf forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
of
Ulva Island, New Zealand Ulva Island (from gd, Eilean na Ulbha) is a small island about long lying within Paterson Inlet, which is part of Stewart Island/Rakiura in New Zealand. It has an area of , of which almost all (261 ha) is part of Rakiura National Park. ...
, 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 In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
regions, conifers often predominate; a widely distributed climax community in the far north of the northern hemisphere is moist
taiga Taiga (; rus, тайга́, p=tɐjˈɡa; relates to Mongolic and Turkic languages), generally referred to in North America as a boreal forest or snow forest, is a biome characterized by coniferous forests consisting mostly of pines, spruce ...
or northern coniferous forest (also called boreal forest). Taiga is the world's largest land
biome A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
, 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 Temperate broadleaf and mixed forest is a temperate climate terrestrial habitat type defined by the World Wide Fund for Nature, with broadleaf tree ecoregions, and with conifer and broadleaf tree mixed coniferous forest ecoregions. These ...
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 over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as e ...
'' 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 atmospheric circulation and precipitation associated with annual latitudinal oscil ...
or monsoon-like climate, where a drier part of the year alternates with a wet period as in the
Amazon rainforest The Amazon rainforest, Amazon jungle or ; es, Selva amazónica, , or usually ; french: Forêt amazonienne; nl, Amazoneregenwoud. In English, the names are sometimes capitalized further, as Amazon Rainforest, Amazon Forest, or Amazon Jungle. ...
, 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 ''Adansonia'' is a genus made up of eight species of medium-to-large deciduous trees known as baobabs ( or ). They are placed in the Malvaceae family, subfamily Bombacoideae. They are native to Madagascar, mainland Africa, and Australia.Trop ...
are well adapted to living in such areas.


