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Principles of terrestrial ecosystem ecology.pdf

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98 5. Carbon Input to Terrestrial Ecosystems<br />

LONG-TERM<br />

CONTROLS<br />

STATE<br />

FACTORS<br />

BIOTA<br />

TIME<br />

PARENT<br />

MATERIAL<br />

CLIMATE<br />

Interactive<br />

controls<br />

Plant<br />

functional<br />

types<br />

Soil<br />

resources<br />

Figure 5.1. The major factors governing gross<br />

primary production (GPP) in <strong>ecosystem</strong>s. These<br />

controls range from the direct controls, which determine<br />

the diurnal and seasonal variations in GPP,<br />

to the interactive controls and state factors, which<br />

are the ultimate causes <strong>of</strong> <strong>ecosystem</strong> differences<br />

in GPP. Thickness <strong>of</strong> the arrows associated with<br />

In this chapter we explore the mechanisms<br />

behind these causal relationships.<br />

Carbon and energy are linked as they move<br />

through <strong>ecosystem</strong>s because the same processes<br />

govern their entry into <strong>ecosystem</strong>s in<br />

photosynthesis, transfer within <strong>ecosystem</strong>s, and<br />

loss from <strong>ecosystem</strong>s. Photosynthesis uses light<br />

energy (i.e., radiation in the visible portion <strong>of</strong><br />

the spectrum) to reduce CO 2 and produce<br />

carbon-containing organic compounds. This<br />

organic carbon and its associated energy are<br />

then transferred among components within the<br />

<strong>ecosystem</strong> and are eventually released to the<br />

atmosphere by respiration or combustion.<br />

The energy content <strong>of</strong> organic matter differs<br />

among carbon compounds, but for whole tissues,<br />

it is relatively constant at about 20kJg -1<br />

<strong>of</strong> ash-free dry mass (Golley 1961, Larcher<br />

1995). The carbon concentration <strong>of</strong> organic<br />

matter is also variable but averages about 45%<br />

in herbaceous tissues and 50% in wood (Gower<br />

et al. 1999). Both the carbon and energy con-<br />

SHORT-TERM<br />

CONTROLS<br />

Direct<br />

controls<br />

Leaf area<br />

Nitrogen<br />

Season<br />

length<br />

Temperature<br />

Light<br />

CO 2<br />

tents <strong>of</strong> organic matter are greatest in materials<br />

such as seeds and animal fat that have high<br />

lipid content and are lowest in tissues with high<br />

concentrations <strong>of</strong> minerals or organic acids<br />

(Fig. 5.2). Because <strong>of</strong> the relative constancy <strong>of</strong><br />

the carbon and energy contents <strong>of</strong> organic<br />

matter, carbon, energy, and biomass have<br />

been used interchangeably as currencies <strong>of</strong> the<br />

carbon and energy dynamics <strong>of</strong> <strong>ecosystem</strong>s.The<br />

preferred units differ among fields <strong>of</strong> <strong>ecology</strong>,<br />

depending on the processes that are <strong>of</strong> greatest<br />

interest or are measured most directly. Production<br />

studies, for example, typically focus on<br />

biomass; trophic studies, on energy; and gas<br />

exchange studies, on carbon.<br />

Photosynthetic Pathways<br />

C 3 Photosynthesis<br />

GPP<br />

direct controls indicates the strength <strong>of</strong> the effect.<br />

The factors that account for most <strong>of</strong> the variation in<br />

GPP among <strong>ecosystem</strong>s are leaf area and length <strong>of</strong><br />

the photosynthetic season, which are ultimately<br />

determined by the interacting effects <strong>of</strong> soil<br />

resources, climate, vegetation, and disturbance<br />

regime.<br />

The rate <strong>of</strong> carbon input to <strong>ecosystem</strong>s depends<br />

on the response <strong>of</strong> photosynthetic biochemistry

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