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The effects of climate change on plants and ecosystems are mediated by plant hydraulic traits, including interspecific and intraspecific variability of trait phenotypes. Yet, integrative and realistic studies of hydraulic traits and climate change are rare. In a semiarid grassland, we assessed the response of several plant hydraulic traits to elevated CO 2 (+200 ppm) and warming (+1.5 to 3°C; day to night). For leaves of five dominant species (three graminoids and two forbs), and in replicated plots exposed to 7 years of elevated CO 2 , warming, or ambient climate, we measured: stomatal density and size, xylem vessel size, turgor loss point, and water potential (pre-dawn). Interspecific differences in hydraulic traits were larger than intraspecific shifts induced by elevated CO 2 and/or warming. Effects of elevated CO 2 were greater than effects of warming, and interactions between treatments were weak or not detected. The forbs showed little phenotypic plasticity. The graminoids had leaf water potentials and turgor loss points that were 10% to 50% less negative under elevated CO 2 ; thus, climate change might cause these species to adjust their drought resistance strategy away from tolerance and toward avoidance. The C4 grass also reduced allocation of leaf area to stomata under elevated CO 2 , which helps explain observations of higher soil moisture. The shifts in hydraulic traits under elevated CO 2 were not, however, simply due to higher soil moisture. Integration of our results with others' indicates that common species in this grassland are more likely to adjust stomatal aperture in response to near-term climate change, rather than anatomical traits; this contrasts with apparent effects of changing CO 2 on plant anatomy over evolutionary time. Future studies should assess how plant responses to drought may be constrained by the apparent shift from tolerance (via low turgor loss point) to avoidance (via stomatal regulation and/or access to deeper soil moisture)., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.)

Rising atmospheric carbon dioxide concentration should stimulate biomass production directly via biochemical stimulation of carbon assimilation, and indirectly via water savings caused by increased plant water-use efficiency. Because of these water savings, the CO 2 fertilization effect (CFE) should be stronger at drier sites, yet large differences among experiments in grassland biomass response to elevated CO 2 appear to be unrelated to annual precipitation, preventing useful generalizations. Here, we show that, as predicted, the impact of elevated CO 2 on biomass production in 19 globally distributed temperate grassland experiments reduces as mean precipitation in seasons other than spring increases, but that it rises unexpectedly as mean spring precipitation increases. Moreover, because sites with high spring precipitation also tend to have high precipitation at other times, these effects of spring and non-spring precipitation on the CO 2 response offset each other, constraining the response of ecosystem productivity to rising CO 2 . This explains why previous analyses were unable to discern a reliable trend between site dryness and the CFE. Thus, the CFE in temperate grasslands worldwide will be constrained by their natural rainfall seasonality such that the stimulation of biomass by rising CO 2 could be substantially less than anticipated.

Temporal variation in soil nitrogen (N) availability affects growth of grassland communities that differ in their use and reuse of N. In a 7-year-long climate change experiment in a semi-arid grassland, the temporal stability of plant biomass production varied with plant N turnover (reliance on externally acquired N relative to internally recycled N). Species with high N turnover were less stable in time compared to species with low N turnover. In contrast, N turnover at the community level was positively associated with asynchrony in biomass production, which in turn increased community temporal stability. Elevated CO 2 and summer irrigation, but not warming, enhanced community N turnover and stability, possibly because treatments promoted greater abundance of species with high N turnover. Our study highlights the importance of plant N turnover for determining the temporal stability of individual species and plant communities affected by climate change., (© 2018 John Wiley & Sons Ltd/CNRS.)

