Global warming is predicted to induce desiccation in many world regions through increases in evaporative demand. Rising CO(2) may counter that trend by improving plant water-use efficiency. However, it is not clear how important this CO(2)-enhanced water use efficiency might be in offsetting warming-induced desiccation because higher CO(2) also leads to higher plant biomass, and therefore greater transpirational surface. Furthermore, although warming is predicted to favour warm-season, C(4) grasses, rising CO(2) should favour C(3), or cool-season plants. Here we show in a semi-arid grassland that elevated CO(2) can completely reverse the desiccating effects of moderate warming. Although enrichment of air to 600 p.p.m.v. CO(2) increased soil water content (SWC), 1.5/3.0 °C day/night warming resulted in desiccation, such that combined CO(2) enrichment and warming had no effect on SWC relative to control plots. As predicted, elevated CO(2) favoured C(3) grasses and enhanced stand productivity, whereas warming favoured C(4) grasses. Combined warming and CO(2) enrichment stimulated above-ground growth of C(4) grasses in 2 of 3 years when soil moisture most limited plant productivity. The results indicate that in a warmer, CO(2)-enriched world, both SWC and productivity in semi-arid grasslands may be higher than previously expected.

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.

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.

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 (WUE(e): 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 WUE(e) in drier years that increased significantly with drought to a maximum WUE(e) 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 WUE(e) may allow us to predict land-surface consequences as large regions become more arid, starting with water-limited, low-productivity grasslands.

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.

There is a need for methodology to warm open-field plots in order to study the likely effects of global warming on ecosystems in the future. Herein, we describe the development of arrays of more powerful and efficient infrared heaters with ceramic heating elements. By tilting the heaters at 451 from horizontal and combining six of them in a hexagonal array, good uniformity of warming was achieved across 3-m-diameter plots. Moreover, there do not appear to be obstacles (other than financial) to scaling to larger plots. The efficiency [g h (%); thermal radiation out per electrical energy in] of these heaters was higher than that of the heaters used in most previous infrared heater experiments and can be described by: g h 5 10 1 25exp(À0.17 u), where u is wind speed at 2 m height (m s À1 ). Graphs are presented to estimate operating costs from degrees of warming, two types of plant canopy, and site windiness. Four such arrays were deployed over plots of grass at Haibei, Qinghai, China and another at Cheyenne, Wyoming, USA, along with corresponding reference plots with dummy heaters. Proportional integral derivative systems with infrared thermometers to sense canopy temperatures of the heated and reference plots were used to control the heater outputs. Over month-long periods at both sites, about 75% of canopy temperature observations were within 0.5 1C of the set-point temperature differences between heated and reference plots. Electrical power consumption per 3-m-diameter plot averaged 58 and 80 kW h day À1 for Haibei and Cheyenne, respectively. However, the desired temperature differences were set lower at Haibei (1.2 1C daytime, 1.7 1C night) than Cheyenne (1.5 1C daytime, 3.0 1C night), and Cheyenne is a windier site. Thus, we conclude that these hexagonal arrays of ceramic infrared heaters can be a successful temperature free-air-controlled enhancement (T-FACE) system for warming ecosystem field plots.

The amplified ''greenhouse effect'' associated with increasing concentrations of greenhouse gases has increased atmospheric temperature by 18C since industrialization (around 1750), and it is anticipated to cause an additional 28C increase by midcentury. Increased biospheric warming is also projected to modify the amount and distribution of annual precipitation and increase the occurrence of both drought and heat waves. The ecological consequences of climate change will vary substantially among ecoregions because of regional differences in antecedent environmental conditions; the rate and magnitude of change in the primary climate change drivers, including elevated carbon dioxide (CO 2), warming and precipitation modification; and nonadditive effects among climate drivers. Elevated atmospheric CO 2 will directly stimulate plant growth and reduce negative effects of drying in a warmer climate by increasing plant water use efficiency; however, the CO 2 effect is mediated by environmental conditions, especially soil water availability. Warming and drying are anticipated to reduce soil water availability, net primary productivity, and other ecosystem processes in the southern Great Plains, the Southwest, and northern Mexico, but warmer and generally wetter conditions will likely enhance these processes in the northern Plains and southern Canada. The Northwest will warm considerably, but annual precipitation is projected to change little despite a large decrease in summer precipitation. Reduced winter snowpack and earlier snowmelt will affect hydrology and riparian systems in the Northwest. Specific consequences of climate change will be numerous and varied and include modifications to forage quantity and quality and livestock production systems, soil C content, fire regimes, livestock metabolism, and plant community composition and species distributions, including range contraction and expansion of invasive species. Recent trends and model projections indicate continued directional change and increasing variability in climate that will substantially affect the provision of ecosystem services on North American rangelands.

Summary Six open‐top chambers were installed on the shortgrass steppe in north‐eastern Colorado, USA from late March until mid‐October in 1997 and 1998 to evaluate how this grassland will be affected by rising atmospheric CO2. Three chambers were maintained at current CO2 concentration (ambient treatment), three at twice ambient CO2, or approximately 720 μmol mol−1 (elevated treatment), and three nonchambered plots served as controls. Above‐ground phytomass was measured in summer and autumn during each growing season, soil water was monitored weekly, and leaf photosynthesis, conductance and water potential were measured periodically on important C3 and C4 grasses. Mid‐season and seasonal above‐ground productivity were enhanced from 26 to 47% at elevated CO2, with no differences in the relative responses of C3/C4 grasses or forbs. Annual above‐ground phytomass accrual was greater on plots which were defoliated once in mid‐summer compared to plots which were not defoliated during the growing season, but there was no interactive effect of defoliation and CO2 on growth. Leaf photosynthesis was often greater in Pascopyrum smithii (C3) and Bouteloua gracilis (C4) plants in the elevated chambers, due in large part to higher soil water contents and leaf water potentials. Persistent downward photosynthetic acclimation in P. smithii leaves prevented large photosynthetic enhancement for elevated CO2‐grown plants. Shoot N concentrations tended to be lower in grasses under elevated CO2, but only Stipa comata (C3) plants exhibited significant reductions in N under elevated compared to ambient CO2 chambers. Despite chamber warming of 2.6 °C and apparent drier chamber conditions compared to unchambered controls, above‐ground production in all chambers was always greater than in unchambered plots. Collectively, these results suggest increased productivity of the shortgrass steppe in future warmer, CO2 enriched environments.