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Principles of terrestrial ecosystem ecology.pdf

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174 7. Terrestrial Decomposition<br />

Methane emission from soils to the atmosphere<br />

is <strong>of</strong> global concern. Methane is 20-fold<br />

more effective in absorbing infrared radiation<br />

than is CO2. Moreover, its concentration in the<br />

atmosphere has risen dramatically in recent<br />

decades, in part as a result <strong>of</strong> increased area <strong>of</strong><br />

rice paddies and reservoirs (see Fig. 15.3). Even<br />

in wetlands, methane accounts for only 5 to<br />

15% <strong>of</strong> the carbon released to the atmosphere<br />

by decomposers. Methane is thus quantitatively<br />

more important in its role as a greenhouse gas<br />

than as a path <strong>of</strong> carbon loss from <strong>ecosystem</strong>s<br />

(see Fig. 6.8).<br />

Methane is even more highly reduced than<br />

are carbohydrates, so it is an effective energy<br />

source for organisms that have access to<br />

oxygen. Another group <strong>of</strong> bacteria (methanotrophs)<br />

that occur in the surface soils <strong>of</strong> wetlands<br />

use this methane as an energy source and<br />

consume much <strong>of</strong> the methane before it diffuses<br />

to the atmosphere. The enzyme system<br />

that converts ammonium to nitrate also reacts<br />

with methane, causing well-aerated soils to be<br />

a net sink for methane. There are therefore<br />

important transfers between methane producers<br />

and consumers that occur both vertically<br />

within poorly drained <strong>ecosystem</strong>s and horizontally<br />

from lowland to upland <strong>ecosystem</strong>s.<br />

Summary<br />

Decomposition is the conversion <strong>of</strong> dead<br />

organic matter into CO2 and inorganic nutrients<br />

through the action <strong>of</strong> leaching, fragmentation,<br />

and chemical alteration. Leaching<br />

removes soluble materials from decomposing<br />

organic matter. Fragmentation by soil animals<br />

breaks large pieces <strong>of</strong> organic matter into<br />

smaller ones that provide a food source for soil<br />

animals and create fresh surfaces for microbial<br />

colonization. Fragmentation also mixes the<br />

decomposing organic matter into the soil.<br />

Chemical alteration <strong>of</strong> dead organic matter is<br />

primarily a consequence <strong>of</strong> the activity <strong>of</strong> bacteria<br />

and fungi, although some chemical reactions<br />

occur spontaneously in the soil without<br />

microbial mediation.<br />

Decomposition rate is regulated by physical<br />

environment, substrate quality, and the compo-<br />

sition <strong>of</strong> the microbial community (including<br />

soil animals). Carbon chemistry is a strong<br />

determinant <strong>of</strong> litter quality; labile substrates,<br />

such as sugars and proteins, decompose<br />

more rapidly than recalcitrant ones, such as<br />

lignin and microbial cell walls. Nitrogen and<br />

phosphorus supply can also constrain the<br />

decomposition <strong>of</strong> labile carbon substrates, such<br />

as agricultural residues and root exudates.<br />

Plants in high-resource environments produce<br />

litter with high litter quality and therefore rapid<br />

decomposition rates. Decomposition rate<br />

declines with time, as recalcitrant substrates are<br />

depleted. Soil animals strongly influence<br />

decomposition by fragmenting litter, consuming<br />

soil microbes, and mixing the litter into<br />

mineral soil. The environmental factors that<br />

favor NPP (warm, moist, fertile soils) also<br />

promote decomposition so there is no clear<br />

relationship between the amount <strong>of</strong> carbon that<br />

accumulates in soils with either NPP or decomposition<br />

rate.<br />

Review Questions<br />

1. What is decomposition, and why is it important<br />

to the functioning <strong>of</strong> <strong>ecosystem</strong>s?<br />

2. What are the three major processes that<br />

give rise to decomposition? What are the<br />

major controls over each <strong>of</strong> these<br />

processes? Which <strong>of</strong> these processes is<br />

directly responsible for most <strong>of</strong> the mass<br />

loss from decomposing litter?<br />

3. What are the major similarities and differences<br />

between bacteria and fungi in the<br />

ways in which they decompose dead<br />

organic matter? How do these two groups<br />

<strong>of</strong> decomposers differ in their response to<br />

moisture and nutrients? Why?<br />

4. What roles do soil animals play in decomposition?<br />

How does this role differ between<br />

protozoans and earthworms?<br />

5. Why do decomposer organisms secrete<br />

enzymes into the soil rather than breaking<br />

down dead organic matter inside their<br />

bodies?<br />

6. What chemical traits determine the quality<br />

<strong>of</strong> soil organic matter? How do carbon<br />

quality and the C:N ratio differ between

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