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Multifactor experiments are often advocated as important for advancing terrestrial biosphere models (TBMs), yet to date, such models have only been tested against single-factor experiments. We applied 10 TBMs to the multifactor Prairie Heating and CO 2 Enrichment (PHACE) experiment in Wyoming, USA. Our goals were to investigate how multifactor experiments can be used to constrain models and to identify a road map for model improvement. We found models performed poorly in ambient conditions; there was a wide spread in simulated above-ground net primary productivity (range: 31-390 g C m -2  yr -1 ). Comparison with data highlighted model failures particularly with respect to carbon allocation, phenology, and the impact of water stress on phenology. Performance against the observations from single-factors treatments was also relatively poor. In addition, similar responses were predicted for different reasons across models: there were large differences among models in sensitivity to water stress and, among the N cycle models, N availability during the experiment. Models were also unable to capture observed treatment effects on phenology: they overestimated the effect of warming on leaf onset and did not allow CO 2 -induced water savings to extend the growing season length. Observed interactive (CO 2  × warming) treatment effects were subtle and contingent on water stress, phenology, and species composition. As the models did not correctly represent these processes under ambient and single-factor conditions, little extra information was gained by comparing model predictions against interactive responses. We outline a series of key areas in which this and future experiments could be used to improve model predictions of grassland responses to global change., (© 2017 John Wiley & Sons Ltd.)

Future ecosystem properties of grasslands will be driven largely by belowground biomass responses to climate change, which are challenging to understand due to experimental and technical constraints. We used a multi-faceted approach to explore single and combined impacts of elevated CO2 and warming on root carbon (C) and nitrogen (N) dynamics in a temperate, semiarid, native grassland at the Prairie Heating and CO2 Enrichment experiment. To investigate the indirect, moisture mediated effects of elevated CO2, we included an irrigation treatment. We assessed root standing mass, morphology, residence time and seasonal appearance/disappearance of community-aggregated roots, as well as mass and N losses during decomposition of two dominant grass species (a C3 and a C4). In contrast to what is common in mesic grasslands, greater root standing mass under elevated CO2 resulted from increased production, unmatched by disappearance. Elevated CO2 plus warming produced roots that were longer, thinner and had greater surface area, which, together with greater standing biomass, could potentially alter root function and dynamics. Decomposition increased under environmental conditions generated by elevated CO2, but not those generated by warming, likely due to soil desiccation with warming. Elevated CO2, particularly under warming, slowed N release from C4-but not C3-roots, and consequently could indirectly affect N availability through treatment effects on species composition. Elevated CO2 and warming effects on root morphology and decomposition could offset increased C inputs from greater root biomass, thereby limiting future net C accrual in this semiarid grassland.

As global changes reorganize plant communities, invasive plants may benefit. We hypothesized that elevated CO2 and warming would strongly influence invasive species success in a semi-arid grassland, as a result of both direct and water-mediated indirect effects. To test this hypothesis, we transplanted the invasive forb Linaria dalmatica into mixed-grass prairie treated with free-air CO2 enrichment and infrared warming, and followed survival, growth, and reproduction over 4 yr. We also measured leaf gas exchange and carbon isotopic composition in L. dalmatica and the dominant native C3 grass Pascopyrum smithii. CO2 enrichment increased L. dalmatica biomass 13-fold, seed production 32-fold, and clonal expansion seven-fold, while warming had little effect on L. dalmatica biomass or reproduction. Elevated CO2 decreased stomatal conductance in P. smithii, contributing to higher soil water, but not in L. dalmatica. Elevated CO2 also strongly increased L. dalmatica photosynthesis (87% versus 23% in P. smithii), as a result of both enhanced carbon supply and increased soil water. More broadly, rapid growth and less conservative water use may allow invasive species to take advantage of both carbon fertilization and water savings under elevated CO2 . Water-limited ecosystems may therefore be particularly vulnerable to invasion as CO2 increases., (No claim to original US goverment works. New Phytologist © 2013 New Phytologist Trust.)

The flux of carbon dioxide (CO2) between terrestrial ecosystems and the atmosphere may ameliorate or exacerbate climate change, depending on the relative responses of ecosystem photosynthesis and respiration to warming temperatures, rising atmospheric CO2, and altered precipitation. The combined effect of these global change factors is especially uncertain because of their potential for interactions and indirectly mediated conditions such as soil moisture. Here, we present observations of CO2 fluxes from a multi-factor experiment in semi-arid grassland that suggests a potentially strong climate - carbon cycle feedback under combined elevated [CO2] and warming. Elevated [CO2] alone, and in combination with warming, enhanced ecosystem respiration to a greater extent than photosynthesis, resulting in net C loss over four years. The effect of warming was to reduce respiration especially during years of below-average precipitation, by partially offsetting the effect of elevated [CO2] on soil moisture and C cycling. Carbon losses were explained partly by stimulated decomposition of soil organic matter with elevated [CO2]. The climate - carbon cycle feedback observed in this semiarid grassland was mediated by soil water content, which was reduced by warming and increased by elevated [CO2]. Ecosystem models should incorporate direct and indirect effects of climate change on soil water content in order to accurately predict terrestrial feedbacks and long-term storage of C in soil.

