The study of how trees grow and develop in terms of genetics; biochemistry; cellular, tissue, and organ functions; and interaction with environmental factors. While many physiological processes are similar in trees and other plants, trees possess unique physiologies that help determine their outward appearance. These physiological processes include carbon relations (photosynthesis, carbohydrate allocation), cold and drought resistance, water relations, and mineral nutrition.
Three characteristics of trees that define their physiology are longevity, height, and simultaneous reproductive and vegetative growth. Trees have physiological processes that are more adaptable than those in the more specialized annual and biennial plants. Height allows trees to successfully compete for light, but at the same time this advantage creates transport and support problems. These problems were solved by the evolution of the woody stem which combines structure and function into a very strong transport system. Simultaneous vegetative and reproductive growth in adult trees causes significant competition for carbohydrates and nutrients, resulting in decreased vegetative growth. Trees accommodate both types of growth by having cyclical reproduction: one year many flowers and seeds are produced, followed by a year or more in which few or no flowers are produced.
While biochemical processes of photosynthesis and carbon assimilation and allocation are the same in trees and other plants, the conditions under which these processes occur in trees are more variable and extreme. In evergreen species, photosynthesis can occur year round as long as the air temperature remains above freezing, while some deciduous species can photosynthesize in the bark of twigs and stem during the winter.
Carbon dioxide fixed into sugars moves through the tree in the phloem and xylem to tissues of high metabolism which vary with season and development. At the onset of growth in the spring, sugars are first mobilized from storage sites, primarily in the secondary xylem (wood) and phloem (inner bark) of the woody twigs, branches, stem, and roots. The sugars, stored as starch, are used to build new leaves and twigs, and if present, flowers. Once the new leaves expand, photosynthesis begins and sugars are produced, leading to additional leaf growth. Activation of the vascular cambium occurs at the same time, producing new secondary xylem and phloem. In late spring, the leaves begin photosynthesizing at their maximum rates, creating excess sugars which are translocated down the stem to support further branch, stem, and root growth. From midsummer through fall until leaf abscission (in deciduous trees) or until temperatures drop to freezing (in evergreen trees), sugars replenish the starch used in spring growth. Root growth may be stimulated at this time by sugar availability and warm soil temperatures. Throughout the winter, starch is used for maintenance respiration, but sparingly since low temperatures keep respiration rates low. See Phloem, Photosynthesis, Xylem
In adult trees, reproductive structures (flowers in angiosperms or strobili in gymnosperms) develop along with new leaves and represent large carbohydrate sinks. Sugars are preferentially utilized at the expense of leaf, stem, and root growth. This reduces the leaf area produced, affecting the amount of sugars produced during that year, thereby reducing vegetative growth even further. The reproductive structures are present throughout the growing season until seed dispersal and continually utilize sugars that would normally go to stem and root growth.
The perennial nature of trees requires them to withstand low temperatures during the winter. Trees develop resistance to freezing through a process of physiological changes beginning in late summer. A tree goes through three sequential stages to become fully cold resistant. The process involves reduced cell hydration along with increased membrane permeability. The first stage is initiated by shortening days and results in shoot growth cessation, bud formation, and metabolic changes. Trees in this stage can survive temperatures down to 23°F (-5°C). The second stage requires freezing temperatures which alter cellular molecules. Starch breakdown is stimulated, causing sugar accumulation. Trees can survive temperatures as low as -13°F (-25°C) at this stage. The last stage occurs after exposure to very low temperatures (-22 to -58°F or -30 to -50°C), which increases soluble protein concentrations that bind cellular water, preventing ice crystallization. Trees can survive temperatures below -112°F (-80°C) in this stage. A few days of warmer temperatures, however, causes trees to revert to the second stage.
Unlike annual plants that survive drought as seeds, trees have evolved traits that allow them to avoid desiccation. These traits include using water stored in the stem, stomatal closure, and shedding of leaves to reduce transpirational area. All the leaves can be shed and the tree survives on stored starch. Another trait of some species is to produce a long tap root that reaches the water table, sometimes tens of meters from the soil surface. On a daily basis, trees must supply water to the leaves for normal physiological function. If the water potential of the leaves drops too low, the stomata close, reducing photosynthesis. To maintain high water potential, trees use water stored in their stems during the morning which is recharged during the night. See Plant-water relations
Transport and support
Trees have evolved a means of combining long-distance transport between the roots and foliage with support through the production of secondary xylem (wood) by the vascular cambium. In older trees the stem represents 60–85% of the aboveground biomass. However, 90% of the wood consists of dead cells. These dead cells function in transport and support of the tree. As these cells develop and mature, they lay down thick secondary walls of cellulose and lignin that provide support, and then they die with the cell lumen becoming an empty tube. The interconnecting cells provide an efficient transport system, capable of moving 106 gal (400 liters) of water per day. The living cells in the wood (ray parenchyma) are the site of starch storage in woody stems and roots. See Plant transport of solutes
Nutrient deficiencies are similar in trees and other plants because of the functions of these nutrients in physiological processes. Tree nutrition is unique because trees require lower concentrations, and they are able to recycle nutrients within various tissues. Trees adapt to areas which are low in nutrients by lowering physiological functions and slowing growth rates. In addition, trees allocate more carbohydrates to root production, allowing them to exploit large volumes of soil in search of limiting nutrients. Proliferation of fine roots at the organic matter-mineral soil interface where many nutrients are released from decomposing organic matter allows trees to recapture nutrients lost by leaf fall. See Plant mineral nutrition, Plant physiology, Tree