Direct and Legacy Effects of Varying Cool-Season Precipitation Totals on Ecosystem Carbon Flux in a Semi-Arid Mixed Grassland.

In the semi-arid grasslands of the southwest United States, annual precipitation is divided between warm-season (July-September) convective precipitation and cool-season (December-March) frontal storms. While evidence suggests shifts in precipitation seasonal distribution, there is a poor understanding of the ecosystem carbon flux responses to cool-season precipitation and the potential legacy effects on subsequent warm-season carbon fluxes. Results from a two-year experiment with three cool-season precipitation treatments (dry, received 5th percentile cool-season total precipitation; normal, 50th; wet, 95th) and constant warm-season precipitation illustrate the direct and legacy effects on carbon fluxes, but in opposing ways. In wet cool-season plots, gross primary productivity (GPP) and ecosystem respiration (ER) were 103% and 127% higher than in normal cool-season plots. In dry cool-season plots, GPP and ER were 47% and 85% lower compared to normal cool-season plots. Unexpectedly, we found a positive legacy effect of the dry cool-season treatment on warm-season carbon flux, resulting in a significant increase in both GPP and ER in the subsequent warm season, compared to normal cool-season plots. Our results reveal positive legacy effects of cool-season drought on warm-season carbon fluxes and highlight the importance of the relatively under-studied cool-growing season and its direct/indirect impact on the ecosystem carbon budget.

Thermography captures the differential sensitivity of dryland functional types to changes in rainfall event timing and magnitude.

Drylands of the southwestern United States are rapidly warming, and rainfall is becoming less frequent and more intense, with major yet poorly understood implications for ecosystem structure and function. Thermography-based estimates of plant temperature can be integrated with air temperature to infer changes in plant physiology and response to climate change. However, very few studies have evaluated plant temperature dynamics at high spatiotemporal resolution in rainfall pulse-driven dryland ecosystems. We address this gap by incorporating high-frequency thermal imaging into a field-based precipitation manipulation experiment in a semi-arid grassland to investigate the impacts of rainfall temporal repackaging. All other factors held constant, we found that fewer/larger precipitation events led to cooler plant temperatures (1.4°C) compared to that of many/smaller precipitation events. Perennials, in particular, were 2.5°C cooler than annuals under the fewest/largest treatment. We show these patterns were driven by: increased and consistent soil moisture availability in the deeper soil layers in the fewest/largest treatment; and deeper roots of perennials providing access to deeper plant available water. Our findings highlight the potential for high spatiotemporal resolution thermography to quantify the differential sensitivity of plant functional groups to soil water availability. Detecting these sensitivities is vital to understanding the ecohydrological implications of hydroclimate change.

Exceptional heat and atmospheric dryness amplified losses of primary production during the 2020 U.S. Southwest hot drought.

Earth's ecosystems are increasingly threatened by "hot drought," which occurs when hot air temperatures coincide with precipitation deficits, intensifying the hydrological, physiological, and ecological effects of drought by enhancing evaporative losses of soil moisture (SM) and increasing plant stress due to higher vapor pressure deficit (VPD). Drought-induced reductions in gross primary production (GPP) exert a major influence on the terrestrial carbon sink, but the extent to which hotter and atmospherically drier conditions will amplify the effects of precipitation deficits on Earth's carbon cycle remains largely unknown. During summer and autumn 2020, the U.S. Southwest experienced one of the most intense hot droughts on record, with record-low precipitation and record-high air temperature and VPD across the region. Here, we use this natural experiment to evaluate the effects of hot drought on GPP and further decompose those negative GPP anomalies into their constituent meteorological and hydrological drivers. We found a 122 Tg C (>25%) reduction in GPP below the 2015-2019 mean, by far the lowest regional GPP over the Soil Moisture Active Passive satellite record. Roughly half of the estimated GPP loss was attributable to low SM (likely a combination of record-low precipitation and warming-enhanced evaporative depletion), but record-breaking VPD amplified the reduction of GPP, contributing roughly 40% of the GPP anomaly. Both air temperature and VPD are very likely to continue increasing over the next century, likely leading to more frequent and intense hot droughts and substantially enhancing drought-induced GPP reductions.

Drought and heat wave impacts on grassland carbon cycling across hierarchical levels.

