Lack of eutrophication in a tallgrass prairie ecosystem over 27 years

Many North American grasslands are receiving atmospheric nitrogen (N) deposition at rates above what are considered critical eutrophication thresholds. Yet, potential changes in grassland function due to anthropogenic N deposition are poorly resolved, especially considering that other dynamic factors such as land use and precipitation can also affect N availability. To better understand whether elevated N deposition has altered ecosystem structure or function in North American grasslands, we analyzed a 27-year record of ecophysiological, community, and ecosystem metrics for an annually burned Kansas tallgrass prairie. Over this time, despite increasing rates of N deposition that are within the range of critical loads for grasslands, there was no evidence of eutrophication. Plant N concentrations did not increase, soil moisture did not decline, forb diversity did not decline, and the relative abundance of dominant grasses did not shift toward more eutrophic species. Neither aboveground primary productivity nor N availability to plants increased. The fates of deposited N in grasslands are still uncertain, and could include management losses through burning and grazing. However, evidence from this grassland indicates that eutrophication of North American grassland ecosystems is not an inevitable consequence of current levels of N deposition

Evidence of physiological decoupling from grassland ecosystem drivers by an encroaching woody shrub

Shrub encroachment of grasslands is a transformative ecological process by which native woody species increase in cover and frequency and replace the herbaceous community. Mechanisms of encroachment are typically assessed using temporal data or experimental manipulations, with few large spatial assessments of shrub physiology. In a mesic grassland in North America, we measured inter- and intra-annual variability in leaf δ¹³C in Cornus drummondii across a grassland landscape with varying fire frequency, presence of large grazers and topographic variability. This assessment of changes in individual shrub physiology is the largest spatial and temporal assessment recorded to date. Despite a doubling of annual rainfall (in 2008 versus 2011), leaf δ¹³C was statistically similar among and within years from 2008-11 (range of −28 to −27‰). A topography*grazing interaction was present, with higher leaf δ¹³C in locations that typically have more bare soil and higher sensible heat in the growing season (upland topographic positions and grazed grasslands). Leaf δ¹³C from slopes varied among grazing contrasts, with upland and slope leaf δ¹³C more similar in ungrazed locations, while slopes and lowlands were more similar in grazed locations. In 2011, canopy greenness (normalized difference vegetation index – NDVI) was assessed at the centroid of individual shrubs using high-resolution hyperspectral imagery. Canopy greenness was highest mid-summer, likely reflecting temporal periods when C assimilation rates were highest. Similar to patterns seen in leaf δ¹³C, NDVI was highest in locations that typically experience lowest sensible heat (lowlands and ungrazed). The ability of Cornus drummondii to decouple leaf physiological responses from climate variability and fire frequency is a likely contributor to the increase in cover and frequency of this shrub species in mesic grassland and may be generalizable to other grasslands undergoing woody encroachment

Extending the Osmometer Method for Assessing Drought Tolerance in Herbaceous Species

Community-scale surveys of plant drought tolerance are essential for understanding semi-arid ecosystems and community responses to climate change. Thus, there is a need for an accurate and rapid methodology for assessing drought tolerance strategies across plant functional types. The osmometer method for predicting leaf osmotic potential at full turgor ((o)), a key metric of leaf-level drought tolerance, has resulted in a 50-fold increase in the measurement speed of this trait; however, the applicability of this method has only been tested in woody species and crops. Here, we assess the osmometer method for use in herbaceous grassland species and test whether (o) is an appropriate plant trait for understanding drought strategies of herbaceous species as well as species distributions along climate gradients. Our model for predicting leaf turgor loss point ((TLP)) from (o) ((TLP)=0.80(o)-0.845) is nearly identical to the model previously presented for woody species. Additionally, (o) was highly correlated with (TLP) for graminoid species ((tlp)=0.944(o)-0.611; r(2)=0.96), a plant functional group previously flagged for having the potential to cause erroneous measurements when using an osmometer. We report that (o), measured with an osmometer, is well correlated with other traits linked to drought tolerance (namely, leaf dry matter content and leaf vulnerability to hydraulic failure) as well as climate extremes linked to water availability. The validation of the osmometer method in an herb-dominated ecosystem paves the way for rapid community-scale surveys of drought tolerance across plant functional groups, which could improve trait-based predictions of ecosystem responses to climate change