Parts and function


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 In botany, the radicle is the first part of a seedling (a growing plant embryo) to emerge from the seed during the process of germination. The radicle is the embryonic root of the plant, and grows downward in the soil (the shoot emerges from ...
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 (embryo ...
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 taproo ...
which goes straight downwards. Within a few weeks
lateral root Lateral roots, emerging from the pericycle (meristematic tissue), extend horizontally from the primary root (radicle) and over time makeup the iconic branching pattern of root systems. They contribute to anchoring the plant securely into the soil ...
s 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 hair Root hair, or absorbent hairs, are outgrowths of epidermal cells, specialized cells at the tip of a plant root. They are lateral extensions of a single cell and are only rarely branched. They are found in the region of maturation, of the root. Root ...
s. These are in immediate contact with the soil particles and can absorb water and nutrients such as
potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosph ...
in solution. The roots require oxygen to respire and only a few species such as
mangrove A mangrove is a shrub or tree that grows in coastal saline water, saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evoluti ...
s and the
pond cypress ''Taxodium ascendens'', also known as pond cypress, is a deciduous conifer of the genus ''Taxodium'', native to North America. Many botanists treat it as a variety of bald cypress, ''Taxodium distichum'' (as ''T. distichum'' var. ''imbricatum'' ...
(''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 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 with the symbol P and atomic number 15. Elemental phosphorus exists in two major forms, white phosphorus and red phosphorus, but because it is highly reactive, phosphorus is never found as a free element on Ear ...
from the fungus, while the fungus obtains the
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or ma ...
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 earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...
, four hundred million years ago, when the first
vascular plant Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They ...
s colonised dry land. Some trees such as
Alder Alders are trees comprising the genus ''Alnus'' in the birch family Betulaceae. The genus comprises about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few sp ...
(''Alnus'' species) have a symbiotic relationship with ''
Frankia ''Frankia'' is a genus of nitrogen-fixing bacteria that live in symbiosis with actinorhizal plants, similar to the ''Rhizobium'' bacteria found in the root nodules of legumes in the family Fabaceae. ''Frankia'' also initiate the forming of root ...
'' species, a filamentous bacterium that can fix nitrogen from the air, converting it into
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...
. They have
actinorhizal Actinorhizal plants are a group of angiosperms characterized by their ability to form a symbiosis with the nitrogen fixing actinomycetota ''Frankia''. This association leads to the formation of nitrogen-fixing root nodules. Classification Actinorh ...
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 growth and differentiation, but also affect apical dominance, axillary bud growth, and le ...
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 Inosculation is a natural phenomenon in which trunks, branches or roots of two trees grow together in a manner biologically similar to the artificial process of grafting. The term is derived from the Latin roots ''in'' + '' ōsculārī'', "to ...
process, a kind of natural
grafting Grafting or graftage is a 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 rootstock. The succ ...
or welding of vegetal tissues. The tests to demonstrate this networking are performed by injecting chemicals, sometimes
radioactive Radioactive decay (also known as nuclear decay, radioactivity, radioactive disintegration, or nuclear disintegration) is the process by which an unstable atomic nucleus loses energy by radiation. A material containing unstable nuclei is consi ...
, 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 Red mangrove may refer to at least three plant species: * ''Rhizophora mangle'' * ''Rhizophora mucronata ''Rhizophora mucronata'' (loop-root mangrove, red mangrove or Asiatic mangrove) is a species of mangrove found on coasts and river banks in ...
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 ''Ficus benghalensis'', commonly known as the banyan, banyan fig and Indian banyan, is a tree native to the Indian Subcontinent. Specimens in India are among the largest trees in the world by canopy coverage. It also known as the " strangler fig" ...
. Many large trees have
buttress root Buttress roots also known as plank roots are large, wide roots on all sides of a shallowly rooted tree. Typically, they are found in nutrient-poor tropical forest soils that may not be very deep. They prevent the tree from falling over (hence t ...
s 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 Aerial roots are roots above the ground. They are almost always adventitious. They are found in diverse plant species, including epiphytes such as orchids (''Orchidaceae''), tropical coastal swamp trees such as mangroves, banyan figs (''Fic ...
, 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 is an impermeable buoyant material, the phellem layer of bark tissue that is harvested for commercial use primarily from ''Quercus suber'' (the cork oak), which is native to southwest Europe and northwest Africa. Cork is composed of sube ...
(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 Cork cambium (pl. cambia or cambiums) is a tissue found in many vascular plants as a part of the epidermis. It is one of the many layers of bark, between the cork and primary phloem. The cork cambium is a lateral meristem and is responsible for ...
or phellogen. The
London plane London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a major se ...
(''Platanus × acerifolia'') periodically sheds its bark in large flakes. Similarly, the bark of the
silver birch ''Betula pendula'', commonly known as silver birch, warty birch, European white birch, or East Asian white birch, is a species of tree in the family Betulaceae, native to Europe and parts of Asia, though in southern Europe, it is only found ...
(''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. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
(''Pinus'' species) the bark exudes sticky
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
which deters attackers whereas in
rubber trees ''Hevea brasiliensis'', the Pará rubber tree, ''sharinga'' tree, seringueira, or most commonly, rubber tree or rubber plant, is a flowering plant belonging to the spurge family Euphorbiaceae originally native to the Amazon basin, but is now pan ...
(''Hevea brasiliensis'') it is a milky
latex Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latexes are found in nature, but synthetic latexes are common as well. In nature, latex is found as a milky fluid found in 10% of all flowering plants (angiosperms ...
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 A pteridophyte is a vascular plant (with xylem and phloem) that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as " cryptogams", meaning that their means of reproduction is hidden. Ferns ...
,
Arecales Arecales is an order of flowering plants. The order has been widely recognised only for the past few decades; until then, the accepted name for the order including these plants was Principes. Taxonomy The APG IV system of 2016 places Dasypogo ...
,
Cycadophyta Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
and
Poales The Poales are a large order of flowering plants in the monocotyledons, and includes families of plants such as the grasses, bromeliads, and sedges. Sixteen plant families are currently recognized by botanists to be part of Poales. Descriptio ...
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 Dutch elm disease (DED) is caused by a member of the sac fungi (Ascomycota) affecting elm trees, and is spread by elm bark beetles. Although believed to be originally native to Asia, the disease was accidentally introduced into Americas, America ...
is caused by a fungus (''
Ophiostoma ''Ophiostoma'' is a genus of fungi within the family Ophiostomataceae. It was circumscribed in 1919 by mycologists Hans Sydow and Paul Sydow. Species *''Ophiostoma adjuncti'' *'' Ophiostoma ainoae'' *'' Ophiostoma allantosporum'' *'' Ophiosto ...
'' species) carried from one
elm Elms are deciduous and semi-deciduous trees comprising the flowering plant genus ''Ulmus'' in the plant family Ulmaceae. They are distributed over most of the Northern Hemisphere, inhabiting the temperate and tropical-montane regions of North ...
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 Phloem (, ) is the living biological tissue, tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This tran ...
and this is involved in the transport of the
sap Sap is a fluid transported in xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a separ ...
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 in vascular plants, the other being phloem. The basic function of xylem is to transport water from roots to stems and leaves, but it also transports nutrients. The word ''xylem'' is derived from ...
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 Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin t ...
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 formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
, a polysaccharide, and most of the remainder is lignin, a complex
polymer A polymer (; Greek '' poly-'', "many" + ''-mer'', "part") is a substance or material consisting of very large molecules called macromolecules, composed of many repeating subunits. Due to their broad spectrum of properties, both synthetic a ...
. 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
bud In botany, a bud is an undeveloped or embryonic shoot and normally occurs in the axil of a leaf or at the tip of a stem. Once formed, a bud may remain for some time in a dormant condition, or it may form a shoot immediately. Buds may be spec ...
s to protect the
meristem The meristem is a type of tissue found in plants. It consists of undifferentiated cells (meristematic cells) capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells conti ...
, 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 over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as e ...
, 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 Lammas growth, also called Lammas leaves, Lammas flush, second shoots, or summer shoots, is a season of renewed growth in some trees in temperate regions put on in July and August (if in the northern hemisphere, January and February if in the sout ...
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.