Droughts and heat waves are increasing in magnitude and frequency, altering the carbon cycle. However, understanding of the underlying response mechanisms remains poor, especially for the combination (hot drought). We conducted a 4-year field experiment to examine both individual and interactive effects of drought and heat wave on carbon cycling of a semiarid grassland across individual, functional group, community and ecosystem levels. Drought did not change below-ground biomass (BGB) or above-ground biomass (AGB) due to compensation effects between grass and non-grass functional groups. However, consistently decreased BGB under heat waves limited such compensation effects, resulting in reduced AGB. Ecosystem CO2 fluxes were suppressed by droughts, attributed to stomatal closure-induced reductions in leaf photosynthesis and decreased AGB of grasses, while CO2 fluxes were little affected by heat waves. Overall the hot drought produced the lowest leaf photosynthesis, AGB and ecosystem CO2 fluxes although the interactions between heat wave and drought were usually not significant. Our results highlight that the functional group compensatory effects that maintain community-level AGB rely on feedback of root system responses, and that plant adjustments at the individual level, together with shifts in composition at the functional group level, co-regulate ecosystem carbon sink strength under climate extremes.

Montane forest productivity across a semiarid climatic gradient.

High-elevation montane forests are disproportionately important to carbon sequestration in semiarid climates where low elevations are dry and characterized by low carbon density ecosystems. However, these ecosystems are increasingly threatened by climate change with seasonal implications for photosynthesis and forest growth. As a result, we leveraged eddy covariance data from six evergreen conifer forest sites in the semiarid western United States to extrapolate the status of carbon sequestration within a framework of projected warming and drying. At colder locations, the seasonal evolution of gross primary productivity (GPP) was characterized by a single broad maximum during the summer that corresponded to snow melt-derived moisture and a transition from winter dormancy to spring activity. Conversely, winter dormancy was transient at warmer locations, and GPP was responsive to both winter and summer precipitation such that two distinct GPP maxima were separated by a period of foresummer drought. This resulted in a predictable sequence of primary limiting factors to GPP beginning with air temperature in winter and proceeding to moisture and leaf area during the summer. Due to counteracting winter (positive) and summer (negative) GPP responses to warming, leaf area index and moisture availability were the best predictors of annual GPP differences across sites. Overall, mean annual GPP was greatest at the warmest site due to persistent vegetation photosynthetic activity throughout the winter. These results indicate that the trajectory of this region's carbon sequestration will be sensitive to reduced or delayed summer precipitation, especially if coupled to snow drought and earlier soil moisture recession, but summer precipitation changes remain highly uncertain. Given the demonstrated potential for seasonally offsetting responses to warming, we project that decadal semiarid montane forest carbon sequestration will remain relatively stable in the absence of severe disturbance.

Terrestrial N2 O emissions and related functional genes under climate change: A global meta-analysis.

Nitrous oxide (N2 O) emissions from soil contribute to global warming and are in turn substantially affected by climate change. However, climate change impacts on N2 O production across terrestrial ecosystems remain poorly understood. Here, we synthesized 46 published studies of N2 O fluxes and relevant soil functional genes (SFGs, that is, archaeal amoA, bacterial amoA, nosZ, narG, nirK and nirS) to assess their responses to increased temperature, increased or decreased precipitation amounts, and prolonged drought (no change in total precipitation but increase in precipitation intervals) in terrestrial ecosystem (i.e. grasslands, forests, shrublands, tundra and croplands). Across the data set, temperature increased N2 O emissions by 33%. However, the effects were highly variable across biomes, with strongest temperature responses in shrublands, variable responses in forests and negative responses in tundra. The warming methods employed also influenced the effects of temperature on N2 O emissions (most effectively induced by open-top chambers). Whole-day or whole-year warming treatment significantly enhanced N2 O emissions, but daytime, nighttime or short-season warming did not have significant effects. Regardless of biome, treatment method and season, increased precipitation promoted N2 O emission by an average of 55%, while decreased precipitation suppressed N2 O emission by 31%, predominantly driven by changes in soil moisture. The effect size of precipitation changes on nirS and nosZ showed a U-shape relationship with soil moisture; further insight into biotic mechanisms underlying N2 O emission response to climate change remain limited by data availability, underlying a need for studies that report SFG. Our findings indicate that climate change substantially affects N2 O emission and highlights the urgent need to incorporate this strong feedback into most climate models for convincing projection of future climate change.

Ecological responses to heavy rainfall depend on seasonal timing and multi-year recurrence.

