Temperate woody species across the angiosperm phylogeny acquire tolerance to water deficit stress during the growing season.

Understanding the capacity of temperate trees to acclimate to limited soil water has become essential in the face of increasing drought risk due to climate change. We documented seasonal - or phenological - patterns in acclimation to water deficit stress in stems and leaves of tree species spanning the angiosperm phylogeny. Over 3 yr of field observations carried out in two US arboreta, we measured stem vulnerability to embolism (36 individuals of 7 Species) and turgor loss point (119 individuals of 27 species) over the growing season. We also conducted a growth chamber experiment on 20 individuals of one species to assess the mechanistic relationship between soil water restriction and acclimation. In three-quarters of species measured, plants became less vulnerable to embolism and/or loss of turgor over the growing season. We were able to stimulate this acclimatory effect by withholding water in the growth chamber experiment. Temperate angiosperms are capable of acclimation to soil water deficit stress, showing maximum vulnerability to soil water deficits following budbreak and becoming more resilient to damage over the course of the growing season or in response to simulated drought. The species-specific tempo and extent of this acclimatory potential constitutes preadaptive climate change resilience.

Phenological physiology: seasonal patterns of plant stress tolerance in a changing climate.

The physiological challenges posed by climate change for seasonal, perennial plants include increased risk of heat waves, postbudbreak freezing ('false springs'), and droughts. Although considerable physiological work has shown that the traits conferring tolerance to these stressors - thermotolerance, cold hardiness, and water deficit stress, respectively - are not static in time, they are frequently treated as such. In this review, I synthesize the recent literature on predictable seasonal - and therefore, phenological - patterns of acclimation and deacclimation to heat, cold, and water-deficit stress in perennials, focusing on woody plants native to temperate climates. I highlight promising, high-throughput techniques for quantifying thermotolerance, cold hardiness, and drought tolerance. For each of these forms of stress tolerance, I summarize the current balance of evidence regarding temporal patterns over the course of a year and suggest a characteristic temporal scale in these responses to environmental stress. In doing so, I offer a synthetic framework of 'phenological physiology', in which understanding and leveraging seasonally recurring (phenological) patterns of physiological stress acclimation can facilitate climate change adaptation and mitigation.

Coupling spectral and resource-use complementarity in experimental grassland and forest communities.

Reflectance spectra provide integrative measures of plant phenotypes by capturing chemical, morphological, anatomical and architectural trait information. Here, we investigate the linkages between plant spectral variation, and spectral and resource-use complementarity that contribute to ecosystem productivity. In both a forest and prairie grassland diversity experiment, we delineated n-dimensional hypervolumes using wavelength bands of reflectance spectra to test the association between the spectral space occupied by individual plants and their growth, as well as between the spectral space occupied by plant communities and ecosystem productivity. We show that the spectral space occupied by individuals increased with their growth, and the spectral space occupied by plant communities increased with ecosystem productivity. Furthermore, ecosystem productivity was better explained by inter-individual spectral complementarity than by the large spectral space occupied by productive individuals. Our results indicate that spectral hypervolumes of plants can reflect ecological strategies that shape community composition and ecosystem function, and that spectral complementarity can reveal resource-use complementarity.

Standardization of electrolyte leakage data and a novel liquid nitrogen control improve measurements of cold hardiness in woody tissue.

