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Principles of terrestrial ecosystem ecology.pdf

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absorption, indicating the large energetic cost<br />

<strong>of</strong> nutrient uptake. Elements required in small<br />

quantities are <strong>of</strong>ten absorbed simply by mass<br />

flow or diffusion into the root cortical cells<br />

(Table 8.2).<br />

Some plant species tap pools <strong>of</strong> nutrients that<br />

are unavailable to other plants. Although all<br />

plants require the same suite <strong>of</strong> nutrients in<br />

similar proportions, nitrogen is available in<br />

several forms (nitrate, ammonium, amino acids,<br />

etc.) that differ in availability among <strong>ecosystem</strong>s.<br />

Many species preferentially absorb<br />

ammonium (and perhaps amino acids) over<br />

nitrate, when all nitrogen forms are equally<br />

available (Table 8.3). Species differ, however, in<br />

their relative preference for these nitrogen<br />

forms, frequently showing a high capacity to<br />

absorb the forms that are most abundant in<br />

the <strong>ecosystem</strong>s to which they are adapted.<br />

Many species that occupy highly organic soils<br />

<strong>of</strong> tundra and boreal forest <strong>ecosystem</strong>s, for<br />

example, preferentially absorb amino acids<br />

(Chapin et al. 1993, Näsholm et al. 1998), although<br />

even agricultural species use amino acid<br />

nitrogen (Näsholm et al. 2000). An important<br />

community consequence <strong>of</strong> species differences<br />

in nitrogen preference is that nitrogen represents<br />

several distinct resources for which<br />

species can compete. Species in the same community<br />

<strong>of</strong>ten have quite different isotopic signatures<br />

<strong>of</strong> tissue nitrogen, because they acquire<br />

Nutrient Uptake 185<br />

Table 8.3. Preference ratios for plant absorption <strong>of</strong> different forms <strong>of</strong> nitrogen a when all forms are equally<br />

available.<br />

+ -<br />

NH4 :NO3<br />

+<br />

Glycine : NH4 Species preferenceb preferenceb References<br />

Arctic vascular plants 1.1 2.1 ± 0.6 (12) Chapin et al. (1993), Kielland (1994)<br />

Arctic nonvascular plants 5.0 ± 1.5 (2) Kielland (1997)<br />

Boreal trees 19.3 ± 5.8 (4) 1.3 Chapin et al. (1986b), Kronzucker<br />

et al. (1997), Näsholm et al. (1998)<br />

Alpine sedges 3.9 ± 1.3 (12) 1.5 ± 0.4 (11) Raab et al. (1999)<br />

Temperate heath 1.0 Read and Bajwa (1985)<br />

Salt marsh 1.3 Morris (1980)<br />

Mediterranean shrub 1.2 Stock and Lewis (1984)<br />

Barley 1.0 Chapin et al. (1993)<br />

Tomato 0.6 Smart and Bloom (1988)<br />

a Assumes all forms <strong>of</strong> nitrogen are equally available.<br />

b A preference ratio >1 indicates that the first form <strong>of</strong> nitrogen is absorbed preferentially over the second. Number <strong>of</strong><br />

species studied in parentheses. Research shows that many plants preferentially absorb glycine (a highly mobile amino<br />

acid) over ammonium and preferentially absorb ammonium over nitrate, when all forms are equally available.<br />

nitrogen from different sources—either different<br />

chemical fractions (nitrate, ammonium,<br />

organic nitrogen) or different soil depths<br />

(Nadelh<strong>of</strong>fer et al. 1996) (Fig. 8.4). Changes in<br />

species composition could therefore alter the<br />

nitrogen pools that are used to support primary<br />

production. In most cases, the species present<br />

are capable <strong>of</strong> using the prevailing forms <strong>of</strong><br />

available nitrogen. If human activities alter<br />

the prevailing form <strong>of</strong> available nitrogen, for<br />

example through nitrate deposition in coniferous<br />

forests, this novel form <strong>of</strong> nitrogen may be<br />

used less effectively by the extant vegetation.<br />

Plant functional type<br />

Woody evergreen<br />

Graminoids,<br />

cryptogams<br />

Woody deciduous<br />

Aquatic<br />

Tissue δ 15 -10 -8 -6 -4 -2 0 2 4<br />

N<br />

Figure 8.4. Concentration <strong>of</strong> 15 N in tissues from different<br />

growth forms <strong>of</strong> boreal plants. (Figure provided<br />

by K. Kielland; Kielland 1999.)

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