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Keith Reinhardt

Publications and source records attributed to Keith Reinhardt.

9 recordsLinked to original sources

Cytotype and local adaptation drive phenotypic variation in two subspecies of big sagebrush (Artemisia tridentata)

Big sagebrush ( Artemisia tridentata ) is a widespread and locally dominant shrub throughout many ecosystems in western North America. A. tridentata ssps. tridentata and wyomingensis are two subspecies whose populations occupy the warm-arid regions of the species range and whose trailing edge is threatened by climate change. Previous studies have presented conflicting results in relation to the genetic control of physiological variation in A. tridentata . Understanding how different genetic factors contribute to physiological variation can provide insight into how these two subspecies may respond to future climate change. To explore possible variation among and within two subspecies of A. tridentata , we measured physiological and morphological traits in A . t . tridentata and A. t. wyomingensis during mid-summer (July), seven years after establishment in a common garden. Contributions to trait variation were quantified for both genetic (subspecies and cytotype) and environmental (climate-of-origin) factors. Measurements revealed an unequal contribution to phenotypic variation by subspecies, cytotype, and climate-of-origin. Ploidy and climate-of-origin were more important than subspecies in driving phenotypic variation in A. tridentata. These findings suggest that A. tridentata has a highly plastic drought response, or that culling (mortality over time due to environmental factors) in the common garden over seven years has led to a lack of genetic diversity within the garden. Understanding what factors drive phenotypic expression in big sagebrush can provide better insight into how climate change may affect migration and extirpation and may aid in the effectiveness of restoration efforts.

Idaho

High-resolution thermal imagery reveals how interactions between crown structure and genetics shape plant temperature

Understanding interactions between environmental stress and genetic variation is crucial to predict the adaptive capacity of species to climate change. Leaf temperature is both a driver and a responsive indicator of plant physiological response to thermal stress, and methods to monitor it are needed. Foliar temperatures vary across leaf to canopy scales and are influenced by genetic factors, challenging efforts to map and model this critical variable. Thermal imagery collected using unoccupied aerial systems (UAS) offers an innovative way to measure thermal variation in plants across landscapes at leaf-level resolutions. We used a UAS equipped with a thermal camera to assess temperature variation among genetically distinct populations of big sagebrush ( Artemisia tridentata ), a keystone plant species that is the focus of intensive restoration efforts throughout much of western North America. We completed flights across a growing season in a sagebrush common garden to map leaf temperature relative to subspecies and cytotype, physiological phenotypes of plants, and summer heat stress. Our objectives were to (1) determine whether leaf-level stomatal conductance corresponds with changes in crown temperature; (2) quantify genetic (i.e., subspecies and cytotype) contributions to variation in leaf and crown temperatures; and (3) identify how crown structure, solar radiation, and subspecies-cytotype relate to leaf-level temperature. When considered across the whole season, stomatal conductance was negatively, non-linearly correlated with crown-level temperature derived from UAS. Subspecies identity best explained crown-level temperature with no difference observed between cytotypes. However, structural phenotypes and microclimate best explained leaf-level temperature. These results show how fine-scale thermal mapping can decouple the contribution of genetic, phenotypic, and microclimate factors on leaf temperature dynamics. As climate-change-induced heat stress becomes prevalent, thermal UAS represents a promising way to track plant phenotypes that emerge from gene-by-environment interactions.

Idaho

Warming temperatures affect meadow-wide nectar resources, with implications for plant-pollinator communities

Nectar production may be a point of sensitivity that can help link primary and secondary trophic responses to climate shifts, and is therefore important to our understanding of ecosystem responses. We evaluated the nectar response of two widespread native forbs, Balsamorhiza sagittata and Eriogonum umbellatum , to experimental warming in a high-elevation sagebrush meadow in the Teton Range, WY, USA, over two years, 2015 and 2016. Warming treatments reduced the occurrence of nighttime freezing and nectar volume but increased sugar concentration in nectar in both species in both years. Warming effects were also evident in a consistent increase in the number of flowers produced by B. sagittata . Our research suggests that warming associated with climate change has the potential to induce shifts in the nectar-feeding community by changing nectar characteristics such as volume and sugar concentration to which nectar feeders are adapted.

