Ecology. Michael Begon

and CAM physiologies. The water‐use efficiency of C₄ plants (the amount of carbon fixed per unit of water transpired) may be double that of C₃ plants.

      tactical changes in stomatal conductance

      The major tactical control of the rates of both photosynthesis and water loss is through changes in stomatal ‘conductance’. These may occur rapidly during the course of a day and allow a very rapid response to immediate water shortages, such that rhythms of stomatal opening may ensure that the above‐ground parts of the plant remain more or less watertight except during controlled periods of active photosynthesis. Stomatal movement may even be triggered directly by conditions at the leaf surface itself – the plant then responds to desiccating conditions at the very site, and at the same time, as the conditions are first sensed.

      coexisting alternative strategies in Australian savannas

The viability of alternative strategies to solve a common problem is nicely illustrated by the trees of seasonally dry tropical forests and woodlands (Eamus, 1999). Communities of this type are found naturally in Africa, the Americas, Australia and India, and as a result of human interference elsewhere in Asia. But whereas, for example, the savannas of Africa and India are dominated by deciduous species, and the Llanos of South America are dominated by evergreens, the savannas of Australia are occupied by roughly equal numbers of species from four groups (Figure 3.11a): evergreens (a full canopy all year), deciduous species (losing all leaves for at least one and usually two to four months each year), semideciduous species (losing around 50% or more of their leaves each year) and brevideciduous species (losing only about 20% of their leaves). At the ends of this continuum, the deciduous species avoid drought in the dry season (April–November in Australia) as a result of their vastly reduced rates of transpiration (Figure 3.11b), but make no net photosynthate at all for around three months, whereas the evergreens maintain a positive carbon balance throughout the year (Figure 3.11c). The alternative, contrasting strategies are clearly sufficiently viable for them to coexist in Australia. Why this is not equally true elsewhere is not known.

Figure 3.11 Alternative strategies for combining photosynthesis and water conservation among trees in Australian savannas. (a) Percentage canopy fullness for deciduous (red), semideciduous (yellow), brevideciduous (purple) and evergreen (blue) trees in Australian savannas throughout the year. (Note that the southern hemisphere dry season runs from around April to November.) (b) Susceptibility to drought as measured by increasingly negative values of ‘predawn water potential’ for deciduous and evergreen trees. (c) Net photosynthesis as measured by the carbon assimilation rate for deciduous and evergreen trees.

      Source: After Eamus (1999).

      3.3.2 Roots as water foragers

Counteracting loss is, of course, only one side of the balance sheet. For most terrestrial plants, the main source of water is the soil and they gain access to it through a root system. We proceed here (and in Section 3.5, on plant nutrient resources) on the basis of plants simply having ‘roots’. In fact, most plants do not have simple, plant‐only roots – they have mycorrhizae: associations of fungal and root tissue in which both partners are crucial to the resource‐gathering properties of the whole. Mycorrhizae, and the respective roles of the plants and the fungi, are discussed in Chapter 13.

      field capacity and the permanent wilting point

Water enters the soil as rain or melting snow and forms a reservoir in the pores between soil particles. What happens to it then depends on the size of the pores, which may hold it by capillary forces against gravity (Figure 3.12). If the pores are wide, as in a sandy soil, much of the water will drain away until it reaches some impediment and accumulates as a rising water table or finds its way into streams or rivers. The water held by soil pores against the force of gravity is called the field capacity of the soil. This is the upper limit of the water that a freely drained soil will retain. However, not all the water retained by soil is available to plants, since they must extract it from those soil pores against the surface tension holding it there, and their ability to do so depends on the structure of their root systems. Hence, there is also a lower limit to the water that can be used in plant growth, determined by the particular plant species present, known as the permanent wilting point – the soil water content at which plants wilt and are unable to recover. The permanent wilting point does not differ much between the plant species of mesic environments (those with a moderate amount of water) or between species of crop plants, but many species native to arid regions have very low permanent wilting points as a result of root systems that allow them to extract significantly more water from the soil, and of leaf morphological adaptations, discussed previously, that give them better water‐holding capacities.

Figure 3.12 Field capacity and the permanent wilting point in soil in relation to pore size and pressure. The status of water in the soil, showing the relationship between the diameter of soil pores that remain water‐filled and the pressure created by the capillary action of those pores that opposes the tendency of water to drain away under the force of gravity. Pressure values are negative because they describe the process of suction. The size of water‐filled pores may be compared in the figure with the sizes of rootlets, root hairs and bacterial cells. Note that for most species of crop plant the permanent wilting point is at approximately −15 bars, but in many other species it reaches −80 bars, depending on their ability to extract water from the narrowest pores.

      roots and the dynamics of water depletion zones

      As a root withdraws water from the soil pores at the root’s surface, it creates water‐depletion zones around it – another example of the RDZs described in Section 3.2.1. These determine gradients of water potential between the interconnected soil pores. Water flows along the gradient into the depleted zones, supplying further water to the root, but this simple process is made much more complex because the more the soil around the roots is depleted of water, the more resistance there is to water flow. Thus, as the root starts to withdraw water from the soil, the first water that it obtains is from the wider pores because they hold the water with weaker capillary forces. This leaves only the narrower, more tortuous pathways, and so the resistance to water flow increases. Thus, when the root draws water from the soil very rapidly, the RDZ may become very sharply defined, because water can move across its boundary only slowly. For this reason, rapidly transpiring plants