Photosynthetic response of two shrubs to rainfall pulses in desert regions of northwestern China

Pulses of rainfall are particularly pivotal in controlling plant physiological processes in ecosystems controlled by limited water, and the response of desert plants to rainfall is a key to understanding the responses of desert ecosystems to global climatic change. We used a portable photosynthesis system to measure the responses of the diurnal course of photosynthesis, light-response curves, and CO₂-response curves of two desert shrubs (Nitraria sphaerocarpa Maxim. and Calligonum mongolicum Turcz) to a rainfall pulse in a desert-oasis ecotone in northwestern China. The photosynthetic parameters, light- and CO₂-response curves differed significantly before and after the rainfall pulse. Their maximum net photosynthetic rate (P _N) values were 23.27 and 32.92 μmol(CO₂) m⁻² s⁻¹ for N. sphaerocarpa and C. mongolicum, respectively, with corresponding maximum stomatal conductance (g _s) values of 0.47 and 0.39 mol(H₂O) m⁻² s⁻¹. The P _N of N. sphaerocarpa after the rainfall was 1.65 to 1.75 times the value before rainfall, whereas those of C. mongolicum increased to approximately 2 times the prerainfall value, demonstrating the importance of the desert plants response by improving their assimilation rate to precipitation patterns under a future climate.     

Verification of a threshold concept of ecologically effective precipitation pulse: From plant individuals to ecosystem

In water-limited ecosystems, an ecologically significant rainfall pulse was defined as a rainfall event that altered both soil water status and plant physiological activity. We developed a new threshold concept of an ecologically effective precipitation pulse (EEPP) applicable to both plant individual and ecosystem scales. The concept was tested in a typical steppe on Inner Mongolia plateau. Two EEPPs, single 3-mm rainfall and 5-mm rainfall, were applied to investigate their effects on soil and plant water status, CO₂ assimilation of five species (four C₃ plants and one C₄ plant), whole-plot soil respiration (R_s), and net ecosystem CO₂ exchange (NEE) on 1 June and 28 July 2009, respectively. Both EEPPs increased leaf water potential (Ψ_l) of all the species, which peaked 1–3 days after rainfall pulses. Soil water content (SWC) in two depths (5 cm and 20 cm) significantly increased after the two EEPPs for 1–3 days. Soil water potential (Ψ_s) within 20‐cm soil layer in EEPP treatments significantly differed (p < 0.05) from control. Net assimilation rates (A_net) of all C₃ plants had a slight increase at the next day after two EEPPs, in contrast to the C₄ species. R_s elevated and peaked 1–3 days later after water supply. Ecosystem net CO₂ absorption rate rose to maximum value 3 days after the 5-mm pulse on 28 July, higher than the response to 3-mm pulse on June 1. The grassland turned to net emission of CO₂ after 3-mm pulse on 28 July. The results supported that there was an ecosystem level threshold for EEPP, and the threshold was temporally variable. It also highlighted the necessity of considering the response threshold of EEPP in rainfall manipulative experiment. In addition, effective rainfall amount was more approriate than total rainfall amount in modeling ecosystem carbon balance.     

Primary Productivity and Water Balance of Grassland Vegetation on Three Soils in a Continuous CO2 Gradient: Initial Results from the Lysimeter CO2 Gradient Experiment

