Jack A. Morgan

Water relations in grassland and desert ecosystems exposed to elevated atmospheric CO2

Atmospheric CO2 enrichment may stimulate plant growth directly through (1) enhanced photosynthesis or indirectly, through (2) reduced plant water consumption and hence slower soil moisture depletion, or the combination of both. Herein we describe gas exchange, plant biomass and species responses of five native or semi-native temperate and Mediterranean grasslands and three semi-arid systems to CO2 enrichment, with an emphasis on water relations. Increasing CO2 led to decreased leaf conductance for water vapor, improved plant water status, altered seasonal evapotranspiration dynamics, and in most cases, periodic increases in soil water content. The extent, timing and duration of these responses varied among ecosystems, species and years. Across the grasslands of the Kansas tallgrass prairie, Colorado shortgrass steppe and Swiss calcareous grassland, increases in aboveground biomass from CO2 enrichment were relatively greater in dry years. In contrast, CO2-induced aboveground biomass increases in the Texas C3/C4 grassland and the New Zealand pasture seemed little or only marginally influenced by yearly variation in soil water, while plant growth in the Mojave Desert was stimulated by CO2 in a relatively wet year. Mediterranean grasslands sometimes failed to respond to CO2-related increased late-season water, whereas semiarid Negev grassland assemblages profited. Vegetative and reproductive responses to CO2 were highly varied among species and ecosystems, and did not generally follow any predictable pattern in regard to functional groups. Results suggest that the indirect effects of CO2 on plant and soil water relations may contribute substantially to experimentally induced CO2-effects, and also reflect local humidity conditions. For landscape scale predictions, this analysis calls for a clear distinction between biomass responses due to direct CO2 effects on photosynthesis and those indirect CO2 effects via soil moisture as documented here.

C4 grasses prosper as carbon dioxide eliminates desiccation in warmed semi-arid grassland

Global warming is predicted to induce desiccation in many world regions through increases in evaporative demand. Rising CO2 may counter that trend by improving plant water-use efficiency. However, it is not clear how important this CO2-enhanced water use efficiency might be in offsetting warming-induced desiccation because higher CO2 also leads to higher plant biomass, and therefore greater transpirational surface. Furthermore, although warming is predicted to favour warm-season, C4 grasses, rising CO2 should favour C3, or cool-season plants. Here we show in a semi-arid grassland that elevated CO2 can completely reverse the desiccating effects of moderate warming. Although enrichment of air to 600 p.p.m.v. CO2 increased soil water content (SWC), 1.5/3.0 °C day/night warming resulted in desiccation, such that combined CO2 enrichment and warming had no effect on SWC relative to control plots. As predicted, elevated CO2 favoured C3 grasses and enhanced stand productivity, whereas warming favoured C4 grasses. Combined warming and CO2 enrichment stimulated above-ground growth of C4 grasses in 2 of 3 years when soil moisture most limited plant productivity. The results indicate that in a warmer, CO2-enriched world, both SWC and productivity in semi-arid grasslands may be higher than previously expected.

Simple additive effects are rare: a quantitative review of plant biomass and soil process responses to combined manipulations of CO2 and temperature

In recent years, increased awareness of the potential interactions between rising atmospheric CO2 concentrations ([ CO2 ]) and temperature has illustrated the importance of multifactorial ecosystem manipulation experiments for validating Earth System models. To address the urgent need for increased understanding of responses in multifactorial experiments, this article synthesizes how ecosystem productivity and soil processes respond to combined warming and [ CO2 ] manipulation, and compares it with those obtained in single factor [ CO2 ] and temperature manipulation experiments. Across all combined elevated [ CO2 ] and warming experiments, biomass production and soil respiration were typically enhanced. Responses to the combined treatment were more similar to those in the [ CO2 ]-only treatment than to those in the warming-only treatment. In contrast to warming-only experiments, both the combined and the [ CO2 ]-only treatments elicited larger stimulation of fine root biomass than of aboveground biomass, consistently stimulated soil respiration, and decreased foliar nitrogen (N) concentration. Nonetheless, mineral N availability declined less in the combined treatment than in the [ CO2 ]-only treatment, possibly due to the warming-induced acceleration of decomposition, implying that progressive nitrogen limitation (PNL) may not occur as commonly as anticipated from single factor [ CO2 ] treatment studies. Responses of total plant biomass, especially of aboveground biomass, revealed antagonistic interactions between elevated [ CO2 ] and warming, i.e. the response to the combined treatment was usually less-than-additive. This implies that productivity projections might be overestimated when models are parameterized based on single factor responses. Our results highlight the need for more (and especially more long-term) multifactor manipulation experiments. Because single factor CO2 responses often dominated over warming responses in the combined treatments, our results also suggest that projected responses to future global warming in Earth System models should not be parameterized using single factor warming experiments.

