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Principles of terrestrial ecosystem ecology.pdf

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alization is the net accumulation <strong>of</strong> inorganic<br />

nitrogen in the soil solution over a given time<br />

interval. Net mineralization occurs when microbial<br />

growth is limited more strongly by carbon<br />

than by nitrogen, whereas net immobilization<br />

occurs when microbial communities are nitrogen<br />

limited.<br />

Net nitrogen mineralization is an excellent<br />

measure <strong>of</strong> the nitrogen supply to plants in<br />

<strong>ecosystem</strong>s with rapid nitrogen turnover, where<br />

there is little competition for nitrogen between<br />

plants and microbes. The annual net mineralization<br />

in the deciduous forests <strong>of</strong> eastern<br />

North America, for example, approximately<br />

equals nitrogen uptake by vegetation<br />

(Nadelh<strong>of</strong>fer et al. 1992). In less fertile <strong>ecosystem</strong>s,<br />

such as arctic tundra, net nitrogen mineralization<br />

rate substantially underestimates the<br />

amount <strong>of</strong> nitrogen that is annually acquired by<br />

plants (Nadelh<strong>of</strong>fer et al. 1992). There are at<br />

least two explanations for this apparent discrepancy.<br />

(1) Plant roots and their mycorrhizal<br />

fungi, which are excluded in net mineralization<br />

assays, may be good competitors with sapro-<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 />

Figure 9.3. The major factors controlling ammonification<br />

(net nitrogen mineralization) in soils. These<br />

controls range from the proximate control over<br />

nitrogen mineralization (the concentration <strong>of</strong> DON,<br />

physical environment, and microbial carbon to<br />

nitrogen ratio) to the state factors and interactive<br />

Plant<br />

uptake <strong>of</strong><br />

DON<br />

Litter<br />

quantity<br />

Carbon quality<br />

Internal Cycling <strong>of</strong> Nitrogen 205<br />

phytic microbes for mineralized nitrogen in the<br />

real world, so this assay may underestimate the<br />

quantity <strong>of</strong> nitrogen that would be mineralized<br />

in the presence <strong>of</strong> roots (Stark 2000). (2) Plants<br />

and their mycorrhizal fungi that absorb amino<br />

acids may also not depend heavily on nitrogen<br />

mineralization to meet their nitrogen requirements<br />

in infertile <strong>ecosystem</strong>s due to absorption<br />

<strong>of</strong> DON, so the low rates <strong>of</strong> net nitrogen mineralization<br />

in these <strong>ecosystem</strong>s may be an<br />

accurate reflection <strong>of</strong> small fluxes that naturally<br />

occur through this pathway.<br />

Nitrogen mineralization rate is controlled by<br />

the availability <strong>of</strong> DON and inorganic nitrogen,<br />

the activity <strong>of</strong> soil microbes, and their relative<br />

demands for carbon and nitrogen. The quantity<br />

and quality <strong>of</strong> organic matter, including its<br />

relative amounts and forms <strong>of</strong> carbon and<br />

nitrogen, that enter the soil are the major<br />

determinants <strong>of</strong> the substrate available for<br />

decomposition (see Chapter 7) and therefore<br />

the substrate available for nitrogen mineralization<br />

(Fig. 9.3). Thus the state factors and interactive<br />

controls that promote productivity and<br />

SHORT-TERM<br />

CONTROLS<br />

(-)<br />

Direct<br />

controls<br />

Dissolved<br />

organic<br />

nitrogen<br />

Microbial<br />

C:N<br />

Temperature<br />

H 2 O<br />

(-)<br />

NET NITROGEN<br />

MINERALIZATION<br />

(AMMONIFICATION)<br />

controls that ultimately determine the differences<br />

among <strong>ecosystem</strong>s in mineralization rates. Thickness<br />

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

influence <strong>of</strong> one factor on another is positive unless<br />

otherwise indicated (-).

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