5.2 Energy Exchange of Organisms

This module develops elementary concepts of thermodynamics as they apply to ecological processes. The First Law of Thermodynamics deals with the conservation of energy and is of fundamental importance. It states that during any biological, physical, or chemical process energy can be neither created nor destroyed. This module and its companion “Application of the First Law to Ecological Systems” serve as a foundation for other modules in the thermodynamics series.

The relationship between an organism and its environment can be studied by examining the exchange of energy in the form of the flow of heat and the transport of mass. Temperature-dependent biological processes, such as the rate of photosynthesis and the production of metabolic heat, are universally recognized as important in ecology and have been the subjects of extensive research. The First Law of Thermodynamics is the basic tool needed to understand how the organism is coupled to the environment and to quantitatively describe the balance of energy inputs, outputs, and production that underlies the temperature dependence of physiological relationships. Thermal biology is broadly applicable but should be particularly relevant for physiologists, behaviorists, and ecologists.

It is well known that organisms have evolved special adaptations to survive in climatically extreme areas such as polar and desert regions. Less emphasis has been placed on the fact that all organisms continually exchange energy with the environment and must be adapted to heat and water vapor fluxes. The climate and particularly the microclimate (climate in the immediate vicinity of the organism) influence these two exchanges.

It is often advantageous for an organism to reduce or increase the flow of heat energy by physiological or behavioral means; otherwise it may not be able to survive. The microclimate and thus heat balance may greatly affect distribution and activity time of plants and animals. The following four examples should clarify these ideas.

Moose are found in temperate forests throughout the northern hemisphere. During the summer this large herbivore selects three kinds of food to meet its nutritional requirements: small herbaceous plants, leaves, and aquatic plants. On Isle Royale in Lake Superior, Michigan, USA, individuals are commonly found eating in ponds or deciduous stands, or resting under conifers. Associated with each of these habitats is a microclimate that depends on the ground temperature, sunshine, wind speed, and humidity. Because of many factors including its large body size and small tolerance for changes on body temperature, the moose can quickly overheat if active during midday. Belovsky (1977) has made calculations indicating that during the summer moose carefully choose feeding times so as to minimize metabolic output with the constraint that the risk of overheating be small.

An example from the plant kingdom is cacti that have evolved special water storage structures to survive in the dry regions of North and South America. Local climate greatly affects the distribution of these plants because low temperatures will freeze the stored water, rupturing the cells and killing the plant. This fact is especially evident if we compare the large Sonoran Desert species with some South American cacti. In the desert around Tucson, Arizona, USA, the Saguaro cactus is limited to extremely low elevations where freezing weather never lasts longer than about 36 hours (MacArthur 1972). Trichocereous terschenki, a species of similar life form, is found at 2,700 meters in the Andes Mountains. Every night the temperature drops below freezing but because of the relatively constant day length the average daily temperature throughout the year is never below freezing. Therefore, even at this high elevation, the tropical latitude ensures that it is never cold enough long enough for the plant cells to freeze and rupture. Both plant species have the same form and thus similar growth and photosynthetic strategies and both can be killed if their cells are ruptured. Different microclimate patterns, however, seem to determine in part the distribution of Saguaro and T. terschenkii cacti.

Dayton (1971) studied an organism that is limited in its distribution by the physical environment. The anemone, Anthopleura elegantissima, is a common member of the rocky intertidal community of the northern Pacific Coast. To investigate the importance of microclimate, Dayton (1971) established aggregates in uncolonized areas during the autumn night tide. Large numbers survived through the winter but turned brown and died from desiccation during March and April when the tides shifted to the daylight hours. Presumably, the anemones received large amounts of radiation from the sun, which evaporated the water from the animals causing desiccation and which simultaneously raised their body temperatures contributing to their death. Thus, anemones that live above mean low water and therefore that are regularly exposed to a terrestrial environment are found underneath or in the cracks and crevices of rocks. These microhabitats receive much less sunshine (direct solar radiation) allowing the animal to survive out of water.

Clark and Lister’s (1975a, b) investigation of the waxy covering of conifer needles provides the final example of the importance of heat balance for biological organisms. Electron microscopy revealed the cuticle of each needle was covered with tubular crystals of wax. Clark and Lister hypothesized that this material was part of the plant’s stress avoidance system. They found (1) that the crystals were very effective in reflecting light in the shorter (blue) wavelengths of the visible spectrum, (2) that in all the species examined the amount of crystals increased with higher elevation (more radiation) and the sunnier habitats and (3) that a theoretical calculation using the diameter of particles shows that a large amount of ultraviolet light would also be reflected (Mies scattering). Two well-known observations agree with Clark and Lister’s evidence: (1) conifers become bluer in color (from increasing wax covering) with elevation (this trend is exemplified by the Blue Spruce Picea pungens Engelm. var. hoopsii) and (2) foresters have found that conifer seedlings derived from low elevation stock will die if transplanted to higher elevations, while plants from higher elevations survive at lower elevations but grow slowly. Clark and Lister have suggested that larger amounts of wax found on the higher elevation plants reduce the uptake of CO2, slowing photosynthesis but allowing the conifers to avoid otherwise intolerable ultraviolet radiation and drought stress.

It should be noted that the patterns we observed in the above examples may not be completely explained by the relation between an organism and the physical environment. Other physical, historical, chemical, and biological factors are also very important. For instance, moose are critically limited by the amount of sodium available, whereas cacti may need specific kinds of soils and pollinators. The physical stress of waves and drift logs prevent anemones from colonizing and in certain climates conifers cannot compete with deciduous trees. The relation between an organism and the physical environment provides a useful baseline or null model for investigating the organism’s ecology.

The above examples have been included to emphasize the importance of the microclimate for both plants and animals. All organisms exchange heat with the environment. In this module we will focus on the First Law of Thermodynamics, the physical principle needed to understand this heat balance. The First Law can also be useful for other kinds of ecological interests such as the energetic coat of locomotion, rate of food assimilation and energy flow in ecosystems. Some discussion of these topics is also included. More biological examples of the physical principles elaborated here can be found in the module titled “Applications of the First Law to Ecological Systems.” The reader may wish to review the principles and definitions of this module, which may be difficult to absorb entirely without the examples of the second module.