Heavy rainfall events are expected to increase in frequency and severity in the future. However, their effects on natural ecosystems are largely unknown, in particular with different seasonal timing of the events and recurrence over multiple years. We conducted a 4 yr manipulative experiment to explore grassland response to heavy rainfall imposed in either the middle of, or late in, the growing season in Inner Mongolia, China. We measured hierarchical responses at individual, community and ecosystem levels. Surprisingly, above-ground biomass remained stable in the face of heavy rainfall, regardless of seasonal timing, whereas heavy rainfall late in the growing season had consistent negative impacts on below-ground and total biomass. However, such negative biomass effects were not significant for heavy rainfall in the middle of the growing season. By contrast, heavy rainfall in the middle of the growing season had greater positive effects on ecosystem CO2 exchanges, mainly reflected in the latter 2 yr of the 4 yr experiment. This two-stage response of CO2 fluxes was regulated by increased community-level leaf area and leaf-level photosynthesis and interannual variability of natural precipitation. Overall, our study demonstrates that ecosystem impacts of heavy rainfall events crucially depend on the seasonal timing and multiannual recurrence. Plant physiological and morphological adjustment appeared to improve the capacity of the ecosystem to respond positively to heavy rainfall.

CO2 exchange and evapotranspiration across dryland ecosystems of southwestern North America.

Global-scale studies suggest that dryland ecosystems dominate an increasing trend in the magnitude and interannual variability of the land CO2 sink. However, such analyses are poorly constrained by measured CO2 exchange in drylands. Here we address this observation gap with eddy covariance data from 25 sites in the water-limited Southwest region of North America with observed ranges in annual precipitation of 100-1000 mm, annual temperatures of 2-25°C, and records of 3-10 years (150 site-years in total). Annual fluxes were integrated using site-specific ecohydrologic years to group precipitation with resulting ecosystem exchanges. We found a wide range of carbon sink/source function, with mean annual net ecosystem production (NEP) varying from -350 to +330 gCm-2 across sites with diverse vegetation types, contrasting with the more constant sink typically measured in mesic ecosystems. In this region, only forest-dominated sites were consistent carbon sinks. Interannual variability of NEP, gross ecosystem production (GEP), and ecosystem respiration (Reco ) was larger than for mesic regions, and half the sites switched between functioning as C sinks/C sources in wet/dry years. The sites demonstrated coherent responses of GEP and NEP to anomalies in annual evapotranspiration (ET), used here as a proxy for annually available water after hydrologic losses. Notably, GEP and Reco were negatively related to temperature, both interannually within site and spatially across sites, in contrast to positive temperature effects commonly reported for mesic ecosystems. Models based on MODIS satellite observations matched the cross-site spatial pattern in mean annual GEP but consistently underestimated mean annual ET by ~50%. Importantly, the MODIS-based models captured only 20-30% of interannual variation magnitude. These results suggest the contribution of this dryland region to variability of regional to global CO2 exchange may be up to 3-5 times larger than current estimates.

Terrestrial carbon balance in a drier world: the effects of water availability in southwestern North America.

Global modeling efforts indicate semiarid regions dominate the increasing trend and interannual variation of net CO2 exchange with the atmosphere, mainly driven by water availability. Many semiarid regions are expected to undergo climatic drying, but the impacts on net CO2 exchange are poorly understood due to limited semiarid flux observations. Here we evaluated 121 site-years of annual eddy covariance measurements of net and gross CO2 exchange (photosynthesis and respiration), precipitation, and evapotranspiration (ET) in 21 semiarid North American ecosystems with an observed range of 100 - 1000 mm in annual precipitation and records of 4-9 years each. In addition to evaluating spatial relationships among CO2 and water fluxes across sites, we separately quantified site-level temporal relationships, representing sensitivity to interannual variation. Across the climatic and ecological gradient, photosynthesis showed a saturating spatial relationship to precipitation, whereas the photosynthesis-ET relationship was linear, suggesting ET was a better proxy for water available to drive CO2 exchanges after hydrologic losses. Both photosynthesis and respiration showed similar site-level sensitivity to interannual changes in ET among the 21 ecosystems. Furthermore, these temporal relationships were not different from the spatial relationships of long-term mean CO2 exchanges with climatic ET. Consequently, a hypothetical 100-mm change in ET, whether short term or long term, was predicted to alter net ecosystem production (NEP) by 64 gCm(-2) yr(-1). Most of the unexplained NEP variability was related to persistent, site-specific function, suggesting prioritization of research on slow-changing controls. Common temporal and spatial sensitivity to water availability increases our confidence that site-level responses to interannual weather can be extrapolated for prediction of CO2 exchanges over decadal and longer timescales relevant to societal response to climate change.