Field studies of atmospheric CO₂ effects on ecosystems usually include few levels of CO₂ and a single soil type, making it difficult to ascertain the shape of responses to increasing CO₂ or to generalize across soil types. The Lysimeter CO₂ Gradient (LYCOG) chambers were constructed to maintain a linear gradient of atmospheric CO₂ (~250 to 500 μl l⁻¹) on grassland vegetation established on intact soil monoliths from three soil series. The chambers maintained a linear daytime CO₂ gradient from 263 μl l⁻¹ at the subambient end of the gradient to 502 μl l⁻¹ at the superambient end, as well as a linear nighttime CO₂ gradient. Temperature variation within the chambers affected aboveground biomass and evapotranspiration, but the effects of temperature were small compared to the expected effects of CO₂. Aboveground biomass on Austin soils was 40% less than on Bastrop and Houston soils. Biomass differences between soils resulted from variation in biomass of Sorghastrum nutans, Bouteloua curtipendula, Schizachyrium scoparium (C₄ grasses), and Solidago canadensis (C₃ forb), suggesting the CO₂ sensitivity of these species may differ among soils. Evapotranspiration did not differ among the soils, but the CO₂ sensitivity of leaf-level photosynthesis and water use efficiency in S. canadensis was greater on Houston and Bastrop than on Austin soils, whereas the CO₂ sensitivity of soil CO₂ efflux was greater on Bastrop soils than on Austin or Houston soils. The effects of soil type on CO₂ sensitivity may be smaller for some processes that are tightly coupled to microclimate. LYCOG is useful for discerning the effects of soil type on the CO₂ sensitivity of ecosystem function in grasslands. Author Contributions: PF conceived study, analyzed data, and wrote the paper. AK, AP analyzed data. DH, VJ, RJ, HJ, and WP conceived study, and conducted research.     

REVIEW: Time lag between photosynthesis and carbon dioxide efflux from soil: a review of mechanisms and controls

CO₂ efflux from soil depends on the availability of organic substances respired by roots and microorganisms. Therefore, photosynthetic activity supplying carbohydrates from leaves to roots and rhizosphere is a key driver of soil CO₂. This fact has been overlooked in most soil CO₂ studies because temperature variations are highly correlated with solar radiation and mask the direct effect of photosynthesis on substrate availability in soil. This review highlights the importance of photosynthesis for rhizosphere processes and evaluates the time lag between carbon (C) assimilation and CO₂ release from soil. Mechanisms and processes contributing to the lag were evaluated. We compared the advantages and shortcomings of four main approaches used to estimate this time lag: (1) interruption of assimilate flow from leaves into the roots and rhizosphere, and analysis of the decrease of CO₂ efflux from soil, (2) time series analysis (TSA) of CO₂ fluxes from soil and photosynthesis proxies, (3) analysis of natural δ¹³C variation in CO₂ with photosynthesis‐related parameters or δ¹³C in the phloem and leaves, and (4) pulse labeling of plants in artificial ¹⁴CO₂ or ¹³CO₂ atmosphere with subsequent tracing of ¹⁴C or ¹³C in CO₂ efflux from soil. We concluded that pulse labeling is the most advantageous approach. It allows clear evaluation not only of the time lag, but also of the label dynamics in soil CO₂, and helps estimate the mean residence time of recently assimilated C in various above‐ and belowground C pools. The impossibility of tracing the phloem pressure–concentration waves by labeling approach may be overcome by its combination with approaches based on TSA of CO₂ fluxes and its δ¹³C with photosynthesis proxies. Numerous studies showed that the time lag for grasses is about 12.5±7.5 (SD) h. The time lag for mature trees was much longer (～4–5 days). Tree height slightly affected the lag, with increasing delay of 0.1 day m^?1. By evaluating bottle‐neck processes responsible for the time lag, we conclude that, for trees, the transport of assimilates in phloem is the rate‐limiting step. However, it was not possible to predict the lag based on the phloem transport rates reported in the literature. We conclude that studies of CO₂ fluxes from soil, especially in ecosystems with a high contribution of root‐derived CO₂, should consider photosynthesis as one of the main drivers of C fluxes. This calls for incorporating photosynthesis in soil C turnover models.     

Increases in Desert Shrub Productivity under Elevated Carbon Dioxide Vary with Water Availability

