Principles of terrestrial ecosystem ecology.pdf
Principles of terrestrial ecosystem ecology.pdf
Principles of terrestrial ecosystem ecology.pdf
Create successful ePaper yourself
Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.
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.)