Wyoming

Snowmelt timing regulates community composition, phenology, and physiological performance of alpine plants

The spatial patterning of alpine plant communities is strongly influenced by the variation in physical factors such as temperature and moisture, which are strongly affected by snow depth and snowmelt patterns. Earlier snowmelt timing and greater soil-moisture limitations may favor wide-ranging species adapted to a broader set of ecohydrological conditions than alpine-restricted species. We asked how plant community composition, phenology, plant water relations, and photosynthetic gas exchange of alpine-restricted and wide-ranging species differ in their responses to a ca. 40-day snowmelt gradient in the Colorado Rocky Mountains ( Lewisia pygmaea , Sibbaldia procumbens , and Hymenoxys grandiflora were alpine-restricted and Artemisia scopulorum , Carex rupestris , and Geum rossii were wide-ranging species). As hypothesized, species richness and foliar cover increased with earlier snowmelt, due to a greater abundance of wide-ranging species present in earlier melting plots. Flowering initiation occurred earlier with earlier snowmelt for 12 out of 19 species analyzed, while flowering duration was shortened with later snowmelt for six species (all but one were wide-ranging species). We observed >50% declines in net photosynthesis from July to September as soil moisture and plant water potentials declined. Early-season stomatal conductance was higher in wide-ranging species, indicating a more competitive strategy for water acquisition when soil moisture is high. Even so, there were no associated differences in photosynthesis or transpiration, suggesting no strong differences between these groups in physiology. Our findings reveal that plant species with different ranges (alpine-restricted vs. wide-ranging) could have differential phenological and physiological responses to snowmelt timing and associated soil moisture dry-down, and that alpine-restricted species’ performance is more sensitive to snowmelt. As a result, alpine-restricted species may serve as better indicator species than their wide-ranging heterospecifics. Overall, alpine community composition and peak % cover are strongly structured by spatio-temporal patterns in snowmelt timing. Thus, near-term, community-wide changes (or variation) in phenology and physiology in response to shifts in snowmelt timing or rates of soil dry down are likely to be contingent on the legacy of past climate on community structure.

Frontiers in Plant Science

Changes in structure and physiological functioning due to experimentally enhanced precipitation seasonality in a widespread shrub species

Semi-arid shrub steppe occupies a vast geographic range that is characterized in part by distinct seasonal patterns in precipitation. Few studies have evaluated how variability in both the amount and timing of precipitation affect the structure and physiology of shrubs in these systems. We quantified changes in foliar crown parameters, xylem anatomy, gas exchange, and hydraulic transport capacity in deep-rooted Artemisia tridentata shrubs following 20 years of experimental manipulations in amount and seasonal timing of precipitation. We hypothesized that shrub growth (total leaf area per shrub and cover of shrub community), hydraulic transport efficiency, and gas exchange would increase in shrubs in irrigated plots compared to non-irrigated control plots, especially for irrigation applied in winter rather than summer. We also predicted similar changes in xylem anatomy (ring width, vessel size and frequency). Most treatment responses entailed changes in plant structure, and were generally consistent with our hypotheses: total-shrub leaf area, shrub basal area, canopy cover, and maximum sapwood-specific branch hydraulic conductivity were more than 2× greater in shrubs in winter-irrigated compared to control plots, while summer irrigation had few effects on these variables. Irrigation in either season did not affect xylem vessel size, but did increase xylem ring width by ~ 2 × and decreased xylem vessel frequency by about half. Anatomical, morphological, and stand-level abundance of A. tridentata appeared much more responsive to irrigation than state changes in gas exchange, particularly when the extra water is received during winter. Thus, it appears for sagebrush that seasonal timing is at least as important as the amount of precipitation, and that responses to changes in precipitation timing occur through changes in carbon allocation more so than changes in leaf-level carbon gain.

Plant Ecology

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere–atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Nature Ecology & Evolution

Response of aboveground carbon balance to long-term, experimental enhancements in precipitation seasonality is contingent on plant community type in cold-desert rangelands

Semi-arid rangelands are important carbon (C) pools at global scales. However, the degree of net C storage or release in water-limited systems is a function of precipitation amount and timing, as well as plant community composition. In northern latitudes of western North America, C storage in cold-desert ecosystems could increase with boosts in wintertime precipitation, in which climate models predict, due to increases in wintertime soil water storage that enhance summertime productivity. However, there are few long-term, manipulative field-based studies investigating how rangelands will respond to altered precipitation amount or timing. We measured aboveground C pools and fluxes at leaf, soil, and ecosystem scales over a single growing season in plots that had 200 mm of supplemental precipitation added in either winter or summer for the past 21 years, in shrub- and exotic-bunchgrass-dominated garden plots. At our cold-desert site (298 mm precipitation during the study year), we hypothesized that increased winter precipitation would stimulate the aboveground C uptake and storage relative to ambient conditions, especially in plots containing shrubs. Our hypotheses were generally supported: ecosystem C uptake and long-term biomass accumulation were greater in winter- and summer-irrigated plots compared to control plots in both vegetation communities. However, substantial increases in the aboveground biomass occurred only in winter-irrigated plots that contained shrubs. Our findings suggest that increases in winter precipitation will enhance C storage of this widespread ecosystem, and moreso in shrub- compared to grass-dominated communities.