Productivity of aridland plants is predicted to increase substantially with rising atmospheric carbon dioxide (CO₂) concentrations due to enhancement in plant water-use efficiency (WUE). However, to date, there are few detailed analyses of how intact desert vegetation responds to elevated CO₂. From 1998 to 2001, we examined aboveground production, photosynthesis, and water relations within three species exposed to ambient (around 38 Pa) or elevated (55 Pa) CO₂ concentrations at the Nevada Desert Free-Air CO₂ Enrichment (FACE) Facility in southern Nevada, USA. The functional types sampled—evergreen (Larrea tridentata), drought-deciduous (Ambrosia dumosa), and winter-deciduous shrubs (Krameria erecta)—represent potentially different responses to elevated CO₂ in this ecosystem. We found elevated CO₂ significantly increased aboveground production in all three species during an anomalously wet year (1998), with relative production ratios (elevated:ambient CO₂) ranging from 1.59 (Krameria) to 2.31 (Larrea). In three below-average rainfall years (1999–2001), growth was much reduced in all species, with only Ambrosia in 2001 having significantly higher production under elevated CO₂. Integrated photosynthesis (mol CO₂ m⁻² y⁻¹) in the three species was 1.26–2.03-fold higher under elevated CO₂ in the wet year (1998) and 1.32–1.43-fold higher after the third year of reduced rainfall (2001). Instantaneous WUE was also higher in shrubs grown under elevated CO₂. The timing of peak canopy development did not change under elevated CO₂; for example, there was no observed extension of leaf longevity into the dry season in the deciduous species. Similarly, seasonal patterns in CO₂ assimilation did not change, except for Larrea. Therefore, phenological and physiological patterns that characterize Mojave Desert perennials—early-season lags in canopy development behind peak photosynthetic capacity, coupled with reductions in late-season photosynthetic capacity prior to reductions in leaf area—were not significantly affected by elevated CO₂. Together, these findings suggest that elevated CO₂ can enhance the productivity of Mojave Desert shrubs, but this effect is most pronounced during years with abundant rainfall when soil resources are most available.     

Temporal variation in CO<Subscript>2</Subscript> efflux from soil and snow surfaces in a Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis>) plantation,central Japan

CO₂ efflux from soil and snow surfaces was measured continuously in a Japanese cedar (Cryptomeria japonica D. Don) forest in central Japan using an open dynamic chamber system. The chamber opens and closes automatically and records measurements based on an open-flow dynamic method. Between May and December, mean soil CO₂ efflux ranged from 1,529 mg CO₂ m⁻² h⁻¹ in September to 255 mg CO₂ m⁻² h⁻¹ in December. The seasonal change in CO₂ efflux from the soil paralleled the seasonal pattern of soil temperature. No marked diurnal trends in soil CO₂ efflux were observed on days without rainfall, whereas significant pulses in soil CO₂ efflux were observed on days with rainfall. In this plantation, soil CO₂ efflux frequently responded to rainfall. Measurements of changes from litter-covered soil to snow-covered surfaces revealed that CO₂ efflux decreased from values of ca. 250 mg CO₂ m⁻² h⁻¹ above soil to less than 33 mg CO₂ m⁻² h⁻¹ above snow. Soil temperature alone explained 66% of the overall variation in soil CO₂ efflux, but explained approximately 85% of the variation when data from two anomalous periods were excluded. Moreover, we found a significant correlation between soil CO₂ efflux and soil moisture (which explained 44% of the overall variation) using a second-order polynomial function. Our results suggest that the seasonality of CO₂ efflux is affected not only by soil temperature and moisture, but also by drying and rewetting cycles and by litterfall pulses.     

Sensitivity of water relations and photosynthesis to summer precipitation pulses for <Emphasis Type="Italic">Artemisia tridentata</Emphasis> and <Emphasis Type="Italic">Purshia tridentata</Emphasis>

For much of the western USA, precipitation occurs in pulses, the nature of which determine soil water potential and plant physiological performance. This research utilized three experiments to examine the sensitivity of photosynthesis and water relations for two widespread Great Basin Desert shrub species, Artemisia tridentata (which has both deep and shallow roots) and Purshia tridentata (which reportedly has only deep roots), to (1) variation in pulse magnitude size, (2) the kinetics of responses to pulses, and (3) the relationship between pulse-size and antecedent soil water content. At the study site in the southwestern Great Basin Desert, USA, summer rainfall exhibits a greater frequency of larger-sized events, and longer inter-pulse intervals, compared to annual patterns. Compared to pre-watering values, stem water potential initially increased by about 2.00 MPa for A. tridentata and 1.00 MPa for P. tridentata following watering to simulate an 11.5 mm rainfall pulse. For the same water addition, stomatal conductance increased by 0.3 mol m⁻² s⁻¹ and photosynthetic CO₂ assimilation increased 8-fold for A. tridentata and 6-fold for P. tridentata. Water potential and photosynthetic gas exchange were maximal for both species 2–3 days following a pulse addition. In comparison to P. tridentata, the increase in photosynthesis for A. tridentata was more pronounced for plants treated incrementally with several small pulses compared to plants treated with one pulse of an equivalent total volume. The results indicate that both species can respond to a range of summer rainfall pulse magnitudes within about 2 days, with A. tridentata generally exhibiting larger responses in comparison to the co-dominant shrub species P. tridentata, which at this study site does indeed have shallow roots. In a future climate, the timing and magnitude of summer rainfall pulses will determine the extent to which these two species undergo changes in water status and photosynthetic carbon uptake, with implications for their fitness.     