Increasing atmospheric [CO 2 ] and temperature are expected to affect the productivity, species composition, biogeochemistry, and therefore the quantity and quality of forage available to herbivores in rangeland ecosystems. Both elevated CO 2 (eCO 2 ) and warming affect plant tissue chemistry through multiple direct and indirect pathways, such that the cumulative outcomes of these effects are difficult to predict. Here, we report on a 7-yr study examining effects of CO 2 enrichment (to 600 ppm) and infrared warming (+1.5°C day/3°C night) under realistic field conditions on forage quality and quantity in a semiarid, mixedgrass prairie. For the three dominant forage grasses, warming effects on in vitro dry matter digestibility (IVDMD) and tissue [N] were detected only in certain years, varied from negative to positive, and were relatively minor. In contrast, eCO 2 substantially reduced IVDMD (two most abundant grasses) and [N] (all three dominant grass species) in most years, except the two wettest years. Furthermore, eCO 2 reduced IVDMD and [N] independent of warming effects. Reduced IVDMD with eCO 2 was related both to reduced [N] and increased acid detergent fiber (ADF) content of grass tissues. For the six most abundant forage species (representing 96% of total forage production), combined warming and eCO 2 increased forage production by 38% and reduced forage [N] by 13% relative to ambient climate. Although the absolute magnitude of the decline in IVDMD and [N] due to combined warming and eCO 2 may seem small (e.g., from 63.3 to 61.1% IVDMD and 1.25 to 1.04% [N] for Pascopyrum smithii), such shifts could have substantial consequences for the rate at which ruminants gain weight during the primary growing season in the largest remaining rangeland ecosystem in North America. With forage production increases, declining forage quality could potentially be mitigated by adaptively increasing stocking rates, and through management such as prescribed burning, fertilization at low rates, and legume interseeding to enhance forage quality., (© 2018 by the Ecological Society of America.)

Climate controls vegetation distribution across the globe, and some vegetation types are more vulnerable to climate change, whereas others are more resistant. Because resistance and resilience can influence ecosystem stability and determine how communities and ecosystems respond to climate change, we need to evaluate the potential for resistance as we predict future ecosystem function. In a mixed-grass prairie in the northern Great Plains, we used a large field experiment to test the effects of elevated CO2, warming, and summer irrigation on plant community structure and productivity, linking changes in both to stability in plant community composition and biomass production. We show that the independent effects of CO2 and warming on community composition and productivity depend on interannual variation in precipitation and that the effects of elevated CO2 are not limited to water saving because they differ from those of irrigation. We also show that production in this mixed-grass prairie ecosystem is not only relatively resistant to interannual variation in precipitation, but also rendered more stable under elevated CO2 conditions. This increase in production stability is the result of altered community dominance patterns: Community evenness increases as dominant species decrease in biomass under elevated CO2. In many grasslands that serve as rangelands, the economic value of the ecosystem is largely dependent on plant community composition and the relative abundance of key forage species. Thus, our results have implications for how we manage native grasslands in the face of changing climate.

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.)

In semiarid western North American riparian ecosystems, increased drought and lower streamflows under climate change may reduce plant growth and recruitment, and favor drought-tolerant exotic species over mesic native species. We tested whether elevated atmospheric CO₂ might ameliorate these effects by improving plant water-use efficiency. We examined the effects of CO₂ and water availability on seedlings of two native (Populus deltoides spp. monilifera, Salix exigua) and three exotic (Elaeagnus angustifolia, Tamarix spp., Ulmus pumila) western North American riparian species in a CO₂-controlled glasshouse, using 1-m-deep pots with different water-table decline rates. Low water availability reduced seedling biomass by 70-97%, and hindered the native species more than the exotics. Elevated CO₂ increased biomass by 15%, with similar effects on natives and exotics. Elevated CO₂ increased intrinsic water-use efficiency (Δ¹³C(leaf) ), but did not increase biomass more in drier treatments than wetter treatments. The moderate positive effects of elevated CO₂ on riparian seedlings are unlikely to counteract the large negative effects of increased aridity projected under climate change. Our results suggest that increased aridity will reduce riparian seedling growth despite elevated CO₂, and will reduce growth more for native Salix and Populus than for drought-tolerant exotic species., (No claim to original US government works. 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.

