Buds and growth
Dormant magnolia (Magnolia sp.) bud
Main article: Bud
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.[58]
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.[58] 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.[59]
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.[60] 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.[61]
Leaves
Buds, leaves, flowers and fruit of oak (Quercus robur)
Buds, leaves and reproductive structures of white fir (Abies alba)
Main article: Leaf
Leaves are structures specialized for photosynthesis and are arranged on the tree in such a way as to maximise their exposure to light without shading each other. They are an important investment by the tree and may be thorny or contain phytoliths, lignins, tannins or poisons to discourage herbivory. Trees have evolved leaves in 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.[62] 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.[63] Many deciduous trees flower before the new leaves emerge.[64] A few trees do not have true leaves but instead have structures with similar external appearance such as Phylloclades – modified stem structures[65] – as seen in the genus Phyllocladus.[66]
Reproduction
Form, leaves and reproductive structures of queen sago Cycas circinalis
Main article: Plant reproduction
Tree forms are found in a wide range of plants and their reproductive strategies are substantially the same as shrub or herbaceous plant forms. Many trees are wind pollinated which may be an evolutionary adaptation to take advantage of increased wind speeds high above the ground, particularly in the case of those that produce pollen before the leaves emerge.[67] A vast quantity of pollen is produced because of the low likelihood of any particular grain landing on an appropriate female flower. Wind-pollinated flowers of broad-leaved trees are characterised by a lack of showy parts, no scent and a copious production of pollen, often with separate male and female flowers, or separate male and female trees. The male flowers may be high up in the tree, often in the form of dangling catkins. The female flowers may be lower down the tree. The pollen of pine trees contains air sacs which give it buoyancy and it has been known to travel as far as 800 kilometres (500 mi).[68] Tree pollen can cause allergies and hay fever.[69]
Seeds
Main article: Seed
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.[70] The great diversity in tree fruits and seeds reflects the many different ways that tree species have evolved to disperse their offspring.
The single extant species of Ginkgophyta (Ginkgo biloba) has fleshy seeds produced at the ends of short branches on female trees,[71] and Gnetum, a tropical and subtropical group of gymnosperms produce seeds at the tip of a shoot axis.[72] 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.[73] 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.[74] Similarly, a number of angiosperms including Acacia cyclops and Acacia mangium have seeds that germinate better after exposure to high temperatures.[75]