Extreme drought alters methane uptake but not methane sink in semi-arid steppes of Inner Mongolia.

Global climate change, particularly drought, is expected to alter grassland methane (CH4) oxidation, a key natural process against atmospheric greenhouse gas accumulation, yet the extent of this effect and its interaction with future atmospheric CH4 concentrations increases remains uncertain. To address this research gap, we measured CH4 flux during an imposed three-month rain-free period corresponding to a 100-year recurrence drought in soil mesocosms collected from 16 different Eurasian steppe sites. We also investigated the abundance and composition of methanotrophs. Additionally, we conducted a laboratory experiment to explore the impact of elevated CH4 concentration on the CH4 uptake capacity of grassland soil under drought conditions. We found that regardless of the type of grassland, CH4 flux was still being absorbed at its peak, meaning that all grasslands functioned as persistent CH4 sinks even when the soil water content (SWC) was <5 %. A bell-shaped relationship between SWC and CH4 uptake was observed in the soils. The average maximum CH4 oxidation rate in the meadow steppe was higher than that in the typical and desert steppe soils during extreme drought. The experimental elevation of atmospheric CH4 concentration counteracted the anticipated reduction in CH4 uptake related to physiological water stress on methanotrophic soil microbes under the drought stress. On the contrary, we found that across the regional scale, nitrogen, phosphorous, and total soil organic content played a crucial role in moderating the duration and magnitude of CH4 uptake with respect to SWC. USC-γ (Upland Soil Cluster γ) and JR-3 (Jasper Ridge Cluster) were the dominant group of soil methanotrophic bacteria in three types of grassland. However, the methanotrophic abundance, rather than the methanotrophic community composition, was the dominant microbiological factor governing CH4 uptake during the drought.

Bark beetle impacts on forest evapotranspiration and its partitioning.

Insect outbreaks affect forest structure and function and represent a major category of forest disturbance globally. However, the resulting impacts on evapotranspiration (ET), and especially hydrological partitioning between the abiotic (evaporation) and biotic (transpiration) components of total ET, are not well constrained. As a result, we combined remote sensing, eddy covariance, and hydrological modeling approaches to determine the effects of bark beetle outbreak on ET and its partitioning at multiple scales throughout the Southern Rocky Mountain Ecoregion (SRME), USA. At the eddy covariance measurement scale, 85 % of the forest was affected by beetles, and water year ET as a fraction of precipitation (P) decreased by 30 % relative to a control site, with 31 % greater reductions in growing season transpiration relative to total ET. At the ecoregion scale, satellite remote sensing masked to areas of >80 % tree mortality showed corresponding ET/P reductions of 9-15 % that occurred 6-8 years post-disturbance, and indicated that the majority of the total reduction occurred during the growing season; the Variable Infiltration Capacity hydrological model showed an associated 9-18 % increase in the ecoregion runoff ratio. Long-term (16-18 year) ET and vegetation mortality datasets extend the length of previously published analyses and allowed for clear characterization of the forest recovery period. During that time, transpiration recovery outpaced total ET recovery, which was lagged in part due to persistently reduced winter sublimation, and there was associated evidence of increasing late summer vegetation moisture stress. Overall, comparison of three independent methods and two partitioning approaches demonstrated a net negative impact of bark beetles on ET, and a relatively greater negative impact on transpiration, following bark beetle outbreak in the SRME.

Indicators of water use efficiency across diverse agroecosystems and spatiotemporal scales.

Understanding the relationship between water and production within and across agroecosystems is essential for addressing several agricultural challenges of the 21st century: providing food, fuel, and fiber to a growing human population, reducing the environmental impacts of agricultural production, and adapting food systems to climate change. Of all human activities, agriculture has the highest demand for water globally. Therefore, increasing water use efficiency (WUE), or producing 'more crop per drop', has been a long-term goal of agricultural management, engineering, and crop breeding. WUE is a widely used term applied across a diverse array of spatial scales, spanning from the leaf to the globe, and over temporal scales ranging from seconds to months to years. The measurement, interpretation, and complexity of WUE varies enormously across these spatial and temporal scales, challenging comparisons within and across diverse agroecosystems. The goals of this review are to evaluate common indicators of WUE in agricultural production and assess tradeoffs when applying these indicators within and across agroecosystems amidst a changing climate. We examine three questions: (1) what are the uses and limitations of common WUE indicators, (2) how can WUE indicators be applied within and across agroecosystems, and (3) how can WUE indicators help adapt agriculture to climate change? Addressing these agricultural challenges will require land managers, producers, policy makers, researchers, and consumers to evaluate costs and benefits of practices and innovations of water use in agricultural production. Clearly defining and interpreting WUE in the most scale-appropriate way is crucial for advancing agroecosystem sustainability.