Effects of Experimental Water Table and Temperature Manipulations on Ecosystem CO2 Fluxes in an Alaskan Rich Fen

Peatlands store 30% of the world’s terrestrial soil carbon (C) and those located at northern latitudes are expected to experience rapid climate warming. We monitored growing season carbon dioxide (CO₂) fluxes across a factorial design of in situ water table (control, drought, and flooded plots) and soil warming (control vs. warming via open top chambers) treatments for 2 years in a rich fen located just outside the Bonanza Creek Experimental Forest in interior Alaska. The drought (lowered water table position) treatment was a weak sink or small source of atmospheric CO₂ compared to the moderate atmospheric CO₂ sink at our control. This change in net ecosystem exchange was due to lower gross primary production and light-saturated photosynthesis rather than increased ecosystem respiration. The flooded (raised water table position) treatment was a greater CO₂ sink in 2006 due largely to increased early season gross primary production and higher light-saturated photosynthesis. Although flooding did not have substantial effects on rates of ecosystem respiration, this water table treatment had lower maximum respiration rates and a higher temperature sensitivity of ecosystem respiration than the control plot. Surface soil warming increased both ecosystem respiration and gross primary production by approximately 16% compared to control (ambient temperature) plots, with no net effect on net ecosystem exchange. Results from this rich fen manipulation suggest that fast responses to drought will include reduced ecosystem C storage driven by plant stress, whereas inundation will increase ecosystem C storage by stimulating plant growth.     

基于改进SW模型的千烟洲人工林蒸散组分拆分及其特征

蒸散组分拆分是准确评估陆地生态系统生产力以及估算水分利用效率的重要基础。利用改进后的Shuttleworth-Wallace模型,将蒸散拆分为植被蒸腾、土壤蒸发和冠层截留蒸发,并采用Monte Carlo随机参数化方案对模型参数进行优化。将模型与千烟洲亚热带人工针叶林站点的2011年涡度相关及小气候观测资料结合,对千烟洲人工林蒸散及其组分进行模拟。研究结果表明:半小时尺度上蒸散量模拟值与实测值的一致性在晴天和雨天都较高。半小时尺度上全年蒸散模拟值与实测值的决定系数、均方根误差和平均偏差为0.73、1.55 mmol m~(-2)s~(-1)和0.21 mmol m~(-2)s~(-1)。蒸散是该生态系统水分输出的最主要贡献项,占全年降水的80%。在蒸散中,植被蒸腾约占总蒸散量的85%,可推测2011年千烟洲人工林生态系统有较高的水分利用效率。该生态系统的蒸腾量季节变化明显,主要受饱和水汽压差和气温两种环境因素以及植被的叶面积指数影响且与三者均呈正相关;土壤蒸发约占总蒸散量的5%,季节变化平缓;模拟的冠层截留蒸发量约占总蒸散量的10%,季节变化大,与降水量呈正相关,与暴雨频次呈负相关,说明冠层无法有效截留强降水。该模型参数较少、时间分辨率高且可以有效模拟蒸散及其组分特征,是陆地生态系统水分循环过程研究有力的模型工具。     

Intraseasonal Variation in Water and Carbon Dioxide Flux Components in a Semiarid Riparian Woodland