Atmospheric concentrations of methane (CH4 ) and nitrous oxide (N2 O) have increased over the last 150 years because of human activity. Soils are important sources and sinks of both potent greenhouse gases where their production and consumption are largely regulated by biological processes. Climate change could alter these processes thereby affecting both rate and direction of their exchange with the atmosphere. We examined how a rise in atmospheric CO2 and temperature affected CH4 and N2 O fluxes in a well-drained upland soil (volumetric water content ranging between 6% and 23%) in a semiarid grassland during five growing seasons. We hypothesized that responses of CH4 and N2 O fluxes to elevated CO2 and warming would be driven primarily by treatment effects on soil moisture. Previously we showed that elevated CO2 increased and warming decreased soil moisture in this grassland. We therefore expected that elevated CO2 and warming would have opposing effects on CH4 and N2 O fluxes. Methane was taken up throughout the growing season in all 5 years. A bell-shaped relationship was observed with soil moisture with highest CH4 uptake at intermediate soil moisture. Both N2 O emission and uptake occurred at our site with some years showing cumulative N2 O emission and other years showing cumulative N2 O uptake. Nitrous oxide exchange switched from net uptake to net emission with increasing soil moisture. In contrast to our hypothesis, both elevated CO2 and warming reduced the sink of CH4 and N2 O expressed in CO2 equivalents (across 5 years by 7% and 11% for elevated CO2 and warming respectively) suggesting that soil moisture changes were not solely responsible for this reduction. We conclude that in a future climate this semiarid grassland may become a smaller sink for atmospheric CH4 and N2 O expressed in CO2 -equivalents., (© 2013 Blackwell Publishing Ltd.)

Nitrogen (N) and phosphorus (P) are essential nutrients for primary producers and decomposers in terrestrial ecosystems. Although climate change affects terrestrial N cycling with important feedbacks to plant productivity and carbon sequestration, the impacts of climate change on the relative availability of N with respect to P remain highly uncertain. In a semiarid grassland in Wyoming, USA, we studied the effects of atmospheric CO(2) enrichment (to 600 ppmv) and warming (1.5/3.0°C above ambient temperature during the day/night) on plant, microbial and available soil pools of N and P. Elevated CO(2) increased P availability to plants and microbes relative to that of N, whereas warming reduced P availability relative to N. Across years and treatments, plant N : P ratios varied between 5 and 18 and were inversely related to soil moisture. Our results indicate that soil moisture is important in controlling P supply from inorganic sources, causing reduced P relative to N availability during dry periods. Both wetter soil conditions under elevated CO(2) and drier conditions with warming can further alter N : P. Although warming may alleviate N constraints under elevated CO(2) , warming and drought can exacerbate P constraints on plant growth and microbial activity in this semiarid grassland., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)

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.

Carbon allocation and N acquisition by plants following defoliation may be linked through plant-microbe interactions in the rhizosphere. Plant C allocation patterns and rhizosphere interactions can also be affected by rising atmospheric CO(2) concentrations, which in turn could influence plant and microbial responses to defoliation. We studied two widespread perennial grasses native to rangelands of western North America to test whether (1) defoliation-induced enhancement of rhizodeposition would stimulate rhizosphere N availability and plant N uptake, and (2) defoliation-induced enhancement of rhizodeposition, and associated effects on soil N availability, would increase under elevated CO(2). Both species were grown at ambient (400 μL L(-1)) and elevated (780 μL L(-1)) atmospheric [CO(2)] under water-limiting conditions. Plant, soil and microbial responses were measured 1 and 8 days after a defoliation treatment. Contrary to our hypotheses, we found that defoliation and elevated CO(2) both reduced carbon inputs to the rhizosphere of Bouteloua gracilis (C(4)) and Pascopyrum smithii (C(3)). However, both species also increased N allocation to shoots of defoliated versus non-defoliated plants 8 days after treatment. This response was greatest for P. smithii, and was associated with negative defoliation effects on root biomass and N content and reduced allocation of post-defoliation assimilate to roots. In contrast, B. gracilis increased allocation of post-defoliation assimilate to roots, and did not exhibit defoliation-induced reductions in root biomass or N content. Our findings highlight key differences between these species in how post-defoliation C allocation to roots versus shoots is linked to shoot N yield, but indicate that defoliation-induced enhancement of shoot N concentration and N yield is not mediated by increased C allocation to the rhizosphere.