Ecosystem resilience despite large-scale altered hydroclimatic conditions

Climate change is predicted to increase both drought frequency and duration, and when coupled with substantial warming, will establish a new hydroclimatological model for many regions. Large-scale, warm droughts have recently occurred in North America, Africa, Europe, Amazonia and Australia, resulting in major effects on terrestrial ecosystems, carbon balance and food security. Here we compare the functional response of above-ground net primary production to contrasting hydroclimatic periods in the late twentieth century (1975–1998), and drier, warmer conditions in the early twenty-first century (2000–2009) in the Northern and Southern Hemispheres. We find a common ecosystem water-use efficiency (WUEe: above-ground net primary production/evapotranspiration) across biomes ranging from grassland to forest that indicates an intrinsic system sensitivity to water availability across rainfall regimes, regardless of hydroclimatic conditions. We found higher WUEe in drier years that increased significantly with drought to a maximum WUEe across all biomes; and a minimum native state in wetter years that was common across hydroclimatic periods. This indicates biome-scale resilience to the interannual variability associated with the early twenty-first century drought—that is, the capacity to tolerate low, annual precipitation and to respond to subsequent periods of favourable water balance. These findings provide a conceptual model of ecosystem properties at the decadal scale applicable to the widespread altered hydroclimatic conditions that are predicted for later this century. Understanding the hydroclimatic threshold that will break down ecosystem resilience and alter maximum WUEe may allow us to predict land-surface consequences as large regions become more arid, starting with water-limited, low-productivity grasslands.

Carbon dioxide enrichment alters plant community structure and accelerates shrub growth in the shortgrass steppe

A hypothesis has been advanced that the incursion of woody plants into world grasslands over the past two centuries has been driven in part by increasing carbon dioxide concentration, [CO2], in Earths atmosphere. Unlike the warm season forage grasses they are displacing, woody plants have a photosynthetic metabolism and carbon allocation patterns that are responsive to CO2, and many have tap roots that are more effective than grasses for reaching deep soil water stores that can be enhanced under elevated CO2. However, this commonly cited hypothesis has little direct support from manipulative experimentation and competes with more traditional theories of shrub encroachment involving climate change, management, and fire. Here, we show that, although doubling [CO2] over the Colorado shortgrass steppe had little impact on plant species diversity, it resulted in an increasingly dissimilar plant community over the 5-year experiment compared with plots maintained at present-day [CO2]. Growth at the doubled [CO2] resulted in an ≈40-fold increase in aboveground biomass and a 20-fold increase in plant cover of Artemisia frigida Willd, a common subshrub of some North American and Asian grasslands. This CO2-induced enhancement of plant growth, among the highest yet reported, provides evidence from a native grassland suggesting that rising atmospheric [CO2] may be contributing to the shrubland expansions of the past 200 years. Encroachment of shrubs into grasslands is an important problem facing rangeland managers and ranchers; this process replaces grasses, the preferred forage of domestic livestock, with species that are unsuitable for domestic livestock grazing.

CO2 ENHANCES PRODUCTIVITY, ALTERS SPECIES COMPOSITION, AND REDUCES DIGESTIBILITY OF SHORTGRASS STEPPE VEGETATION

The impact of increasing atmospheric CO2 concentrations has been studied in a number of field experiments, but little information exists on the response of semiarid rangelands to CO2, or on the consequences for forage quality. This study was initiated to study the CO2 response of the shortgrass steppe, an important semiarid grassland on the western edge of the North American Great Plains, used extensively for livestock grazing. The experiment was conducted for five years on native vegetation at the USDA-ARS Central Plains Experimental Range in northeastern Colorado, USA. Three perennial grasses dominate the study site, Bouteloua gracilis, a C4 grass, and two C3 grasses, Pascopyrum smithii and Stipa comata. The three species comprise 88% of the aboveground phytomass. To evaluate responses to rising atmospheric CO2, we utilized six open-top chambers, three with ambient air and three with air CO2 enriched to 720 μmol/mol, as well as three unchambered controls. We found that elevated CO2 enhanced production o...

Rhizosphere priming: a nutrient perspective.