Abstract We investigated how the distribution of precipitation over a growing season influences the coupling of carbon and water cycle components in a semiarid floodplain woodland dominated by the deep-rooted velvet mesquite (Prosopis velutina). Gross ecosystem production (GEP) and ecosystem respiration (R _eco) were frequently uncoupled because of their different sensitivities to growing season rainfall. Soon after the first monsoon rains, R _eco was high and was not proportional to slight increases in GEP. During the wettest month of the growing season (July), the system experienced a net carbon loss equivalent to 46% of the carbon accumulated over the 6-month study period (114 g C m⁻²; May–October). It appears that a large CO₂ efflux and a rapid water loss following precipitation early in the growing season and a later CO₂ gain is a defining characteristic of seasonally dry ecosystems. The relative contribution of plant transpiration (T) to total evapotranspiration (ET) (T/ET) was 0.90 for the entire growing season, with T/ET reaching a value of 1 during dry conditions and dropping to as low as 0.65 when the soil surface was wet. The evaporation fraction (E) was equivalent to 31% of the precipitation received during the study period (253 mm) whereas trees and understory vegetation transpired 38 and 31%, respectively, of this water source. The water-use efficiency of the vegetation (GEP/T) was higher later in the growing season when the C4 grassy understory was fully developed. The influence of rain on net ecosystem production (NEP) can be interpreted as the proportion of precipitation that is transpired by the plant community; the water-use efficiency of the vegetation and the precipitation fraction that is lost by evaporation.     

Productivity responses to altered rainfall patterns in a C<Subscript>4</Subscript>-dominated grassland

Rainfall variability is a key driver of ecosystem structure and function in grasslands worldwide. Changes in rainfall patterns predicted by global climate models for the central United States are expected to cause lower and increasingly variable soil water availability, which may impact net primary production and plant species composition in native Great Plains grasslands. We experimentally altered the timing and quantity of growing season rainfall inputs by lengthening inter-rainfall dry intervals by 50%, reducing rainfall quantities by 30%, or both, compared to the ambient rainfall regime in a native tallgrass prairie ecosystem in northeastern Kansas. Over three growing seasons, increased rainfall variability caused by altered rainfall timing with no change in total rainfall quantity led to lower and more variable soil water content (0–30 cm depth), a ~10% reduction in aboveground net primary productivity (ANPP), increased root to shoot ratios, and greater canopy photon flux density at 30 cm above the soil surface. Lower total ANPP primarily resulted from reduced growth, biomass and flowering of subdominant warm-season C₄ grasses while productivity of the dominant C₄ grass Andropogon gerardii was relatively unresponsive. In general, vegetation responses to increased soil water content variability were at least equal to those caused by imposing a 30% reduction in rainfall quantity without altering the timing of rainfall inputs. Reduced ANPP most likely resulted from direct effects of soil moisture deficits on root activity, plant water status, and photosynthesis. Altered rainfall regimes are likely to be an important element of climate change scenarios in this grassland, and the nature of interactions with other climate change elements remains a significant challenge for predicting ecosystem responses to climate change.     

Landscape and Ecosystem-Level Controls on Net Carbon Dioxide Exchange along a Natural Moisture Gradient in Canadian Low Arctic Tundra

Our understanding of the controls and magnitudes of regional CO₂ exchanges in the Arctic are limited by uncertainties due to spatial heterogeneity in vegetation across the landscape and temporal variation in environmental conditions through the seasons. We measured daytime net ecosystem CO₂ exchange and each of its component fluxes in the three major tundra ecosystem-types that typically occur along natural moisture gradients in the Canadian Low Arctic biweekly during the full snow-free season of 2004. In addition, we used a plant-removal treatment to compare the contribution of bulk soil organic matter to total respiratory CO₂ loss among these ecosystems. Net CO₂ exchange rates varied strongly, but not consistently, among ecosystems in the spring and summer phases as a result of ecosystem-specific and differing responses of gross photosynthesis and respiration to temporal variation in environmental conditions. Overall, net carbon gain was largest in the wet sedge ecosystem and smallest in the dry heath. Our measures of CO₂ flux variation within each ecosystem were frequently most closely correlated with air or soil temperatures during each seasonal phase. Nevertheless, a particularly large rainfall event in early August rapidly decreased respiration rates and stimulated gross photosynthetic rates, resulting in peak rates of net carbon gain in all ecosystems. Finally, the bulk soil carbon contribution to total respiration was relatively high in the birch hummock ecosystem. Together, these results demonstrate that the relative influences of moisture and temperature as primary controls on daytime net ecosystem CO₂ exchange and its component fluxes differ in fundamental ways between the landscape and ecosystem scales. Furthermore, they strongly suggest that carbon cycling responses to environmental change are likely to be highly ecosystem-specific, and thus to vary substantially across the low arctic landscape. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Leaf gas exchange and water status responses of a native and non-native grass to precipitation across contrasting soil surfaces in the Sonoran Desert

