Solar Radiation and Productivity in Tropical Ecosystems*

In thermodynamic terms, ecosystems are machines supplied with energy from an external source, usually the sun. When the input of energy to an ecosystem is exactly equal to its total output of energy, the state of equilibrium which exists is a special case of the First Law of Thermodynamics. The Second Law is relevant oo. It implies that in every spontaneous process, physical or chemical, the production of 'useful' energy, which could be harnessed in a form such as mechanical work, must be accompanied by a simultaneous 'waste' of heat. No biological system can break or evade this law. The heat produced by a respiring cell is an inescapable component of cellular metabolism, the cost which Nature has to pay for creating biological order out of physical chaos in the environment of plants and animals. Dividing the useful energy of a thermodynamic process by the total energy involved gives a figure for the efficiency of the process, and this procedure has been widely used to analyse the flow of energy in ecosystems. For example, the efficiency with which a stand of plants produces dry matter by photosynthesis can be defined as the ratio of chemical energy stored in the assimilates to radiant energy absorbed by foliage during the period of assimilation. The choice of absorbed energy as a base for calculating efficiency is convenient but arbitrary. To derive an efficiency depending on the environment of a particular site as well as oil the nature of the vegetation, dry matter production can be related to the receipt of solar energy at the top of the earth's atmosphere. This exercise was attempted by Professor William Thomson, later Lord Kelvin, in 1852. 'The author estimates the mechanical value of the solar heat which, were none of it absorbed by the atmosphere, would fall annually on each square foot of land, at 530 000 000 foot pounds; and infers that probably a good deal more, 1/1000 of the solar heat, which actually falls on growing plants, is converted into mechanical effect.' Outside the earth's atmosphere, a surface kept at right angles to the sun's rays receives energy at a mean rate of 1360 W m-2 or 1f36 kJ m-2 s-1, a figure known as the solar constant. As the energy stored by plants is about 17 kJ per gram of dry matter, the solar constant is equivalent to the production of dry matter at a rate of about 1 g m-2 every 12 s, 7 kg m-2 per day, or 2 6 t m-2 year-'. The annual yield of agricultural crops ranges from a maximum of 30-60 t ha-' in field experiments to less than I t ha-' in some forms of subsistence farming. When these rates are expressed as a fraction of the integrated solar constant, the efficiencies of agricultural systems lie between 0-2 and 0 004%, a range including Kelvin's estimate of 0-1%. Conventional estimates of efficiency interms of the amount of solar radiation incident at the earth's surface provide ecologists and agronomists with a method for comparing plant productivity under different systems of land use and management and in different * Opening paper read at IBP/UNESCO Meeting on Productivity ofTropical Ecosystems, Makerere University, Uganda, September 1970.