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Principles of terrestrial ecosystem ecology.pdf

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256 11. Trophic Dynamics<br />

eating rate may be constrained by processing<br />

time. Animals that are more selective and have<br />

a lower feeding rate generally eat food that is<br />

more digestible, contributing to their shorter<br />

passage time.<br />

Assimilation Efficiency<br />

Assimilation efficiency depends on both the<br />

quality <strong>of</strong> the food and the physiology <strong>of</strong> the<br />

consumer. Assimilation efficiency is the proportion<br />

<strong>of</strong> ingested energy that is digested<br />

and assimilated (An) into the bloodstream<br />

(Fig. 11.7).<br />

E<br />

assim =<br />

A<br />

I<br />

(11.4)<br />

Unassimilated material returns to the soil as<br />

feces, a component <strong>of</strong> the detrital input to<br />

<strong>ecosystem</strong>s.<br />

Assimilation efficiencies are <strong>of</strong>ten higher (5<br />

to 80%) than consumption efficiencies (0.1 to<br />

50%). Carnivores feeding on vertebrates tend<br />

to have higher assimilation efficiencies (about<br />

80%) than do <strong>terrestrial</strong> herbivores (5 to 20%),<br />

because carnivores eat food that has less structural<br />

material than is present in <strong>terrestrial</strong><br />

plants. Carnivores that kill large prey can avoid<br />

eating indigestible parts such as bones, whereas<br />

most <strong>terrestrial</strong> herbivores consume the indigestible<br />

cell wall structure in combination with<br />

cell contents. Among herbivores, species that<br />

feed on seeds, which have high concentrations<br />

<strong>of</strong> digestible, energy-rich storage reserves, have<br />

a higher assimilation efficiency than those<br />

feeding on leaves. Leaf-feeding herbivores,<br />

in turn, have higher assimilation efficiencies<br />

than those feeding on wood, which has higher<br />

concentrations <strong>of</strong> cellulose and lignin. Many<br />

aquatic herbivores have a particularly high<br />

assimilation efficiency (up to 80%) because <strong>of</strong><br />

the low allocation to structure in many algae<br />

and other aquatic plants. Even in aquatic<br />

<strong>ecosystem</strong>s, however, herbivores that feed on<br />

well-defended species have low assimilation<br />

efficiencies. Assimilation efficiencies <strong>of</strong> herbivores<br />

feeding on cyanobacteria, for example,<br />

can be as low as 20%.<br />

The physiological properties <strong>of</strong> a consumer<br />

strongly influence assimilation efficiency. Rumi-<br />

n<br />

n<br />

nants, which carry a vat <strong>of</strong> cellulose-digesting<br />

microbes (the rumen) have a higher assimilation<br />

efficiency (about 50%) than do most<br />

nonruminant herbivores. One reason for the<br />

high assimilation efficiency <strong>of</strong> ruminants is the<br />

greater processing time than in nonruminants<br />

<strong>of</strong> similar size, giving more time for microbial<br />

breakdown <strong>of</strong> food. Homeotherms typically<br />

have higher assimilation efficiencies than do<br />

poikilotherms due to the warmer, more constant<br />

gut temperature, which promotes digestion<br />

and assimilation. Homeotherms therefore<br />

have an advantage over poikilotherms in both<br />

consumption and assimilation efficiency.<br />

Production Efficiency<br />

Production efficiency is determined primarily<br />

by animal metabolism. Production efficiency<br />

is the proportion <strong>of</strong> assimilated energy that<br />

is converted to animal production (Fig. 11.7).<br />

Production efficiency includes both growth <strong>of</strong><br />

individuals and reproduction to produce new<br />

individuals.<br />

od<br />

Eprod<br />

=<br />

A<br />

Pr<br />

(11.5)<br />

Assimilated energy that is not incorporated<br />

into production is lost to the environment as<br />

respiratory heat. Production efficiencies for<br />

individual animals vary 50-fold from less than<br />

1 to greater than 50% (Table 11.2) and differ<br />

most dramatically between homeotherms (Eprod<br />

1 to 3%) and poikilotherms (Eprod 10 to 50%).<br />

Homeotherms expend most <strong>of</strong> their assimilated<br />

energy maintaining a relatively constant body<br />

temperature. This high constant body temperature<br />

makes their activity less dependent on<br />

environmental temperature and increases their<br />

capacity to catch prey and avoid predation but<br />

makes homeotherms inefficient in producing<br />

new animal biomass. Among homeotherms,<br />

production efficiency decreases with decreasing<br />

body size because a small size results in a high<br />

surface to volume ratio and therefore a high<br />

rate <strong>of</strong> heat loss from the warm animal to the<br />

cold environment. In contrast, the production<br />

efficiency <strong>of</strong> poikilotherms is relatively high<br />

(about 25%) and tends to decrease with<br />

increasing body size. Some large-bodied<br />

n<br />

n

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