Modified enzyme activity assay to determine biofilm biomass.

An assay of potential exoproteolytic enzyme activity was modified to quantitatively measure the biomass of attached biofilm. The assay utilized the nonfluorescent compound L-leucine-beta-naphthylamide (LLbetaN) that becomes fluorescent when bacterial exoenzymes break the peptide bond, releasing the fluorochrome beta-naphthylamine. Fluorescence development was measured by pumping the liquid phase of a biofilm sample through a fluorescence detector and recording the detector output using a personal computer. A significant linear relationship was shown to exist between the rate of fluorescence development and the biofilm's biomass as carbon, determined using total direct cell counts, measured cell volumes and an existing relationship between cell volume and cell carbon. The technique was used to measure biofilm biomass for experiments where iron oxides were the substratum. Biofilm biomass measurements made using heterotrophic plate counts (HPCs) on R2A medium were shown to correlate well to biomass measurements made using the modified enzyme assay. The technique was shown to be sufficiently sensitive to measure biomass on samples containing little biofilm biomass, such as those exposed to free chlorine. While granular and porous media were used for the experiments presented, small biofilm coupons could easily be used to measure biofilm biomass, expanding the number of possible applications for the enzyme assay technique.

Minimizing biofilm in the presence of iron oxides and humic substances.

Based upon circumstantial evidence linking elevated coliform bacteria counts in drinking water distribution systems with unlined cast iron pipe, it was hypothesized that adsorption of humic substances by iron oxide containing corrosion products (CPs) can stimulate and/or support biofilm development. Using porous media consisting of iron-oxide-coated glass beads (IOCBs) or actual iron CPs, experiments were performed to evaluate the effectiveness of different corrosion control and disinfection treatments in reducing biofilm when humic substances were the carbon source. Free chlorine was the most effective treatment in minimizing biofilm. Addition of phosphate alone did not significantly reduce biofilm using the CPs, but there was weak evidence it did using the IOCBs. The combination of free chlorine and phosphate was more effective at minimizing biofilm than free chlorine alone when CPs were the media. The presence of humic substances was a major factor when considering biofilm minimization based on results of experiments using both types of iron oxide media. The combination of humic substances and CPs led to an increase in biofilm biomass when free chlorine was not present, similar to conditions that could occur at distribution system dead-ends. Treatment to raise the pH to 9 did not reduce biofilm in experiments using both media, and actually increased biofilm in the experiment using CPs under the conditions tested.

Temperature and wetland plant species effects on wastewater treatment and root zone oxidation.

Constructed wetlands are widely used for wastewater treatment, but there is little information on processes affecting their performance in cold climates, effects of plants on seasonal performance, or plant selection for cold regions. We evaluated the effects of three plant species on seasonal removal of dissolved organic matter (OM) (measured by chemical oxygen demand and dissolved organic carbon) and root zone oxidation status (measured by redox potential [Eh] and sulfate [SO4(2-)]) in subsurface-flow wetland (SSW) microcosms. A series of 20-d incubations of simulated wastewater was conducted during a 28-mo greenhouse study at temperatures from 4 to 24 degrees C. Presence and species of plants strongly affected seasonal differences in OM removal and root zone oxidation. All plants enhanced OM removal compared with unplanted controls, but plant effects and differences among species were much greater at 4 degrees C, during dormancy, than at 24 degrees C, during the growing season. Low temperatures were associated with decreased OM removal in unplanted controls and broadleaf cattail (Typha latifolia L.) microcosms and with increased removal in beaked sedge (Carex rostrata Stokes) and hardstem bulrush [Schoenoplectus acutus (Muhl. ex Bigelow) A. & D. Löve var. acutus] microcosms. Differences in OM removal corresponded to species' apparent abilities to increase root zone oxygen supply. Sedge and bulrush significantly raised Eh values and SO4(2-) concentrations, particularly at 4 degrees C. These results add to evidence that SSWs can be effective in cold climates and suggest that plant species selection may be especially important to optimizing SSW performance in cold climates.