Potential nitrogen constraints on soil carbon sequestration under low and elevated atmospheric CO2.

The interaction between nitrogen cycling and carbon sequestration is critical in predicting the consequences of anthropogenic increases in atmospheric CO2 (hereafter, Ca). The progressive N limitation (PNL) theory predicts that carbon sequestration in plants and soils with rising Ca may be constrained by the availability of nitrogen in many ecosystems. Here we report on the interaction between C and N dynamics during a four-year field experiment in which an intact C3/C4 grassland was exposed to a gradient in Ca from 200 to 560 micromol/mol. There were strong species effects on decomposition dynamics, with C loss positively correlated and N mineralization negatively correlated with Ca for litter of the C3 forb Solanum dimidiatum, whereas decomposition of litter from the C4 grass Bothriochloa ischaemum was unresponsive to Ca. Both soil microbial biomass and soil respiration rates exhibited a nonlinear response to Ca, reaching a maximum at approximately 440 micromol/mol Ca. We found a general movement of N out of soil organic matter and into aboveground plant biomass with increased Ca. Within soils we found evidence of C loss from recalcitrant soil C fractions with narrow C:N ratios to more labile soil fractions with broader C:N ratios, potentially due to decreases in N availability. The observed reallocation of N from soil to plants over the last three years of the experiment supports the PNL theory that reductions in N availability with rising Ca could initially be overcome by a transfer of N from low C:N ratio fractions to those with higher C:N ratios. Although the transfer of N allowed plant production to increase with increasing Ca, there was no net soil C sequestration at elevated Ca, presumably because relatively stable C is being decomposed to meet microbial and plant N requirements. Ultimately, if the C gained by increased plant production is rapidly lost through decomposition, the shift in N from older soil organic matter to rapidly decomposing plant tissue may limit net C sequestration with increased plant production.

Increasing CO2 from subambient to superambient concentrations alters species composition and increases above-ground biomass in a C3 /C4 grassland.

• The glacial-to-present increase in atmospheric CO2 concentration is likely to have stimulated plant production, but experimental tests in natural ecosystems are lacking. • We measured above-ground biomass production, plant nitrogen (N) accumulation, and species dynamics in a C3 /C4 grassland exposed for 4 yr (1997-2000) to a continuous gradient in CO2 from 200-560 mol mol-1 . • Biomass increased with CO2 concentration in 1997-99. Biomass increases ranged between 121 and 161 g m-2 per 100 mol mol-1 rise in CO2 and were similar at subambient and superambient concentrations. Biomass responses to CO2 were determined by different species or functional groups of species during different years. Increasing CO2 accelerated a successional shift initiated by release from grazing in which C3 forbs increased at the expense of a C4 grass. Effects of CO2 on tissue N concentration varied among species and functional groups, but CO2 did not alter total N in above-ground tissues. • Results imply that rising CO2 has stimulated plant production and accelerated successional change and that grasslands will remain sensitive to rising CO2 for several decades.

Nonlinear grassland responses to past and future atmospheric CO(2).

Carbon sequestration in soil organic matter may moderate increases in atmospheric CO(2) concentrations (C(a)) as C(a) increases to more than 500 micromol mol(-1) this century from interglacial levels of less than 200 micromol mol(-1) (refs 1 6). However, such carbon storage depends on feedbacks between plant responses to C(a) and nutrient availability. Here we present evidence that soil carbon storage and nitrogen cycling in a grassland ecosystem are much more responsive to increases in past C(a) than to those forecast for the coming century. Along a continuous gradient of 200 to 550 micromol mol(-1) (refs 9, 10), increased C(a) promoted higher photosynthetic rates and altered plant tissue chemistry. Soil carbon was lost at subambient C(a), but was unchanged at elevated C(a) where losses of old soil carbon offset increases in new carbon. Along the experimental gradient in C(a) there was a nonlinear, threefold decrease in nitrogen availability. The differences in sensitivity of carbon storage to historical and future C(a) and increased nutrient limitation suggest that the passive sequestration of carbon in soils may have been important historically, but the ability of soils to continue as sinks is limited.

Growth rate and survivorship of drought: CO2 effects on the presumed tradeoff in seedlings of five woody legumes.

