Introduction

In order to understand the distribution and abundance of a species we need to know its history (Chapter 1), the resources it requires (Chapter 3), the individuals' rates of birth, death and migration (Chapters 4 and 6), their interactions with their own and other species (Chapters 5 and 8-13) and the effects of environmental conditions. This chapter deals with the limits placed on organisms by environmental conditions.

A condition is as an abiotic environmental factor that influences the functioning of living organisms. Examples include temperature, relative humidity, pH, salinity and the concentration of pollutants. A condition may be modified by the presence of other organisms. For example, temperature, humidity and soil pH may be altered under a forest canopy. But unlike resources, conditions are not consumed or used up by organisms.

For some conditions we can recognize an optimum concentration or level at which an organism performs best, with its activity tailing off at both lower and higher levels (Figure 2.1a). But we need to define what we mean by 'performs best'. From an evolutionary point of view, 'optimal' conditions are those under which individuals leave most descendants (are fittest), but these are often impossible to determine in practice because measures of fitness should be made over several generations. Instead, we more often measure the effect of conditions on some key property like the activity of an enzyme, the respiration rate of a tissue, the growth rate of individuals or their rate of reproduction. However, the effect of variation in conditions on these various properties will often not be the same; organisms can usually survive over a wider range of conditions than permit them to grow or reproduce (Figure 2.1a).

The precise shape of a species' response will vary from condition to condition. The generalized form of response, shown in Figure 2.1a, is appropriate for conditions like temperature and pH

conditions may be altered - but not consumed

Figure 2.1 Response curves illustrating the effects of a range of environmental conditions on individual survival (S), growth (G) and reproduction (R). (a) Extreme conditions are lethal; less extreme conditions prevent growth; only optimal conditions allow reproduction. (b) The condition is lethal only at high intensities; the reproduction-growth-survival sequence still applies. (c) Similar to (b), but the condition is required by organisms, as a resource, at low concentrations.

Figure 2.1 Response curves illustrating the effects of a range of environmental conditions on individual survival (S), growth (G) and reproduction (R). (a) Extreme conditions are lethal; less extreme conditions prevent growth; only optimal conditions allow reproduction. (b) The condition is lethal only at high intensities; the reproduction-growth-survival sequence still applies. (c) Similar to (b), but the condition is required by organisms, as a resource, at low concentrations.

in which there is a continuum from an adverse or lethal level (e.g. freezing or very acid conditions), through favorable levels of the condition to a further adverse or lethal level (heat damage or very alkaline conditions). There are, though, many environmental conditions for which Figure 2.1b is a more appropriate response curve: for instance, most toxins, radioactive emissions and chemical pollutants, where a low-level intensity or concentration of the condition has no detectable effect, but an increase begins to cause damage and a further increase may be lethal. There is also a different form of response to conditions that are toxic at high levels but essential for growth at low levels (Figure 2.1c). This is the case for sodium chloride - an essential resource for animals but lethal at high concentrations - and for the many elements that are essential micronutrients in the growth of plants and animals (e.g. copper, zinc and manganese), but that can become lethal at the higher concentrations sometimes caused by industrial pollution.

In this chapter, we consider responses to temperature in much more detail than other conditions, because it is the single most important condition that affects the lives of organisms, and many of the generalizations that we make have widespread relevance. We move on to consider a range of other conditions, before returning, full circle, to temperature because of the effects of other conditions, notably pollutants, on global warming. We begin, though, by explaining the framework within which each of these conditions should be understood here: the ecological niche.

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