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.

Predicting net C balance under future global change scenarios requires a comprehensive understanding of how ecosystem photosynthesis (gross primary production; GPP) and respiration (Re) respond to elevated atmospheric [CO(2)] and altered water availability. We measured net ecosystem exchange of CO(2) (NEE), GPP and Re under ambient and elevated [CO(2)] in a northern mixed-grass prairie (Wyoming, USA) during dry intervals and in response to simulated precipitation pulse events. Elevated [CO(2)] resulted in higher rates of both GPP and Re across the 2006 growing season, and the balance of these two fluxes (NEE) accounted for cumulative growing season C uptake (-14.4 +/- 8.3 g C m(-2)). Despite lower GPP and Re, experimental plots under ambient [CO(2)] had greater cumulative uptake (-36.2 +/- 8.2 g C m(-2)) than plots under elevated [CO(2)]. Non-irrigated control plots received 50% of average precipitation during the drought of 2006, and had near-zero NEE (1.9 +/- 6.4 g C m(-2)) for the growing season. Elevated [CO(2)] extended the magnitude and duration of pulse-related increases in GPP, resulting in a significant [CO(2)] treatment by pulse day interaction, demonstrating the potential for elevated [CO(2)] to increase the capacity of this ecosystem to respond to late-season precipitation. However, stimulation of Re throughout the growing season under elevated [CO(2)] reduced net C uptake compared to plots under ambient [CO(2)]. These results indicate that although elevated [CO(2)] stimulates gross rates of ecosystem C fluxes, it does not necessarily enhance net C uptake, and that C cycle responses in semi-arid grasslands are likely to be more sensitive to changes in precipitation than atmospheric [CO(2)].

In semi-arid regions, where plants using both C(3) and C(4) photosynthetic pathways are common, the stable C isotope ratio (delta(13)C) of ecosystem respiration (delta(13)C(R)) is strongly variable seasonally and inter-annually. Improved understanding of physiological and environmental controls over these variations will improve C cycle models that rely on the isotopic composition of atmospheric CO(2). We hypothesized that timing of precipitation events and antecedent moisture interact with activity of C(3) and C(4) grasses to determine net ecosystem CO(2) exchange (NEE) and delta(13)C(R). Field measurements included CO(2) and delta(13)C fluxes from the whole ecosystem and from patches of different plant communities, biomass and delta(13)C of plants and soils over the 2000 and 2001 growing seasons. NEE shifted from C source to sink in response to rainfall events, but this shift occurred after a time lag of up to 2 weeks if a dry period preceded the rainfall. The seasonal average of delta(13)C(R) was higher in 2000 (-16 per thousand) than 2001 (20 per thousand), probably due to drier conditions during the 2000 growing season (79.7 mm of precipitation from April up to and including July) than in 2001 (189 mm). During moist conditions, delta(13)C averaged -22 per thousand from C(3) patches, -16 per thousand from C(4) patches, and -19 per thousand from mixed C(3) and C(4) patches. However, during dry conditions the apparent spatial differences were not obvious, suggesting reduced autotrophic activity in C(4) grasses with shallow rooting depth, soon after the onset of dry conditions. Air and soil temperatures were negatively correlated with delta(13)C(R); vapor pressure deficit was a poor predictor of delta(13)C(R), in contrast to more mesic ecosystems. Responses of respiration components to precipitation pulses were explained by differences in soil moisture thresholds between C(3) and C(4) species. Stable isotopic composition of respiration in semi-arid ecosystems is more temporally and spatially variable than in mesic ecosystems owing to dynamic aspects of pulse precipitation episodes and biological drivers.

Although global change is known to influence plant invasion, little is known about interactions between altered precipitation and invasion. In the North American mixedgrass prairie, invasive species are often abundant in wet and nitrogen (N)-rich areas, suggesting that predicted changes in precipitation and N deposition could exacerbate invasion. Here, this possibility was tested by seeding six invasive species into experimental plots of mixedgrass prairie treated with a factorial combination of increased snow, summer irrigation, and N addition. Without added snow, seeded invasive species were rarely observed. Snow addition increased average above-ground biomass of Centaurea diffusa from 0.026 to 66 g m(-2), of Gypsophila paniculata from 0.1 to 7.3 g m(-2), and of Linaria dalmatica from 5 to 101 g m(-2). Given added snow, summer irrigation increased the density of G. paniculata, and N addition increased the density and biomass of L. dalmatica. Plant density responses mirrored those of plant biomass, indicating that increases in biomass resulted, in part, from increases in recruitment. In contrast to seeded invasive species, resident species did not respond to snow addition. These results suggest that increases in snowfall or variability of snowfall may exacerbate forb invasion in the mixedgrass prairie.