Traits that promote rapid growth and seedling recruitment when water is plentiful may become a liability when seedlings encounter drought. We tested the hypothesis that CO2 enrichment reinforces any tradeoff between growth rate and drought tolerance by exaggerating interspecific differences in maximum relative growth rate (RGR) and survivorship of drought among seedlings of five woody legumes. We studied invasive species of grasslands that differ in distribution along a rainfall gradient. Survivorship of drought at ambient CO2 concentration ([CO2]) was negatively related to RGR in well-watered seedlings in one of two experiments, but the relationship was weak because interspecific differences in RGR were small. Contrary to our hypothesis, there was no significant relationship among well-watered seedlings between RGR at ambient [CO2] and either the relative or absolute increase in RGR at elevated [CO2]. As predicted, however, CO2 enrichment reinforced interspecific differences in survivorship of seedlings exposed to similar rates of soil water depletion. Doubling [CO2] improved seedling survivorship of the most drought-tolerant species throughout the period of soil water depletion, but did not consistently affect survivorship of more drought-sensitive species. Midday xylem pressure potentials of drought-treated seedlings were less negative at elevated [CO2] than at ambient [CO2], but no other measured trait was consistently correlated with improved survivorship at high [CO2]. Carbon dioxide enrichment may not reinforce species differences in RGR, but could exaggerate interspecific differences in drought tolerance. To the extent that seedling persistence in grasslands correlates with drought survivorship, our results indicate a positive effect of CO2 enrichment on recruitment of woody legumes that are currently tolerant of drought.

Genetic variation for carbon isotope composition in honey mesquite (Prosopis glandulosa).

Carbon isotope composition (delta(13)C) is a useful surrogate for integrated, plant water-use efficiency (WUE) when measured on plants grown in a common environment. In a variety of species, genetic variation in delta(13)C has been linked to the distribution of genotypes across gradients in atmospheric and soil water. We examined genetic variation for delta(13)C in seedlings of honey mesquite (Prosopis glandulosa Torr.), an invasive grassland shrub that thrives in the southwestern USA. Fifteen maternal families, representing progeny of 15 adult trees, were studied in three common garden experiments in a greenhouse. The 990-km east-west transect along which the adult trees were located encompasses a wide precipitation gradient, and includes mesic grassland, semiarid grassland, and Chihuahuan desert ecosystems. Genetic variation for delta(13)C in mesquite was substantial, with the rank order of half-sib families based on delta(13)C relatively stable across experiments, which were conducted under different environmental conditions. Conversely, rankings of families by mean seedling height (an index of growth rate) varied markedly among experiments. Seedlings derived from Chihuahuan desert adults emerged more quickly and had more negative delta(13)C (indicative of lower WUE) than seedlings derived from the other regions. Although delta(13)C and seedling height were not correlated, these results suggest that mesquite genotypes at the drier, western extreme of the species' range are adapted for quicker emergence and possibly faster growth than genotypes from mesic areas. Together, these traits may facilitate exploitation of infrequent precipitation events.

Growth, water relations, and survival of drought-exposed seedlings from six maternal families of honey mesquite (Prosopis glandulosa): responses to CO(2) enrichment.

Low water availability is a leading contributor to mortality of woody seedlings on grasslands, including those of the invasive shrub Prosopis. Increasing atmospheric CO(2) concentration could favor some genotypes of this species over others if there exists intraspecific variation in the responsiveness of survivorship to CO(2). To investigate such variation, we studied effects of CO(2) enrichment on seedling survival in response to uniform rates of soil water depletion in six maternal families of honey mesquite (P. glandulosa Torr. var. glandulosa). Three families each from the arid and mesic extremes of the species' distribution in the southwestern United States were studied in environmentally controlled glasshouses. Relative water content at turgor loss and osmotic potential were not affected by CO(2) treatment. Increased atmospheric CO(2) concentration, however, increased growth, leaf production and area, and midday xylem pressure potential, and apparently reduced transpiration per unit leaf area of seedlings as soil dried. Consequently, CO(2) enrichment about doubled the fraction of seedlings that survived soil water depletion. Maternal families of honey mesquite differed in percentage survival of drought and in several other characteristics, but differences were of similar or of smaller magnitude compared with differences between CO(2) treatments. There was no evidence for genetic variation in the responsiveness of survivorship to CO(2). By increasing seedling survival of drought, increasing atmospheric CO(2) concentration could increase the abundance of honey mesquite where establishment is limited by water availability. Genetic types with superior ability to survive drought today, however, apparently will maintain that advantage in the future.