Idaho

Linking carbon and water limitations to drought-induced mortality of Pinus flexilis seedlings

Survival of tree seedlings at high elevations has been shown to be limited by thermal constraints on carbon balance, but it is unknown if carbon relations also limit seedling survival at lower elevations, where water relations may be more important. We measured and modeled carbon fluxes and water relations in first-year Pinus flexilis seedlings in garden plots just beyond the warm edge of their natural range, and compared these with dry-mass gain and survival across two summers. We hypothesized that mortality in these seedlings would be associated with declines in water relations, more so than with carbon-balance limitations. Rather than gradual declines in survivorship across growing seasons, we observed sharp, large-scale mortality episodes that occurred once volumetric soil-moisture content dropped below 10%. By this point, seedling water potentials had decreased below −5 MPa, seedling hydraulic conductivity had decreased by 90% and seedling hydraulic resistance had increased by >900%. Additionally, non-structural carbohydrates accumulated in aboveground tissues at the end of both summers, suggesting impairments in phloem-transport from needles to roots. This resulted in low carbohydrate concentrations in roots, which likely impaired root growth and water uptake at the time of critically low soil moisture. While photosynthesis and respiration on a leaf area basis remained high until critical hydraulic thresholds were exceeded, modeled seedling gross primary productivity declined steadily throughout the summers. At the time of mortality, modeled productivity was insufficient to support seedling biomass-gain rates, metabolism and secondary costs. Thus the large-scale mortality events that we observed near the end of each summer were most directly linked with acute, episodic declines in plant hydraulic function that were linked with important changes in whole-seedling carbon relations.

Tree Physiology

Desert shrub responses to experimental modification of precipitation seasonality and soil depth: relationship to the two-layer model and ecohydrological niche

1. Ecohydrological niches are important for understanding plant community responses to climate shifts, particularly in dry lands. According to the two-layer hypothesis, selective use of deep-soil water increases growth or persistence of woody species during warm and dry summer periods and thereby contributes to their coexistence with shallow-rooted herbs in dry ecosystems. The resource-pool hypothesis further suggests that shallow-soil water benefits growth of all plants while deep-soil water primarily enhances physiological maintenance and survival of woody species. Few studies have directly tested these by manipulating deep-soil water availability and observing the long-term outcomes. 2. We predicted that factors promoting infiltration and storage of water in deep soils, specifically greater winter precipitation and soil depth, would enhance Artemisia tridentata (big sagebrush) in cold, winter-wet/summer-dry desert. Sagebrush responses to 20 years of winter irrigation were compared to summer- or no irrigation, on plots having relatively deep or shallow soils (2 m vs. 1 m depths). 3. Winter irrigation increased sagebrush cover, and crown and canopy volumes, but not density (individuals/plot) compared to summer or no irrigation, on deep-soil plots. On shallow-soil plots, winter irrigation surprisingly decreased shrub cover and size, and summer irrigation had no effect. Furthermore, multiple regression suggested that the variations in growth were related (i) firstly to water in shallow soils (0-0.2 m) and secondly to deeper soils (> 1 m deep) and (ii) more by springtime than by midsummer soil water. Water-use efficiency increased considerably on shallow soils without irrigation and was lowest with winter irrigation. 4. Synthesis . Sagebrush was more responsive to the seasonal timing of precipitation than to total annual precipitation. Factors that enhanced deep-water storage (deeper soils plus more winter precipitation) led to increases in Artemisia tridentata that were consistent with the two-layer hypothesis, and the contribution of shallow water to growth on these plots was consistent with the resource-pool hypothesis. However, shallow-soil water also had negative effects on sagebrush, suggesting an ecohydrological trade-off not considered in these or related theories. The interaction between precipitation timing and soil depth indicates that increased winter precipitation could lead to a mosaic of increases and decreases in A. tridentata across landscapes having variable soil depth.

Idaho