The effects of climate change on plants and ecosystems are mediated by plant hydraulic traits, including interspecific and intraspecific variability of trait phenotypes. Yet, integrative and realistic studies of hydraulic traits and climate change are rare. In a semiarid grassland, we assessed the response of several plant hydraulic traits to elevated CO2 (+200 ppm) and warming (+1.5 to 3°C; day to night). For leaves of five dominant species (three graminoids and two forbs), and in replicated plots exposed to 7 years of elevated CO2, warming, or ambient climate, we measured: stomatal density and size, xylem vessel size, turgor loss point, and water potential (pre‐dawn). Interspecific differences in hydraulic traits were larger than intraspecific shifts induced by elevated CO2 and/or warming. Effects of elevated CO2 were greater than effects of warming, and interactions between treatments were weak or not detected. The forbs showed little phenotypic plasticity. The graminoids had leaf water potentials and turgor loss points that were 10% to 50% less negative under elevated CO2; thus, climate change might cause these species to adjust their drought resistance strategy away from tolerance and toward avoidance. The C4 grass also reduced allocation of leaf area to stomata under elevated CO2, which helps explain observations of higher soil moisture. The shifts in hydraulic traits under elevated CO2 were not, however, simply due to higher soil moisture. Integration of our results with others' indicates that common species in this grassland are more likely to adjust stomatal aperture in response to near‐term climate change, rather than anatomical traits; this contrasts with apparent effects of changing CO2 on plant anatomy over evolutionary time. Future studies should assess how plant responses to drought may be constrained by the apparent shift from tolerance (via low turgor loss point) to avoidance (via stomatal regulation and/or access to deeper soil moisture). [ABSTRACT FROM AUTHOR]

Global changes that alter soil water availability may have profound effects on semiarid ecosystems. Although both elevated CO2 (eCO2) and warming can alter water availability, often in opposite ways, few studies have measured their combined influence on the amount, timing, and temporal variability of soil water. Here, we ask how free air CO2 enrichment (to 600 ppmv) and infrared warming (+ 1.5 °C day, + 3 °C night) effects on soil water vary within years and across wet-dry periods in North American mixed-grass prairie. We found that eCO2 and warming interacted to influence soil water and that those interactions varied by season. In the spring, negative effects of warming on soil water largely offset positive effects of eCO2. As the growing season progressed, however, warming reduced soil water primarily (summer) or only (autumn) in plots treated with eCO2. These interactions constrained the combined effect of eCO2 and warming on soil water, which ranged from neutral in spring to positive in autumn. Within seasons, eCO2 increased soil water under drier conditions, and warming decreased soil water under wetter conditions. By increasing soil water under dry conditions, eCO2 also reduced temporal variability in soil water. These temporal patterns explain previously observed plant responses, including reduced leaf area with warming in summer, and delayed senescence with eCO2 plus warming in autumn. They also suggest that eCO2 and warming may favor plant species that grow in autumn, including winter annuals and C3 graminoids, and species able to remain active under the dry conditions moderated by eCO2. [ABSTRACT FROM AUTHOR]

Plant roots mediate the impacts of environmental change on ecosystems, yet knowledge of root responses to environmental change is limited because few experiments evaluate multiple environmental factors and their interactions. Inferences about root functions are also limited because root length dynamics are rarely measured.Using a 5‐year experiment in a mixed‐grass prairie, we report the responses of root biomass, length and life span to elevated carbon dioxide (CO2), warming, elevated CO2 and warming combined, and irrigation. Root biomass was quantified using soil cores and root length dynamics were assessed using minirhizotrons. By comparing root dynamics with published results for soil resources and above‐ground productivity, we provide mechanistic insights into how climate change might impact grassland ecosystems.In the upper soil layer, 0–15 cm depth, both irrigation and elevated CO2 alone increased total root length by twofold, but irrigation decreased root biomass and elevated CO2 had only small positive effects on root biomass. The large positive effects of irrigation and elevated CO2 alone on total root length were due to increases in both root length production and root life span. The increased total root length and life span under irrigation and elevated CO2 coincided with apparent shifts from water limitation of plant growth to nitrogen limitation. Warming alone had minimal effects on root biomass, length and life span in this shallow soil layer. Warming and elevated CO2 combined increased root biomass and total root length by c. 25%, but total root length in this treatment was lower than expected if the effects of CO2 and warming alone were additive. Treatment effects on total root length and root life span varied with soil depth and root diameter.Synthesis. Sub‐additive effects of CO2 and warming suggest studies of elevated CO2 alone might overestimate the future capacity of grassland root systems to acquire resources. In this mixed‐grass prairie, elevated CO2 with warming stimulated total root length and root life span in deeper soils, likely enhancing plant access to more stable pools of growth‐limiting resources, including water and phosphorus. Thus, these root responses help explain previous observations of higher, and more stable, above‐ground productivity in these projected climate conditions. Sub‐additive effects of CO2 and warming suggest studies of elevated CO2 alone might overestimate the future capacity of grassland root systems to acquire resources. In this mixed‐grass prairie, elevated CO2 with warming stimulated total root length and root life span in deeper soils, likely enhancing plant access to more stable pools of growth‐limiting resources, including water and phosphorus. Thus, these root responses help explain previous observations of higher, and more stable, above‐ground productivity in these projected climate conditions [ABSTRACT FROM AUTHOR]

