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Principles of terrestrial ecosystem ecology.pdf

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6<br />

Terrestrial Production Processes<br />

The balance between gross primary production and <strong>ecosystem</strong> respiration determines<br />

the net carbon accumulation by the biosphere. This chapter describes the<br />

factors that regulate the carbon balance <strong>of</strong> <strong>terrestrial</strong> vegetation and <strong>ecosystem</strong>s.<br />

Introduction<br />

The carbon balance <strong>of</strong> vegetation and <strong>ecosystem</strong>s<br />

governs the productivity <strong>of</strong> the biosphere<br />

and the impact <strong>of</strong> <strong>ecosystem</strong>s on the Earth<br />

System. Plant production determines the<br />

amount <strong>of</strong> energy available to sustain all organisms,<br />

including humans. We depend on plant<br />

production directly for food and fiber and indirectly<br />

because <strong>of</strong> the critical role <strong>of</strong> vegetation<br />

in all <strong>ecosystem</strong> processes (see Chapter 16).<br />

Much <strong>of</strong> the carbon produced by plants eventually<br />

moves to the soil, where it influences the<br />

capacity <strong>of</strong> soils to retain water and nutrients<br />

and therefore to support plant production (see<br />

Chapter 3). Carbon cycling through <strong>ecosystem</strong>s<br />

also directly affects Earth’s climate by modifying<br />

the concentration <strong>of</strong> atmospheric CO2 (see<br />

Chapter 2). Because <strong>of</strong> the many critical roles<br />

<strong>of</strong> carbon balance in the biosphere and the<br />

Earth System, the recent rapid change in<br />

carbon cycling <strong>of</strong> plants and <strong>ecosystem</strong>s is an<br />

issue <strong>of</strong> fundamental societal importance.<br />

Overview<br />

Carbon that enters <strong>ecosystem</strong>s as gross primary<br />

production (GPP) accumulates within the <strong>ecosystem</strong>,<br />

returns to the atmosphere via respira-<br />

tion or disturbance, or is transported laterally<br />

to other <strong>ecosystem</strong>s. About half <strong>of</strong> GPP is<br />

respired by plants to provide the energy that<br />

supports their growth and maintenance<br />

(Schlesinger 1997, Waring and Running 1998).<br />

Net primary production (NPP) is the net<br />

carbon gain by vegetation and equals the difference<br />

between GPP and plant respiration.<br />

Plants lose carbon through several pathways<br />

besides respiration (Fig. 6.1). The largest <strong>of</strong><br />

these releases is typically the transfer <strong>of</strong> carbon<br />

from plants to the soil. This occurs through litterfall<br />

(the shedding <strong>of</strong> plant parts and death <strong>of</strong><br />

plants), root exudation (the secretion <strong>of</strong> soluble<br />

organic compounds by roots into the soil), and<br />

carbon transfers to microbes that are symbiotically<br />

associated with roots (e.g., mycorrhizae<br />

and nitrogen-fixing bacteria). These carbon<br />

transfers from plants to soil eventually give rise<br />

to soil organic matter (SOM), which is typically<br />

the largest pool <strong>of</strong> <strong>ecosystem</strong> carbon. Herbivores<br />

also remove carbon from plants. Herbivory<br />

<strong>of</strong>ten accounts for 5 to 10% <strong>of</strong> NPP<br />

in <strong>terrestrial</strong> <strong>ecosystem</strong>s but can be less than<br />

1% in some forests or greater than 50% in<br />

some grasslands (see Chapter 11). Herbivores<br />

account for most <strong>of</strong> the carbon loss from plants<br />

in aquatic <strong>ecosystem</strong>s (see Chapter 10). Plants<br />

also release carbon to the atmosphere through<br />

emission <strong>of</strong> volatile organic compounds or by<br />

123

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