Trading water for carbon in the future : effects of elevated CO2 and warming on leaf hydraulic traits in a semiarid grassland

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)

Water Availability Dictates How Plant Traits Predict Demographic Rates

A major goal in ecology is to make generalizable predictions of organism responses to environmental variation based on their traits. However, straightforward relationships between traits and fitness are rare and likely to vary with environmental context. Characterizing how traits mediate demographic responses to the environment may enhance the predictions of organism responses to global change. We synthesized 15 years of demographic data and species-level traits in a shortgrass steppe to determine whether the effects of leaf and root traits on growth and survival depended on seasonal water availability. We predicted that (1) species with drought-tolerant traits, such as lower leaf turgor loss point (TLP) and higher leaf and root dry matter content (LDMC and RDMC), would be more likely to survive and grow in drier years due to higher wilting resistance, (2) these traits would not predict fitness in wetter years, and (3) traits that more directly measure physiological mechanisms of water use such as TLP would best predict demographic responses. We found that graminoids with more negative TLP and higher LDMC and RDMC had higher survival rates in drier years. Forbs demonstrated similar yet more variable responses. Graminoids grew larger in wetter years, regardless of traits. However, in both wet and dry years, graminoids with more negative TLP and higher LDMC and RDMC grew larger than less negative TLP and low LDMC and RDMC species. Traits significantly mediated the impact of drought on survival, but not growth, suggesting that survival could be a stronger driver of species\u27 drought response in this system. TLP predicted survival in drier years, but easier to measure LDMC and RDMC were equal or better predictors. These results advance our understanding of the mechanisms by which drought drives population dynamics, and show that abiotic context determines how traits drive fitness

Shifts in Plant Functional Composition Following Long-term Drought in Grasslands

1. Plant traits can provide unique insights into plant performance at the community scale. Functional composition, defined by both functional diversity and community-weighted trait means (CWMs), can affect the stability of above-ground net primary production (ANPP) in response to climate extremes. Further complexity arises, however, when functional composition itself responds to environmental change. The duration of climate extremes, such as drought, is expected to increase with rising global temperatures; thus, understanding the impacts of long-term drought on functional composition and the corresponding effect that has on ecosystem function could improve predictions of ecosystem sensitivity to climate change. 2. We experimentally reduced growing season precipitation by 66% across six temperate grasslands for 4 years and measured changes in three indices of functional diversity (functional dispersion, richness and evenness), community-weighted trait means and phylogenetic diversity (PD). Specific leaf area (SLA), leaf nitrogen content (LNC) and (at most sites) leaf turgor loss point (pi(TLP)) were measured for species cumulatively representing similar to 90% plant cover at each site. 3. Long-term drought led to increased community functional dispersion in three sites, with negligible effects on the remaining sites. Species re-ordering following the mortality/senescence of dominant species was the main driver of increased functional dispersion. The response of functional diversity was not consistently matched by changes in phylogenetic diversity. Community-level drought strategies (assessed as CWMs) largely shifted from drought tolerance to drought avoidance and/or escape strategies, as evidenced by higher community-weighted pi(TLP), SLA and LNC. Lastly, ecosystem drought sensitivity (i.e. relative reduction in ANPP in drought plots) was positively correlated with community-weighted SLA and negatively correlated with functional diversity. 4. Synthesis. Increased functional diversity following long-term drought may stabilize ecosystem functioning in response to future drought. However, shifts in community-scale drought strategies may increase ecosystem drought sensitivity, depending on the nature and timing of drought. Thus, our results highlight the importance of considering both functional diversity and abundance-weighted traits means of plant communities as their collective effect may either stabilize or enhance ecosystem sensitivity to drought

Growth and survival during drought: the link between hydraulic architecture and drought tolerance in grasses