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, "plant stone") are rigid, microscopic structures made of silica, found in some plant tissues and persisting after the decay of the plant. These plants take up silica from the soil, whereupon it is deposited within different ...
s, lignins,
tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'' ...
s or poisons 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 and the red and yellow pigments already present in the blades become apparent. Synthesis in the leaf of a
plant hormone Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
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
Phylloclade Phylloclades and cladodes are flattened, photosynthetic shoots, which are usually considered to be modified branches. The two terms are used either differently or interchangeably by different authors. '' Phyllocladus'', a genus of conifer, is nam ...
s – modified stem structures – as seen in the genus ''
Phyllocladus ''Phyllocladus'', the celery pines, is a small genus of conifers, now usually treated in the family Podocarpaceae.James E. Eckenwalder. 2009. ''Conifers of the World''. Timber Press: Portland, OR, USA. .'' ''Species occur mainly in New Zealand, ...
''.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 Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, ...
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 In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors ...
. 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'' (giant sequoia; also known as giant redwood, Sierra redwood, Sierran redwood, California big tree, Wellingtonia or simply big treea nickname also used by John Muir) is the sole living species in the genus ''Sequoiade ...
'', 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 A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech-oak family Fagaceae. The genus ''Betula'' contains 30 ...
are small and have papery wings to aid dispersal by the wind.
Ash Ash or ashes are the solid remnants of fires. Specifically, ''ash'' refers to all non-aqueous, non- gaseous residues that remain after something burns. In analytical chemistry, to analyse the mineral and metal content of chemical samples, ash ...
trees and
maple ''Acer'' () is a genus of trees and shrubs commonly known as maples. The genus is placed in the family Sapindaceae.Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 nd more or less continuously updated since http ...
s have larger seeds with blade shaped wings which spiral down to the ground when released. The Ceiba pentandra, kapok tree has cottony threads to catch the breeze. 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, 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 Delonix regia, flame tree ''Delonix regia'' does not rely on fire but 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 the seeds on the surface of water. Mangroves often grow in water and some species have propagules, which are buoyant fruits with seeds that start germinating before becoming detached 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 Crataegus, 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. Nut (fruit), Nuts may be gathered by animals such as squirrels that hoarding (animal behavior), cache any not immediately consumed. Many of these caches are never revisited, the nut-casing softens with rain and frost, and the seed germinates 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 single extant species of Ginkgophyta (''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 period. The first tree may have been ''Wattieza'', fossils of which have been found in New York (state), New York State 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 period. The gymnosperms include conifers, cycads, gnetales and ginkgos and these may have appeared as a result of a Gene duplication#As an evolutionary event, whole genome duplication event which took place about 319 million years ago. Ginkgophyta 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 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 the Cretaceous period. These began to dominance (ecology), 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 ice ages 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 ice age.