BACKGROUND: A variety of basic and applied research programs in plant biology require the accurate and reliable determination of plant tissue cold hardiness. Over the past 50 years, the electrolyte leakage method has emerged as a popular and practical method for quantifying the amount of damage inflicted on plant tissue by exposure to freezing temperatures. Numerous approaches for carrying out this method and analyzing the resultant data have emerged. These include multiple systems for standardizing and modeling raw electrolyte leakage data and multiple protocols for boiling or autoclaving samples in order to maximize leakage as a positive control. We compare four different routines for standardization of leakage data and assess a novel control method-immersion in liquid nitrogen in lieu of traditional autoclaving-and apply them to woody twigs collected from 12 maple (Acer) species in early spring. We compare leakage data from these samples using each of four previously published forms of data analysis and autoclaving vs. liquid nitrogen controls and validate each of these approaches against visual estimates of freezing damage and differential thermal analysis. RESULTS: Through presentation of our own data and re-analysis of previously published findings, we show that standardization of raw data against estimates of both minimum and maximum attainable freezing damage allows for reliable estimation of cold hardiness at the species level and across studies in diverse systems. Furthermore, use of our novel liquid nitrogen control produces data commensurate across studies and enhances the consistency and realism of the electrolyte leakage method, especially for very cold hardy samples. CONCLUSION: Future leakage studies that relativize data against minimum and maximum leakage and that employ our updated liquid nitrogen control will contribute generalizable, repeatable, and realistic data to the existing body of cold hardiness research in woody plants. Data from studies conducted using a liquid nitrogen (and not an autoclaving) control can still be compared to previously published data, especially when raw data are standardized using the best-performing approach among those we assessed. Electrolyte leakage of woody twigs emerges as a useful technique for quickly assessing the probability of tissue death in response to freezing in dormant plants. Differential thermal analysis may provide different and complementary information on cold hardiness.

Context Dependence of Local Adaptation to Abiotic and Biotic Environments: A Quantitative and Qualitative Synthesis.

Understanding how spatially variable selection shapes adaptation is an area of long-standing interest in evolutionary ecology. Recent meta-analyses have quantified the extent of local adaptation, but the relative importance of abiotic and biotic factors in driving population divergence remains poorly understood. To address this gap, we combined a quantitative meta-analysis and a qualitative metasynthesis to (1) quantify the magnitude of local adaptation to abiotic and biotic factors and (2) characterize major themes that influence the motivation and design of experiments that seek to test for local adaptation. Using local-foreign contrasts as a metric of local adaptation (or maladaptation), we found that local adaptation was greater in the presence than in the absence of a biotic interactor, especially for plants. We also found that biotic environments had stronger effects on fitness than abiotic environments when ignoring whether those environments were local versus foreign. Finally, biotic effects were stronger at low latitudes, and abiotic effects were stronger at high latitudes. Our qualitative analysis revealed that the lens through which local adaptation has been examined differs for abiotic and biotic factors. It also revealed biases in the design and implementation of experiments that make quantitative results challenging to interpret and provided directions for future research.

Drought and freezing vulnerability of the isolated hybrid aspen Populus x smithii relative to its parental species, P. tremuloides and P. grandidentata.

AIM: We assessed the vulnerability of an isolated, relictual Pleistocene hybrid aspen population of conservation interest (Populus x. smithii) and the nearest populations of its parent species (Populus grandidentata and Populus tremuloides) to springtime post-bud break freezing and growing season drought stress. Response to these stressors in the three taxa was compared in terms of avoidance and tolerance. LOCATION: North American Midwest; USA. METHODS: Unique genets from the hybrid Niobrara River population and from the two parental populations were propagated in a common garden from rhizome cuttings. We tracked their phenology before and after bud break and measured their vulnerability to freezing (stem electrolyte leakage and leaf chlorophyll fluorescence) and to drought (stem hydraulic conductance, leaf osmotic potential, stomatal pore index, and gas exchange). RESULTS: Populus grandidentata was slower to leaf out, showed lower vulnerability to stem freezing and drought-induced cavitation, but exhibited a lower capacity to tolerate drought stress through leaf resistance traits compared to P. tremuloides. Hybrids were similar to P. grandidentata in their overwintering strategy, exhibiting later bud break, and in their higher resistance to stem freezing damage, but they were more similar to P. tremuloides in their higher vulnerability to drought-induced cavitation. The hybrids shared various leaf-level gas exchange traits with both parents. All aspens showed limited loss of leaf photosynthetic function following moderate freezing. MAIN CONCLUSIONS: The Niobrara River hybrid population is vulnerable to drought due to its combination of inherited drought avoidance and tolerance traits. As climate changes, P. x smithii will likely suffer from increased drought stress, while being unaffected by frost during warmer springs. The two parental species contrast in their survival mechanisms in response to climatic stress, with P. tremuloides tending toward freezing tolerance but drought avoidance and P. grandidentata tending toward freezing avoidance and drought tolerance.

