In light of the changes in precipitation and soil water availability expected with climate change, understanding the mechanisms underlying plant responses to water deficit is essential. Toward that ...end we have conducted an integrative analysis of responses to drought stress in the perennial C4 grass and biofuel crop, Panicum virgatum (switchgrass). Responses to soil drying and re-watering were measured at transcriptional, physiological, and metabolomic levels. To assess the interaction of soil moisture with diel light: dark cycles, we profiled gene expression in drought and control treatments under pre-dawn and mid-day conditions.
Soil drying resulted in reduced leaf water potential, gas exchange, and chlorophyll fluorescence along with differential expression of a large fraction of the transcriptome (37%). Many transcripts responded differently depending on time of day (e.g. up-regulation pre-dawn and down-regulation mid-day). Genes associated with C4 photosynthesis were down-regulated during drought, while C4 metabolic intermediates accumulated. Rapid changes in gene expression were observed during recovery from drought, along with increased water use efficiency and chlorophyll fluorescence.
Our findings demonstrate that drought responsive gene expression depends strongly on time of day and that gene expression is extensively modified during the first few hours of drought recovery. Analysis of covariation in gene expression, metabolite abundance, and physiology among plants revealed non-linear relationships that suggest critical thresholds in drought stress responses. Future studies may benefit from evaluating these thresholds among diverse accessions of switchgrass and other C4 grasses.
1. Quantification of stomatal responses to environmental variables, in particular to soil water status, is needed to model carbon and water exchange rates between plants and the atmosphere. 2. Models ...based on stomatal optimality theory successfully describe leaf gas exchange under different environmental conditions, but the effects of water availability on the key optimization parameter the marginal water use efficiency (WUE), λ = əA/əEhas resisted complete theoretical treatment. Building on previous optimal leaf gas exchange models, we developed an analytical equation to estimate λ from gas exchange observations along gradients of soil water availability. This expression was then used in a meta-analysis of about 50 species to investigate patterns of variation in λ. 3. Assuming that cuticular water losses are negligible λ increases under mild water stress but decreases when severe water stress limits photosynthesis. When cuticular conductance is considered, however, λ increases monotonically with increasing water stress, in agreement with previous theoretical predictions. Moreover, the shape of these response curves to soil water availability changes with plant functional type and climatic conditions. In general, λ is lower in species grown in dry climates, indicating lower marginal WUE. 4. The proposed parameterization provides a framework to assess the responses of leaf gas exchange to changes in water availability. Moreover, the model can be extended to scale leaflevel fluxes to the canopy and ecosystem level.
Amplification of the hydrological cycle as a consequence of global warming is forecast to lead to more extreme intra-annual precipitation regimes characterized by larger rainfall events and longer ...intervals between events. We present a conceptual framework, based on past investigations and ecological theory, for predicting the consequences of this underappreciated aspect of climate change. We consider a broad range of terrestrial ecosystems that vary in their overall water balance. More extreme rainfall regimes are expected to increase the duration and severity of soil water stress in mesic ecosystems as intervals between rainfall events increase. In contrast, xeric ecosystems may exhibit the opposite response to extreme events. Larger but less frequent rainfall events may result in proportional reductions in evaporative losses in xeric systems, and thus may lead to greater soil water availability. Hydric (wetland) ecosystems are predicted to experience reduced periods of anoxia in response to prolonged intervals between rainfall events. Understanding these contingent effects of ecosystem water balance is necessary for predicting how more extreme precipitation regimes will modify ecosystem processes and alter interactions with related global change drivers.
How ecosystem productivity and species richness are interrelated is one of the most debated subjects in the history of ecology. Decades of intensive study have yet to discern the actual mechanisms ...behind observed global patterns. Here, by integrating the predictions from multiple theories into a single model and using data from 1,126 grassland plots spanning five continents, we detect the clear signals of numerous underlying mechanisms linking productivity and richness. We find that an integrative model has substantially higher explanatory power than traditional bivariate analyses. In addition, the specific results unveil several surprising findings that conflict with classical models. These include the isolation of a strong and consistent enhancement of productivity by richness, an effect in striking contrast with superficial data patterns. Also revealed is a consistent importance of competition across the full range of productivity values, in direct conflict with some (but not all) proposed models. The promotion of local richness by macroecological gradients in climatic favourability, generally seen as a competing hypothesis, is also found to be important in our analysis. The results demonstrate that an integrative modelling approach leads to a major advance in our ability to discern the underlying processes operating in ecological systems.