Doctor of PhilosophyDepartment of AgronomyP.V. Vara PrasadThe pathway for the movement of water through plants, from the soil matrix to the atmosphere, constitutes the hydraulic architecture of a plant. The linkage between the hydraulic architecture of woody plants and drought tolerance has received considerable attention, but much less work has been done on grasses. I investigated the linkage between the hydraulic architecture of grasses to physiological patterns of water use across a range of species and conditions. The rate of stomatal conductance (g[subscript]s) and photosynthesis (A) increased acropetally along the leaves of 5 grass species, which is a unique feature of this growth form. The internal structure of leaves also changed acropetally in order to minimize the pressure gradient across the mesophyll that would otherwise occur as a result of increasing g[subscript]s. The resistance to water movement through the mesophyll represented 80-90% of leaf resistance in six genotypes of Sorghum bicolor L. (Moench). This resistance was most important in controlling g[subscript]s and A when water was readily available, but as soil-moisture decreased it was the efficient transport of water through the xylem that was most important in maintaining plant function. I also investigated the relationship between hydraulic architecture and stomatal responses of grasses to increasing Vapor Pressure Deficit (D). Grasses with a larger proportion of their hydraulic resistance within the xylem were less sensitive to increasing D and plants with high root conductance maintained higher rates of gas exchange D increased. Finally, I investigated the tolerance of grasses to extreme drought events to test if there was a trade-off between drought tolerance and growth in grasses. Plants with drought tolerant leaf traits typically sacrificed the ability to move water efficiently through their leaves. Having drought tolerant leaves did not limit the plants ability to have high rates of gas exchange, and, in fact, the most drought tolerant plants had the high rates of g[subscript]s when expressed on a mass basis. Leaf-level drought tolerance did contribute to species’ occurrence, as the drought intolerant species I studied are not commonly found in low precipitation systems. The results presented here highlight the importance of studying the hydraulic architecture of plants to provide a better understanding of what controls plant function across a range of environmental conditions

Perennial ponds are not an important source of water or dissolved organic matter to groundwaters with high arsenic concentrations in West Bengal, India

[1] Arsenic (As) concentrations and stable hydrogen (δ2H) and oxygen (δ18O) isotope ratios of surface and groundwaters from a representative site in West Bengal, India, are reported. Shallow groundwaters (≤35 m) from the study site have among the highest As concentrations measured in the entire Bengal Basin, reaching values in excess of 4600 μg kg−1. Stable isotope ratios of waters from constructed, perennial ponds indicate the ponds are chiefly recharged during the summer monsoon, and subsequently undergo extensive evaporation during the dry (winter) season. In contrast, groundwaters with high As concentrations plot along the local meteoric water line (LMWL) near where the annual, volume-weighted mean precipitation values for δ2H and δ18O would plot. The stable isotope data demonstrate that groundwaters are directly recharged by local precipitation without significant evaporation, and thus are not recharged by, nor mixed with, the pond waters. Furthermore, reactive transport modeling indicates that dissolved organic matter (DOM) derived from pond waters does not fuel microbial respiration and As mobilization at depth in the underlying aquifer because travel times for pond-derived DOM exceed groundwater ages by thousands of years. Instead, organic matter within the aquifer sediments must drive dissimilatory iron reduction and As release to groundwaters

Temperature Induced Shifts in Leaf Water Relations and Growth Efficiency Indicate Climate Change may Limit Aspen Growth in the Colorado Rockies

Higher temperatures and evaporative demand forecasted for Colorado forests by the end of the century suggest that soil water limitation increasingly will negatively impact whole plant performance. At the same time, upslope or poleward migration of plant ranges in response to warming may result in species experiencing cooler overnight temperatures, particularly if extremes in climate increase. In 2014 we established three experimental gardens along a temperature/elevation gradient to test the implications of shifting temperatures on tree function. Quaking aspen (Populus tremuloides) seedlings from a mid-elevation population were established in all three gardens and leaves were sampled monthly during the 2017 growing season. From these, we quantified the magnitude and timing of osmotic regulation, the relationships between leaf osmotic potential (Ψosm), midday leaf water potential (Ψmid), and soil moisture conditions under different temperature regimes, and growth efficiency (change in basal area/growing day). We observed a strong relationship between Ψosm and soil moisture, and a strong seasonal decline in Ψosm at the warmest and intermediate sites while the coldest site experienced a later increase in osmolytes associated with the highest degree of freeze tolerance. Growth efficiency was highest at the intermediate-temperature site – closest in elevation to the seed source location – but declined asymmetrically with warming or cooling. The novel abiotic conditions at both non-local sites resulted in declines in growth efficiency, suggesting that aspen will experience suboptimal conditions whether it stays in areas experiencing warming temperatures or if it migrates upslope to areas with colder overnight temperatures