Non-symbiotic soil microbes are more strongly influenced by altered tree biodiversity than arbuscular mycorrhizal fungi during initial forest establishment.

While the relationship between plant and microbial diversity has been well studied in grasslands, less is known about similar relationships in forests, especially for obligately symbiotic arbuscular mycorrhizal (AM) fungi. To assess the effect of varying tree diversity on microbial alpha- and beta-diversity, we sampled soil from plots in a high-density tree diversity experiment in Minnesota, USA, 3 years after establishment. About 3 of 12 tree species are AM hosts; the other 9 primarily associate with ectomycorrhizal fungi. We used phospho- and neutral lipid fatty acid analysis to characterize the biomass and functional identity of the whole soil bacterial and fungal community and high throughput sequencing to identify the species-level richness and composition of the AM fungal community. We found that plots of differing tree composition had different bacterial and fungal communities; plots with conifers, and especially Juniperus virginiana, had lower densities of several bacterial groups. In contrast, plots with a higher density or diversity of AM hosts showed no sign of greater AM fungal abundance or diversity. Our results indicate that early responses to plant diversity vary considerably across microbial groups, with AM fungal communities potentially requiring longer timescales to respond to changes in host tree diversity.

Species richness and traits predict overyielding in stem growth in an early-successional tree diversity experiment.

Over the last two decades, empirical work has established that higher biodiversity can lead to greater primary productivity; however, the importance of different aspects of biodiversity in contributing to such relationships is rarely elucidated. We assessed the relative importance of species richness, phylogenetic diversity, functional diversity, and identity of neighbors for stem growth 3 yr after seedling establishment in a tree diversity experiment in eastern Minnesota. Generally, we found that community-weighted means of key functional traits (including mycorrhizal association, leaf nitrogen and calcium, and waterlogging tolerance) as well as species richness were strong, independent predictors of stem biomass growth. More phylogenetically diverse communities did not consistently produce more biomass than expected, and the trait values or diversity of individual functional traits better predicted biomass production than did a multidimensional functional diversity metric. Furthermore, functional traits and species richness best predicted growth at the whole-plot level (12 m2 ), whereas neighborhood composition best predicted growth at the focal tree level (0.25 m2 ). The observed effects of biodiversity on growth appear strongly driven by positive complementary effects rather than by species-specific selection effects, suggesting that synergistic species' interactions rather than the influence of a few important species may drive overyielding.

Relationships between Spatial Metrics and Plant Diversity in Constructed Freshwater Wetlands.

The diversity of plant species and their distribution in space are both thought to have important effects on the function of wetland ecosystems. However, knowledge of the relationships between plant species and spatial diversity remains incomplete. In this study, we investigated relationships between spatial pattern and plant species diversity over a five year period following the initial restoration of experimental wetland ecosystems. In 2003, six identical and hydrologically-isolated 0.18 ha wetland "cells" were constructed in former farmland in northeast Ohio. The systems were subjected to planting treatments that resulted in different levels of vascular plant species diversity among cells. Plant species diversity was assessed through annual inventories. Plant spatial pattern was assessed by digitizing low-altitude aerial photographs taken at the same time as the inventories. Diversity metrics derived from the inventories were significantly related to certain spatial metrics derived from the photographs, including cover type diversity and contagion. We found that wetlands with high cover type diversity harbor higher plant species diversity than wetlands with fewer types of patches. We also found significant relationships between plant species diversity and spatial patterning of patch types, but the direction of the effect differed depending on the diversity metric used. Links between diversity and spatial pattern observed in this study suggest that high-resolution aerial imagery may provide wetland scientists with a useful tool for assessing plant diversity.