Arid and semi-arid ecosystems of the southwestern US are undergoing changes in vegetation composition and are predicted to experience shifts in climate. To understand implications of these current and predicted changes, we conducted a precipitation manipulation experiment on the Santa Rita Experimental Range in southeastern Arizona. The objectives of our study were to determine how soil surface and seasonal timing of rainfall events mediate the dynamics of leaf-level photosynthesis and plant water status of a native and non-native grass species in response to precipitation pulse events. We followed a simulated precipitation event (pulse) that occurred prior to the onset of the North American monsoon (in June) and at the peak of the monsoon (in August) for 2002 and 2003. We measured responses of pre-dawn water potential, photosynthetic rate, and stomatal conductance of native (Heteropogon contortus) and non-native (Eragrostis lehmanniana) C₄ bunchgrasses on sandy and clay-rich soil surfaces. Soil surface did not always amplify differences in plant response to a pulse event. A June pulse event lead to an increase in plant water status and photosynthesis. Whereas the August pulse did not lead to an increase in plant water status and photosynthesis, due to favorable soil moisture conditions facilitating high plant performance during this period. E. lehmanniana did not demonstrate heightened photosynthetic performance over the native species in response to pulses across both soil surfaces. Overall accumulated leaf-level CO₂ response to a pulse event was dependent on antecedent soil moisture during the August pulse event, but not during the June pulse event. This work highlights the need to understand how desert species respond to pulse events across contrasting soil surfaces in water-limited systems that are predicted to experience changes in climate.     

Wetting-induced soil CO2 emission pulses are driven by interactions among soil temperature,carbon, and nitrogen limitation in the Colorado Desert

Warming-induced changes in precipitation regimes, coupled with anthropogenically enhanced nitrogen (N) deposition, are likely to increase the prevalence, duration, and magnitude of soil respiration pulses following wetting via interactions among temperature and carbon (C) and N availability. Quantifying the importance of these interactive controls on soil respiration is a key challenge as pulses can be large terrestrial sources of atmospheric carbon dioxide (CO₂) over comparatively short timescales. Using an automated sensor system, we measured soil CO₂ flux dynamics in the Colorado Desert—a system characterized by pronounced transitions from dry-to-wet soil conditions—through a multi-year series of experimental wetting campaigns. Experimental manipulations included combinations of C and N additions across a range of ambient temperatures and across five sites varying in atmospheric N deposition. We found soil CO₂ pulses following wetting were highly predictable from peak instantaneous CO₂ flux measurements. CO₂ pulses consistently increased with temperature, and temperature at time of wetting positively correlated to CO₂ pulse magnitude. Experimentally adding N along the N deposition gradient generated contrasting pulse responses: adding N increased CO₂ pulses in low N deposition sites, whereas adding N decreased CO₂ pulses in high N deposition sites. At a low N deposition site, simultaneous additions of C and N during wetting led to the highest observed soil CO₂ fluxes reported globally at 299.5 μmol CO₂ m⁻² s⁻¹. Our results suggest that soils have the capacity to emit high amounts of CO₂ within small timeframes following infrequent wetting, and pulse sizes reflect a non-linear combination of soil resource and temperature interactions. Importantly, the largest soil CO₂ emissions occurred when multiple resources were amended simultaneously in historically resource-limited desert soils, pointing to regions experiencing simultaneous effects of desertification and urbanization as key locations in future global C balance.     