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, [CO(2)], in Earth's atmosphere. Unlike the warm season forage grasses they are displacing, woody plants have a photosynthetic metabolism and carbon allocation patterns that are responsive to CO(2), and many have tap roots that are more effective than grasses for reaching deep soil water stores that can be enhanced under elevated CO(2). 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 [CO(2)] 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 [CO(2)]. Growth at the doubled [CO(2)] resulted in an approximately 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 CO(2)-induced enhancement of plant growth, among the highest yet reported, provides evidence from a native grassland suggesting that rising atmospheric [CO(2)] 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.

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.

Levels of UV were manipulated in a native shortgrass steppe using open-sided structures with tops that either passed or blocked wavelengths shorter than approximately 370 nm. Precipitation was controlled to create a drought or a very wet year. Subplots were either nondefoliated or defoliated to simulate grazing by livestock, which is the primary land use. Plant community productivity and forage quality were assessed in response to the two climate change variables (UV, precipitation) and grazing stress. Productivity and seasonal standing biomass of the dominant grass species were negatively affected by passing versus blocking UV, but only in the dry year. Another species was negatively affected by passing UV in the wet year, indicating the potential for future shifts in species composition. Forage quality for ruminants increased when UV was passed compared with blocked, as determined by in vitro digestible dry matter, depending on species and precipitation. Nitrogen concentrations and soluble and fiber components of vegetation also displayed some UV effects, but they were generally small and depended on species, season or amount of precipitation (or all). Grazing treatment had large positive effects on current-year productivity only in the wet year and some small positive effects on quality in both wet and dry years. Interactions between UV and grazing treatment were not observed.

We investigated the influence of long-term (56 years) grazing on organic and inorganic carbon (C) and nitrogen (N) contents of the plant-soil system (to 90 cm depth) in shortgrass steppe of northeastern Colorado. Grazing treatments included continuous season-long (May-October) grazing by yearling heifers at heavy (60-75% utilization) and light (20-35% utilization) stocking rates, and nongrazed exclosures. The heavy stocking rate resulted in a plant community that was dominated (75% of biomass production) by the C4 grass blue grama (Bouteloua gracilis), whereas excluding livestock grazing increased the production of C3 grasses and prickly pear cactus (Opuntia polycantha). Soil organic C (SOC) and organic N were not significantly different between the light grazing and nongrazed treatments, whereas the heavy grazing treatment was 7.5 Mg ha(-1) higher in SOC than the nongrazed treatment. Lower ratios of net mineralized N to total organic N in both grazed compared to nongrazed treatments suggest that long-term grazing decreased the readily mineralizable fraction of soil organic matter. Heavy grazing affected soil inorganic C (SIC) more than the SOC. The heavy grazing treatment was 23.8 Mg ha(-1) higher in total soil C (0-90 cm) than the nongrazed treatment, with 68% (16.3 Mg ha(-1)) attributable to higher SIC, and 32% (7.5 Mg ha(-1)) to higher SOC. These results emphasize the importance in semiarid and arid ecosystems of including inorganic C in assessments of the mass and distribution of plant-soil C and in evaluations of the impacts of grazing management on C sequestration.

To model the effect of increasing atmospheric CO2 on semi-arid grasslands, the gas exchange responses of leaves to seasonal changes in soil water, and how they are modified by CO2, must be understood for C3 and C4 species that grow in the same area. In this study, open-top chambers were used to investigate the photosynthetic and stomatal responses of Pascopyrum smithii (C3) and Bouteloua gracilis (C4) grown at 360 (ambient CO2) and 720 micro mol mol-1 CO2 (elevated CO2) in a semi-arid shortgrass steppe. Assimilation rate (A) and stomatal conductance (gs) at the treatment CO2 concentrations and at a range of intercellular CO2 concentrations and leaf water potentials (psileaf) were measured over 4 years with variable soil water content caused by season and CO2 treatment. Carboxylation efficiency of ribulose bisphosphate carboxylase/oxygenase (Vc,max), and ribulose bisphosphate regeneration capacity (Jmax) were reduced in P. smithii grown in elevated CO2, to the degree that A was similar in elevated and ambient CO2 (when soil moisture was adequate). Photosynthetic capacity was not reduced in B. gracilis under elevated CO2, but A was nearly saturated at ambient CO2. There were no stomatal adaptations independent of photosynthetic acclimation. Although photosynthetic capacity was reduced in P. smithii growing in elevated CO2, reduced gs and transpiration improved soil water content and psileaf in the elevated CO2 chambers, thereby improving A of both species during dry periods. These results suggest that photosynthetic responses of C3 and C4 grasses in this semi-arid ecosystem will be driven primarily by the effect of elevated CO2 on plant and soil water relations.