Roots (
Pinus resinosa) with spread of roots visible, as a result of soil erosion 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 to emerge from the seed during the process of
germination. This develops into a
taproot which goes straight downwards. Within a few weeks
lateral roots branch out of the side of this and grow horizontally through the upper layers of the soil. In most trees, the taproot eventually withers away and the wide-spreading laterals remain. Near the tip of the finer roots are single cell
root hairs. These are in immediate contact with the soil particles and can absorb water and nutrients such as
potassium in solution. The roots require oxygen to
respire and only a few species such as
mangroves and the
pond cypress (
Taxodium ascendens) can live in permanently waterlogged soil. In the soil, the roots encounter the
hyphae of fungi. Many of these are known as
mycorrhiza and form a
mutualistic relationship with the tree roots. Some are specific to a single tree species, which will not flourish in the absence of its mycorrhizal associate. Others are generalists and associate with many species. The tree acquires minerals such as
phosphorus from the fungus, while the fungus obtains the
carbohydrate products of photosynthesis from the tree. The hyphae of the fungus can link different trees and a network is formed, transferring nutrients and signals from one place to another. The fungus promotes growth of the roots and helps protect the trees against predators and pathogens. It can also limit damage done to a tree by pollution as the fungus accumulate
heavy metals within its tissues. Fossil evidence shows that roots have been associated with mycorrhizal fungi since the early
Paleozoic, four hundred million years ago, when the first
vascular plants colonised dry land. '') Some trees such as
Alder (
Alnus species) have a
symbiotic relationship with
Frankia species, a filamentous bacterium that can fix nitrogen from the air, converting it into
ammonia. They have
actinorhizal root nodules on their roots in which the bacteria live. This process enables the tree to live in low nitrogen habitats where they would otherwise be unable to thrive. The plant hormones called
cytokinins initiate root nodule formation, in a process closely related to mycorrhizal association. It has been demonstrated that some trees are interconnected through their root system, forming a colony. The interconnections are made by the
inosculation process, a kind of natural
grafting or welding of vegetal tissues. The tests to demonstrate this networking are performed by injecting chemicals, sometimes
radioactive, into a tree, and then checking for its presence in neighbouring trees. The roots are, generally, an underground part of 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. 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. (
Taxus baccata) showing 27 annual growth rings, pale
sapwood and dark
heartwood 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 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, a
polysaccharide, and most of the remainder is lignin, a complex
polymer. A transverse section through a tree trunk or a horizontal core will show concentric circles of lighter or darker wood – tree rings. 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, 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. Synthesis in the leaf of a
plant hormone called
auxin also ceases. This causes the cells at the junction of the
petiole and the twig to weaken until the joint breaks and the leaf floats to the ground. In tropical and subtropical regions, many trees keep their leaves all year round. Individual leaves may fall intermittently and be replaced by new growth but most leaves remain intact for some time. Other tropical species and those in arid regions may shed all their leaves annually, such as at 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.
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 ), ash (Fraxinus) and maple (Acer'') 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, 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. These float on the water and may become lodged on emerging mudbanks and successfully take root.
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, and also enriches the forest floor with wood ash and removes competing vegetation. Similarly, a number of angiosperms including
Acacia cyclops and
Acacia mangium have seeds that germinate better after exposure to high temperatures. The single extant species of
Ginkgophyta (
Ginkgo biloba) has fleshy seeds produced at 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 ==