A meta-analysis of plant physiological and growth responses to temperature and elevated CO<Subscript>2</Subscript>

Atmospheric carbon dioxide (CO₂) and global mean temperature are expected to be significantly higher by the end of the 21st century. Elevated CO₂ (eCO₂) and higher temperature each affect plant physiology and growth, but their interactive effects have not been reviewed statistically with respect to higher chronic mean temperatures and abrupt heat stress. In this meta-analysis, we examined the effect of CO₂ on the physiology and growth of plants subjected to different temperature treatments. The CO₂ treatments were categorized into ambient (<400 ppm) or elevated (>560 ppm) levels, while temperature treatments were categorized into ambient temperature (AT), elevated temperature (ET; AT + 1.4–6°C), or heat stress (HS; AT + >8°C). Plant species were grouped according to photosynthetic pathways (C₃, C₄), functional types (legumes, non-legumes), growth forms (herbaceous, woody), and economic purposes (crop, non-crop). eCO₂ enhanced net photosynthesis at AT, ET, and HS in C₃ species (especially at the HS level), but in C₄ species, it had no effect at AT, a positive effect at ET, and a negative effect at HS. The positive effect of eCO₂ on net photosynthesis was greater for legumes than for non-legumes at HS, for non-crops than crops at ET, and for woody than herbaceous species at ET and HS. Total (W _T) and above- (W _AG) and below-ground (W _BG) biomass were increased by eCO₂ for most species groups at all temperatures, except for C₄ species and W _BG of legumes at HS. Hence, eCO₂ × heat effects on growth were often not explained by effects on net photosynthesis. Overall, the results show that eCO₂ effects on plant physiology and growth vary under different temperature regimes, among functional groups and photosynthetic pathways, and among response variables. These findings have important implications for biomass accumulation and ecosystem functioning in the future when the CO₂ level is higher and climate extremes, such as heat waves, become more frequent.     

Wetting and drying cycles drive variations in the stable carbon isotope ratio of respired carbon dioxide in semi-arid grassland

In semi-arid regions, where plants using both C₃ and C₄ photosynthetic pathways are common, the stable C isotope ratio (δ¹³C) of ecosystem respiration (δ¹³C_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 CO₂. We hypothesized that timing of precipitation events and antecedent moisture interact with activity of C₃ and C₄ grasses to determine net ecosystem CO₂ exchange (NEE) and δ¹³C_R. Field measurements included CO₂ and δ¹³C fluxes from the whole ecosystem and from patches of different plant communities, biomass and δ¹³C 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 δ¹³C_R was higher in 2000 (−16‰) than 2001 (20‰), 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, δ¹³C averaged −22‰ from C₃ patches, −16‰ from C₄ patches, and −19‰ from mixed C₃ and C₄ patches. However, during dry conditions the apparent spatial differences were not obvious, suggesting reduced autotrophic activity in C₄ grasses with shallow rooting depth, soon after the onset of dry conditions. Air and soil temperatures were negatively correlated with δ¹³C_R; vapor pressure deficit was a poor predictor of δ¹³C_R, in contrast to more mesic ecosystems. Responses of respiration components to precipitation pulses were explained by differences in soil moisture thresholds between C₃ and C₄ 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.     

Environmental effects on CO2 efflux from riparian tundra in the northern foothills of the Brooks Range,Alaska, USA

Summary Carbon dioxide efflux and soil microenvironmental factors were measured diurnally in Carex aquatilus-and Eriophorum angustifolium-dominated riparian tundra communities to determine the relative importance of soil environmental factors controlling ecosystem carbon dioxide exchange with the atmosphere. Measurements were made weekly between 18 June and 24 July 1990. Diurnal patterns in carbon dioxide efflux were best explained by changes in soil temperature, while seasonal changes in efflux were correlated with changes in depth to water table, depth to frozen soil and soil moisture. Carbon dioxide efflux rates were lowest early in the growing season when high water tables and low soil temperatures limited microbial and root activity. Individual rainfall events that raised the water table were found to strongly reduce carbon dioxide efflux. As the growing season progressed, rainfall was low and depth to water table and soil temperatures increased. In response, carbon dioxide efflux increased strongly, attaining rates late in the season of approximately 10 g CO₂ m^–2 day^–1. These rates are as high as maxima recorded for other arctic sites. A mathematical model is developed which demonstrates that soil temperature and depth to water table may be used as efficient predictors of ecosystem CO₂ efflux in this habitat. In parallel with the field measurements of CO₂ efflux, microbial respiration was studied in the laboratory as a function of temperature and water content. Estimates of microbial respiration per square meter under field conditions were made by adjusting for potential respiring soil volume as water table changed and using measured soil temperatures. The results indicate that the effect of these factors on microbial respiration may explain a large part of the diurnal and seasonal variation observed in CO₂ efflux. As in coastal tundra sites, environmental changes that alter water table depth in riparian tundra communities will have large effects on ecosystem CO₂ efflux and carbon balance.     

