06.12.2012 Views

Principles of terrestrial ecosystem ecology.pdf

Principles of terrestrial ecosystem ecology.pdf

Principles of terrestrial ecosystem ecology.pdf

SHOW MORE
SHOW LESS

You also want an ePaper? Increase the reach of your titles

YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.

212 9. Terrestrial Nutrient Cycling<br />

chemodenitrification converts NO2 - (nitrite)<br />

abiotically to nitric oxide gas (NO) where NO2 -<br />

accumulates in the soil at low pH. Chemodenitrification<br />

is typically much less important than<br />

biological denitrification.<br />

The sequence <strong>of</strong> NO3 - reduction is NO3 - Æ<br />

NO2 - Æ NO Æ N2O Æ N2. The last three products,<br />

particularly N2O and N2, are released as<br />

gases to the atmosphere (Fig. 9.4). Most denitrifiers<br />

have the enzymatic potential to carry<br />

out the entire reductive sequence but produce<br />

variable proportions <strong>of</strong> N2O and N2, depending<br />

in part on the relative availability <strong>of</strong> oxidant<br />

(NO3 - ) versus reductant (organic carbon).<br />

When NO3 - is relatively more abundant than<br />

labile organic carbon, more N2O than N2 is produced.<br />

Other factors that favor N2O over N2<br />

production include low pH, low temperature,<br />

and high oxygen. Although NO is <strong>of</strong>ten<br />

released during denitrification in laboratory<br />

incubations, this seldom occurs in nature<br />

because its diffusion to the air is impeded<br />

by water-filled pore spaces. Some <strong>of</strong> the NO<br />

that is produced serves as a substrate for<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 />

Indirect<br />

controls<br />

Plant NO 3 -<br />

uptake<br />

Litter quantity<br />

Carbon quality<br />

Root / microbial<br />

respiration<br />

Temperature<br />

H 2 O<br />

Soil texture<br />

Figure 9.7. The major factors controlling denitrification<br />

in soils. These controls range from concentrations<br />

<strong>of</strong> substrates that directly control nitrification<br />

to the interactive controls such as climate and disturbance<br />

regime that are the ultimate determinants<br />

further reduction to N2O or N2 by denitrifying<br />

bacteria.<br />

The three conditions required for significant<br />

denitrification are low oxygen, high nitrate concentration,<br />

and a supply <strong>of</strong> organic carbon (Fig.<br />

9.7) (Del Grosso et al. 2000). In most nonflooded<br />

soils, oxygen availability exerts the<br />

strongest control over denitrification. Oxygen<br />

supply is reduced by high soil water content,<br />

which impedes the diffusion <strong>of</strong> oxygen through<br />

soil pores. Soil moisture, in turn, is controlled<br />

by other environmental factors such as slope<br />

position, soil texture, and the balance between<br />

precipitation and evapotranspiration. Soil<br />

oxygen concentration is also sensitive to its<br />

rate <strong>of</strong> consumption by soil microbes and roots.<br />

It is consumed most quickly in warm, moist<br />

environments.<br />

The second major control over denitrification<br />

is an adequate supply <strong>of</strong> the substrate<br />

NO3 - . Because nitrification is a primarily an<br />

aerobic process, the low-oxygen conditions that<br />

are optimal for denitrification frequently limit<br />

NO3 - supply. Some wetlands, for example, have<br />

SHORT-TERM<br />

CONTROLS<br />

(-)<br />

DIRECT<br />

CONTROLS<br />

(-)<br />

Nitrate<br />

concentration<br />

(-)<br />

Labile<br />

carbon<br />

Oxygen<br />

concentration<br />

DENITRIFICATION<br />

<strong>of</strong> denitrification rate. Thickness <strong>of</strong> the arrows<br />

indicates the strength <strong>of</strong> effects. The influence <strong>of</strong><br />

one factor on another is positive unless otherwise<br />

indicated (-).<br />

(-)

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!