Invasive plant species often dominate native species in competition, augmenting other potential advantages such as release from natural enemies. Resource pre-emption may be a particularly important ...mechanism for establishing dominance over competitors of the same functional type. We hypothesized that competitive success of an exotic grass against native grasses is mediated by establishing an early size advantage. We tested this prediction among four perennial C4 warm-season grasses: the exotic weed Johnsongrass (Sorghum halepense), big bluestem (Andropogon gerardii), little bluestem (Schizachyrium scoparius) and switchgrass (Panicum virgatum). We predicted that a) the competitive effect of Johnsongrass on target species would be proportional to their initial biomass difference, b) competitive effect and response would be negatively correlated and c) soil fertility would have little effect on competitive relationships. In a greenhouse, plants of the four species were grown from seed either alone or with one Johnsongrass neighbor at two fertilizer levels and periodically harvested. The first two hypotheses were supported: The seedling biomass of single plants at first harvest (50 days after seeding) ranked the same way as the competitive effect of Johnsongrass on target species: Johnsongrass < big bluestem < little bluestem/switchgrass, while Johnsongrass responded more strongly to competition from Johnsongrass than from native species. At final harvest, native plants growing with Johnsongrass attained between 2-5% of their single-plant non-root biomass, while Johnsongrass growing with native species attained 89% of single-plant non-root biomass. Fertilization enhanced Johnsongrass' competitive effects on native species, but added little to the already severe competitive suppression. Accelerated early growth of Johnsongrass seedlings relative to native seedlings appeared to enable subsequent resource pre-emption. Size-asymmetric competition and resource-pre-emption may be a critical mechanism by which exotic invasive species displace functionally similar native species and alter the functional dynamics of native communities.
Increased nutrient inputs due to anthropogenic activity are expected to increase primary productivity across terrestrial ecosystems, but changes in allocation aboveground versus belowground with ...nutrient addition have different implications for soil carbon (C) storage. Thus, given that roots are major contributors to soil C storage, understanding belowground net primary productivity (BNPP) and biomass responses to changes in nutrient availability is essential to predicting carbon–climate feedbacks in the context of interacting global environmental changes. To address this knowledge gap, we tested whether a decade of nitrogen (N) and phosphorus (P) fertilization consistently influenced aboveground and belowground biomass and productivity at nine grassland sites spanning a wide range of climatic and edaphic conditions in the continental United States. Fertilization effects were strong aboveground, with both N and P addition stimulating aboveground biomass at nearly all sites (by 30% and 36%, respectively, on average). P addition consistently increased root production (by 15% on average), whereas other belowground responses to fertilization were more variable, ranging from positive to negative across sites. Site‐specific responses to P were not predicted by the measured covariates. Atmospheric N deposition mediated the effect of N fertilization on root biomass and turnover. Specifically, atmospheric N deposition was positively correlated with root turnover rates, and this relationship was amplified with N addition. Nitrogen addition increased root biomass at sites with low N deposition but decreased it at sites with high N deposition. Overall, these results suggest that the effects of nutrient supply on belowground plant properties are context dependent, particularly with regard to background N supply rates, demonstrating that site conditions must be considered when predicting how grassland ecosystems will respond to increased nutrient loading from anthropogenic activity.
Long-term climate change and periodic environmental extremes threaten food and fuel security
and global crop productivity
. Although molecular and adaptive breeding strategies can buffer the effects ...of climatic stress and improve crop resilience
, these approaches require sufficient knowledge of the genes that underlie productivity and adaptation
-knowledge that has been limited to a small number of well-studied model systems. Here we present the assembly and annotation of the large and complex genome of the polyploid bioenergy crop switchgrass (Panicum virgatum). Analysis of biomass and survival among 732 resequenced genotypes, which were grown across 10 common gardens that span 1,800 km of latitude, jointly revealed extensive genomic evidence of climate adaptation. Climate-gene-biomass associations were abundant but varied considerably among deeply diverged gene pools. Furthermore, we found that gene flow accelerated climate adaptation during the postglacial colonization of northern habitats through introgression of alleles from a pre-adapted northern gene pool. The polyploid nature of switchgrass also enhanced adaptive potential through the fractionation of gene function, as there was an increased level of heritable genetic diversity on the nondominant subgenome. In addition to investigating patterns of climate adaptation, the genome resources and gene-trait associations developed here provide breeders with the necessary tools to increase switchgrass yield for the sustainable production of bioenergy.
