Principles of terrestrial ecosystem ecology.pdf
Principles of terrestrial ecosystem ecology.pdf
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