Vegetation greenness, detected using digital photography, is useful for monitoring phenology of plant growth, carbon uptake and water loss at the ecosystem level. Assessing ecosystem phenology by greenness is especially useful in spatially extensive, water-limited ecosystems such as the grasslands of the western United States, where productivity is moisture dependent and may become increasingly vulnerable to future climate change., We used repeat photography and a novel means of quantifying greenness in digital photographs to assess how the individual and combined effects of warming and elevated CO2 impact ecosystem phenology (greenness and plant cover) in a semi-arid grassland over an 8-year period., Climate variability within and among years was the proximate driver of ecosystem phenology. Individual and combined effects of warming and elevated CO2 were significant at times, but mediated by variation in both intra- and interannual precipitation. Specifically, warming generally enhanced plant cover and greenness early in the growing season but often had a negative effect during the middle of the summer, offsetting the early season positive effects. The individual effects of elevated CO2 on plant cover and greenness were generally neutral., Opposing seasonal variations in the effects of warming and less so elevated CO2 cancelled each other out over an entire growing season, leading to no net effect of treatments on annual accumulation of greenness. The main effect of elevated CO2 dampened quickly, but warming continued to affect plant cover and plot greenness throughout the experiment. The combination of warming and elevated CO2 had a generally positive effect on greenness, especially early in the growing season and in later years of the experiment, enhanced annual greenness accumulation. However, interannual precipitation variation had larger effect on greenness, with two to three times greater greenness in wet years than in dry years., Synthesis. Seasonal variation in timing and amount of precipitation governs grassland phenology, greenness and the potential for carbon uptake. Our results indicate that concurrent changes in precipitation regimes mediate vegetation responses to warming and elevated atmospheric CO2 in semi-arid grasslands. Even small changes in vegetation phenology and greenness in response to warming and rising atmospheric CO2 concentrations, such as those we report here, can have large consequences for the future of grasslands. [ABSTRACT FROM AUTHOR]

In semiarid western North American riparian ecosystems, increased drought and lower streamflows under climate change may reduce plant growth and recruitment, and favor drought-tolerant exotic species over mesic native species. We tested whether elevated atmospheric CO2 might ameliorate these effects by improving plant water-use efficiency., We examined the effects of CO2 and water availability on seedlings of two native ( Populus deltoides spp. monilifera, Salix exigua) and three exotic ( Elaeagnus angustifolia, Tamarix spp., Ulmus pumila) western North American riparian species in a CO2-controlled glasshouse, using 1-m-deep pots with different water-table decline rates., Low water availability reduced seedling biomass by 70-97%, and hindered the native species more than the exotics. Elevated CO2 increased biomass by 15%, with similar effects on natives and exotics. Elevated CO2 increased intrinsic water-use efficiency (Δ13Cleaf), but did not increase biomass more in drier treatments than wetter treatments., The moderate positive effects of elevated CO2 on riparian seedlings are unlikely to counteract the large negative effects of increased aridity projected under climate change. Our results suggest that increased aridity will reduce riparian seedling growth despite elevated CO2, and will reduce growth more for native Salix and Populus than for drought-tolerant exotic species. [ABSTRACT FROM AUTHOR]