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Principles of terrestrial ecosystem ecology.pdf

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11<br />

Trophic Dynamics<br />

Introduction<br />

Although <strong>terrestrial</strong> animals consume a relatively<br />

small proportion <strong>of</strong> net primary production<br />

(NPP), they strongly affect energy flow<br />

and nutrient cycling. In earlier chapters we<br />

emphasized the interactions between plants<br />

and soil microbes, because these two groups<br />

directly account for about 90% <strong>of</strong> the energy<br />

transfers in most <strong>terrestrial</strong> <strong>ecosystem</strong>s. Plants<br />

use solar energy to reduce CO 2 to organic<br />

matter, most <strong>of</strong> which senesces, dies, and<br />

directly enters the soil, where it is broken down<br />

by bacteria and fungi. Similarly, most nutrient<br />

transfers in <strong>ecosystem</strong>s involve uptake by<br />

plants and return to the soil as dead organic<br />

matter, where nutrients are released and made<br />

available by microbial mineralization. In most<br />

<strong>terrestrial</strong> <strong>ecosystem</strong>s the uncertainties in our<br />

estimates <strong>of</strong> primary production and decomposition<br />

exceed the total energy transfers from<br />

plants to animals. It is perhaps for this reason<br />

that <strong>terrestrial</strong> <strong>ecosystem</strong> ecologists have frequently<br />

ignored animals in classical studies <strong>of</strong><br />

production and biogeochemical cycles. Aquatic<br />

ecologists, however, have been unable to ignore<br />

animals because most <strong>of</strong> the energy and nutrients<br />

are transferred from plants to animals<br />

rather than directly from plants to dead organic<br />

matter (see Chapter 10). Perhaps for this<br />

244<br />

Trophic dynamics govern the movement <strong>of</strong> carbon, nutrients, and energy among<br />

organisms in an <strong>ecosystem</strong>. This chapter describes the controls over trophic dynamics<br />

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

reason aquatic <strong>ecosystem</strong> ecologists have generally<br />

led the theoretical developments in this<br />

aspect <strong>of</strong> <strong>ecosystem</strong> <strong>ecology</strong>.<br />

Understanding the factors governing energy<br />

and nutrient transfer to animals has societal<br />

implications. Most human populations depend<br />

heavily on high-protein foods derived from<br />

animals. The exponentially increasing human<br />

population requires more food in a world<br />

where many people already face an inadequate<br />

food supply. An ecologically viable strategy for<br />

efficiently providing food to feed the growing<br />

human population requires a good understanding<br />

<strong>of</strong> the ecological principles regulating<br />

the efficiency with which plants and animals<br />

support their growth and maintenance.<br />

Overview<br />

Energy transfers define the trophic structure <strong>of</strong><br />

<strong>ecosystem</strong>s. The simplest way to visualize the<br />

energetic interactions among organisms in an<br />

<strong>ecosystem</strong> is to trace the fate <strong>of</strong> a packet <strong>of</strong><br />

energy from the time it enters the <strong>ecosystem</strong><br />

until it leaves—without worrying about the<br />

identity <strong>of</strong> the organisms involved (Lindeman<br />

1942). Trophic transfers involve the feeding <strong>of</strong><br />

one organism on another or on dead organic<br />

matter. Plants are called primary producers or

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