Spatial heterogeneity in ecosystem structure and productivity in a moist Kenyan savanna

Topographic variations and disturbances are key sources of spatial heterogeneity in the ecosystem and may influence its functioning, productivity, and carbon (C) storage. In water controlled ecosystems, structural and functional heterogeneity become distinct during drought when the ecosystem processes are operating at their limits. We examined spatial heterogeneity arising from grazing, abandoned cropland, presence of Acacia trees, and termite mounds (termitaria). Soil water content (SWC) was significantly (P < 0.05) higher in termitaria and fenced (un-grazed) plots. Higher soil nitrogen (N) content occurred in the Acacia, termitaria, and fenced plots while total biomass was highest in the fenced plot. The termitaria plots showed the highest net ecosystem CO₂ exchange (NEE), ecosystem respiration (R _eco), and gross primary production (GPP) and were the only plots that were net CO₂ sinks. Except in fenced plots, maximum GPP was positively correlated with SWC and green biomass in all the other plots. Green biomass and R _eco were positively correlated with SWC. Shifting cultivation (abandoned farmlands) negatively affected soil quality, ecosystem CO₂ assimilation, and productivity. Removal of grazing (cattle) from the ecosystem negatively influenced GPP, while the presence of termitaria and Acacia trees facilitated soil water and N availability and ecosystem productivity. We concluded that soil water availability was responsible for most of the localized differences in the savanna and has a strong influence on ecosystem C capture and storage. We recommend that future studies on savanna productivity and ecosystem CO₂ fluxes should consider heterogeneity in the ecosystem in order to avoid bias and increase the accuracy of any estimates made.     

The Influence of Spatial Patterns of Soil Moisture on the Grass and Shrub Responses to a Summer Rainstorm in a Chihuahuan Desert Ecotone

The cycling of surface water, energy, nutrients, and carbon is different between semiarid grassland and shrubland ecosystems. Although differences are evident when grasslands are compared to shrublands, the processes that contribute to this transition are more challenging to document. We evaluate how surface redistribution of precipitation and plant responses to the resulting infiltration patterns could contribute to the changes that occur during the transition from grassland to shrubland. We measured soil water potential under grasses (Bouteloua eriopoda), shrubs (Larrea tridentata) and bare soil and changes in plant water relations and gas exchange following a 15 mm summer storm in the grassland–shrubland ecotone at the Sevilleta National Wildlife Refuge in central New Mexico USA. Following the storm, soil water potential (Ψ_s) increased to 30 cm depth beneath both grass and shrub canopies, with the greatest change observed in the top 15 cm of the soil. The increase in Ψ_s was greater beneath grass canopies than beneath shrub canopies. Ψ_s under bare soil increased only to 5 cm depth. The substantial redistribution of rainfall and different rooting depths of the vegetation resulted in high Ψ_s throughout most of the rooting volume of the grasses whereas soil moisture was unchanged throughout a large portion of the shrub rooting volume. Consistent with this pattern, predawn water potential (Ψ_PD) of grasses increased more than 5 MPa to greater than −1 MPa whereas Ψ_PD of shrubs increased to −2.5 MPa, a change of less than 2 MPa. Transpiration increased roughly linearly with Ψ_PD in both grasses and shrubs. In grasses, assimilation was strongly correlated with Ψ_PD whereas there was no relationship in shrubs where assimilation showed no significant response to the pulse of soil moisture following the storm. These data show that preferential redistribution of water to grass canopies enhances transpiration and assimilation by grasses following large summer storms. This process may inhibit shrubland expansion at the ecotone during periods without extreme drought.