Rhizosphere priming is the change in decomposition of soil organic matter (SOM) caused by root activity. Rhizosphere priming plays a crucial role in soil carbon (C) dynamics and their response to global climate change. Rhizosphere priming may be affected by soil nutrient availability, but rhizosphere priming itself can also affect nutrient supply to plants. These interactive effects may be of particular relevance in understanding the sustained increase in plant growth and nutrient supply in response to a rise in atmospheric CO2 concentration. We examined how these interactions were affected by elevated CO2 in two similar semiarid grassland field studies. We found that an increase in rhizosphere priming enhanced the release of nitrogen (N) through decomposition of a larger fraction of SOM in one study, but not in the other. We postulate that rhizosphere priming may enhance N supply to plants in systems that are N limited, but that rhizosphere priming may not occur in systems that are phosphorus (P) limited. Under P limitation, rhizodeposition may be used for mobilization of P, rather than for decomposition of SOM. Therefore, with increasing atmospheric CO2 concentrations, rhizosphere priming may play a larger role in affecting C sequestration in N poor than in P poor soils.

Next generation of elevated [CO2] experiments with crops: A critical investment for feeding the future world

A rising global population and demand for protein-rich diets are increasing pressure to maximize agricultural productivity. Rising atmospheric [CO(2)] is altering global temperature and precipitation patterns, which challenges agricultural productivity. While rising [CO(2)] provides a unique opportunity to increase the productivity of C(3) crops, average yield stimulation observed to date is well below potential gains. Thus, there is room for improving productivity. However, only a fraction of available germplasm of crops has been tested for CO(2) responsiveness. Yield is a complex phenotypic trait determined by the interactions of a genotype with the environment. Selection of promising genotypes and characterization of response mechanisms will only be effective if crop improvement and systems biology approaches are closely linked to production environments, that is, on the farm within major growing regions. Free air CO(2) enrichment (FACE) experiments can provide the platform upon which to conduct genetic screening and elucidate the inheritance and mechanisms that underlie genotypic differences in productivity under elevated [CO(2)]. We propose a new generation of large-scale, low-cost per unit area FACE experiments to identify the most CO(2)-responsive genotypes and provide starting lines for future breeding programmes. This is necessary if we are to realize the potential for yield gains in the future.

Soil-atmosphere exchange of CH4, CO2, NOx, and N2O in the Colorado shortgrass steppe under elevated CO2

In late March 1997, an open-top-chamber (OTC) CO2 enrichment study was begun in the Colorado shortgrass steppe. The main objectives of the study were to determine the effect of elevated CO2 (∼720 μmol mol−1) on plant production, photosynthesis, and water use of this mixed C3/C4 plant community, soil nitrogen (N) and carbon (C) cycling and the impact of changes induced by CO2 on trace gas exchange. From this study, we report here our weekly measurements of CO2, CH4, NOx and N2O fluxes within control (unchambered), ambient CO2 and elevated CO2 OTCs. Soil water and temperature were measured at each flux measurement time from early April 1997, year round, through October 2000. Even though both C3 and C4 plant biomass increased under elevated CO2 and soil moisture content was typically higher than under ambient CO2 conditions, none of the trace gas fluxes were significantly altered by CO2 enrichment. Over the 43 month period of observation NOx and N2O flux averaged 4.3 and 1.7 in ambient and 4.1 and 1.7 μg N m−2 hr −1 in elevated CO2 OTCs, respectively. NOx flux was negatively correlated to plant biomass production. Methane oxidation rates averaged −31 and −34 μg C m−2 hr−1 and ecosystem respiration averaged 43 and 44 mg C m−2 hr−1 under ambient and elevated CO2, respectively, over the same time period.

Contrasting effects of elevated CO2 and warming on nitrogen cycling in a semiarid grassland

SUMMARY *Simulation models indicate that the nitrogen (N) cycle plays a key role in how other ecosystem processes such as plant productivity and carbon (C) sequestration respond to elevated CO(2) and warming. However, combined effects of elevated CO(2) and warming on N cycling have rarely been tested in the field. *Here, we studied N cycling under ambient and elevated CO(2) concentrations (600 micromol mol(-1)), and ambient and elevated temperature (1.5 : 3.0 degrees C warmer day:night) in a full factorial semiarid grassland field experiment in Wyoming, USA. We measured soil inorganic N, plant and microbial N pool sizes and NO(3)(-) uptake (using a (15)N tracer). *Soil inorganic N significantly decreased under elevated CO(2), probably because of increased microbial N immobilization, while soil inorganic N and plant N pool sizes significantly increased with warming, probably because of increased N supply. We observed no CO(2 )x warming interaction effects on soil inorganic N, N pool sizes or NO(3)(-) uptake in plants and microbes. *Our results indicate a more closed N cycle under elevated CO(2) and a more open N cycle with warming, which could affect long-term N retention, plant productivity, and C sequestration in this semiarid grassland.