Ecology

Trees are an important part of the terrestrial ecosystem, providing essential habitats including many kinds of forest for communities of organisms. Epiphyte, Epiphytic plants such as ferns, some mosses, liverworts, orchids and some species of parasite, 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 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 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 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 resistance, nonhost neighbours from its insect herbivore, 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'') are used to make cocoa solids, 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 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, 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 List of vegetable oils#Nut oils, 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'') is most often used in the production of a sweet liquid, 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 the inserted spigots. It is piped to a sugarhouse where it is heated to concentrate it and improve its flavour. Similarly in northern Europe the spring rise in the Birch sap, 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, 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 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 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 food, 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. Smoking (cooking), 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 Curing (food preservation), 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 of wood by heating it in the absence of air in a kiln. 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 blacksmiths 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 Wood veneer, veneers of wood together with adhesives to form composite materials. 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 Medium-density fibreboard, fibreboard, hardboard, Particle board, chipboard and plywood. The wood of conifers is known as softwood while that of broad-leaved trees is hardwood.


Art

Besides inspiring artists down the centuries, trees have been used to create art. Living trees have been used in bonsai and in tree shaping, and both living and dead specimens have been sculpted into sometimes fantastic shapes.


Bonsai

is the practice of Hòn Non Bộ, ''hòn non bộ'' originated in China and spread to Japan more than a thousand years ago, there are 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 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 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 List of species used in bonsai, species 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, Zelkova serrata, 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-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, Khasi people of Meghalaya, India using the roots of the Ficus elastica, rubber tree (''Ficus elastica'').


Bark

Cork (material), 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 (leather), 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 precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'' ...
, 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 medicinal plant, plant sources, many of them from the bark of trees. Quinine originates from the cinchona tree (''Cinchona'') and was for a long time the remedy of choice for the treatment of malaria. 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 (genus), ''Cassia'' spp.), the cocoa tree (''Theobroma cacao''), the tree of life (''Camptotheca acuminata'') and the downy birch (''Betula pubescens''). The papery bark of the white birch tree (''Betula papyrifera'') was used extensively by Indigenous peoples of the Americas, Native Americans. Wigwams were covered by it and canoes 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, 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 herbivores. Many trees produce it when injured but the main source of the latex used to make natural rubber 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 terpenes and is produced mostly by coniferous trees. It is used in varnishes, for making small castings and in ten-pin bowling balls. When heated, the terpenes are driven off and the remaining product is called "rosin" and is used by stringed instrumentalists on their Bow (music), bows. Some resins contain essential oils and are used in incense 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'') 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 moisture stress, 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 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 Celtic tree worship, Celts, certain trees, especially the oak, Fraxinus, ash and Crataegus, 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, 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, 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, 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 (biblical), tree of life and the knowledge of good and evil was planted by God in the Garden of Eden. 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 deity, tree deities, such as Puliyidaivalaiyamman, the Tamil People, Tamil deity of the tamarind, tamarind tree, or Kadambariyamman, associated with the kadamba 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 , but the tallest known specimen on earth is believed to be a coast redwood (''Sequoia sempervirens'') at Redwood National Park, California. It has been named Hyperion (tree), 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'') 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'') growing in the White Mountains (California), 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, 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

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