Grassland production is sensitive to both precipitation and plant N accumulation and utilization, such that change in one variable influences grassland response to the second variable. We ...investigated effects of interannual variation in precipitation on the response of ‘community’-scale values of relative growth rate (RGR) to two multiplicative components of RGR, nitrogen productivity (NP; rate of change in biomass/plant N), an index of N utilization efficiency, and plant N concentration (N), in two grasslands in Texas, USA. Grasslands included a planted mixture of perennial grass and forb species and monoculture of the perennial C
4
grass
Panicum virgatum
that was invaded by multiple plant species. RGR and its N components were measured at the spatial scale of 7-m diameter circular patches near the spring peak in mixture biomass during each of 5 years. We found that RGR varied substantially among patches and years and between the planted mixture and monoculture. RGR variation was strongly correlated with variation in NP. Precipitation during the 3 months prior to RGR measurement mediated the RGR response to NP by altering the correlation between NP and N in both grasslands. Reduced precipitation led to more negative NP-N correlation coefficients, which reduced proportional change in RGR per change in NP by as much as 30% even in the absence of a precipitation effect on means of RGR and NP. Our results highlight an under-appreciated aspect of the pervasive role of precipitation in grassland growth that was mediated via change in the growth benefit derived from plant N.
Climate changes, including chronic changes in precipitation amounts, will influence plant physiology and growth. However, such precipitation effects on switchgrass, a major bioenergy crop, have not ...been well investigated. We conducted a two-year precipitation simulation experiment using large pots (95 L) in an environmentally controlled greenhouse in Nashville, TN. Five precipitation treatments (ambient precipitation, and -50%, -33%, +33%, and +50% of ambient) were applied in a randomized complete block design with lowland "Alamo" switchgrass plants one year after they were established from tillers. The growing season progression of leaf physiology, tiller number, height, and aboveground biomass were determined each growing season. Precipitation treatments significantly affected leaf physiology, growth, and aboveground biomass. The photosynthetic rates in the wet (+50% and +33%) treatments were significantly enhanced by 15.9% and 8.1%, respectively, than the ambient treatment. Both leaf biomass and plant height were largely increased, resulting in dramatically increases in aboveground biomass by 56.5% and 49.6% in the +50% and +33% treatments, respectively. Compared to the ambient treatment, the drought (-33% and -50%) treatments did not influence leaf physiology, but the -50% treatment significantly reduced leaf biomass by 37.8%, plant height by 16.3%, and aboveground biomass by 38.9%. This study demonstrated that while switchgrass in general is a drought tolerant grass, severe drought significantly reduces Alamo's growth and biomass, and that high precipitation stimulates its photosynthesis and growth.
Rising atmospheric CO2 concentration directly stimulates plant productivity and affects nutrient dynamics in the soil. However, the influence of CO2 enrichment on soil bacterial communities remains ...elusive, likely due to their complex interactions with a wide range of plant and soil properties. Here, we investigated the bacterial community response to a decade long preindustrial‐to‐future CO2 gradient (250–500 ppm) among three contrasting soil types using 16S rRNA gene amplicon sequencing. In addition, we examined the effect of seasonal variation and plant species composition on bacterial communities. We found that Shannon index (H’) and Faith's phylogenetic diversity (PD) did not change in response to the CO2 gradient (R2 = 0.01, p > 0.05). CO2 gradient and season also had a negligible effect on overall community structure, although silty clay soil communities were better structured on a CO2 gradient (p < 0.001) among three soils. Similarly, CO2 gradient had no significant effect on the relative abundance of different phyla. However, we observed soil‐specific variation of CO2 effects in a few individual families. For example, the abundance of Pirellulaceae family decreased linearly with CO2 gradient, but only in sandy loam soils. Conversely, the abundance of Micromonosporaceae and Gaillaceae families increased with CO2 gradient in clay soils. Soil water content (SWC) and nutrient properties were the key environmental constraints shaping bacterial community structure, one manifestation of which was a decline in bacterial diversity with increasing SWC. Furthermore, the impact of plant species composition on community structure was secondary to the strong influence of soil properties. Taken together, our findings indicate that bacterial communities may be largely unresponsive to indirect effects of CO2 enrichment through plants. Instead, bacterial communities are strongly regulated by edaphic conditions, presumably because soil differences create distinct environmental niches for bacteria.
We investigated the bacterial community response to a decade‐long preindustrial‐to‐future CO2 gradient (250–500 ppm) among three contrasting soil types using 16S rRNA gene amplicon sequencing. We found that bacterial communities may be largely unresponsive to indirect effects of CO2 enrichment through plants. Instead, bacterial communities are strongly regulated by edaphic conditions, presumably because soil differences create distinct environmental niches for bacteria.
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