Differences in heat tolerance, water use efficiency and growth among Douglas-fir families and varieties evidenced by GWAS and common garden studies.

Severe and frequent heat and drought events challenge the survival and development of long-generation trees. In this study, we investigated the genomic basis of heat tolerance, water use efficiency and growth by performing genome-wide association studies in coastal Douglas-fir (Pseudotsuga menziesii) and intervarietal (menziesii × glauca) hybrid seedlings. GWAS results identified 32 candidate genes involved in primary and secondary metabolism, abiotic stress and signaling, among other functions. Water use efficiency (inferred from carbon isotope discrimination), photosynthetic capacity (inferred from %N), height and heat tolerance (inferred from electrolyte leakage in a heat stress experiment) were significantly different among Douglas-fir families and varieties. High-elevation seed sources had increased water use efficiency, which could be a result of higher photosynthetic capacity. Similarly, families with greater heat tolerance also had higher water use efficiency and slower growth, suggesting a conservative growth strategy. Intervarietal hybrids showed increased heat tolerance (lower electrolyte leakage at 50 and 55 °C) and higher water use efficiency compared with coastal families, suggesting that hybridization might be a source of pre-adapted alleles to warming climates and should be considered for large-scale reforestation projects under increasingly arid conditions.

Detecting patterns of post-fire pine regeneration in a Madrean Sky Island with field surveys and remote sensing.

The American Southwest is experiencing drastic increases in aridity and wildfire incidence, triggering conversion of some frequent surface forests to non-forest. Extensive research has focused on these dynamics in regional ponderosa pine forests, but we know much less about Madrean pine-oak forests, which are broadly distributed from the Sierra Madre in Mexico to the Sky Island mountain ranges in the U.S. Increased fire incidence and drought in these forests are limiting pine regeneration and driving conversion of biodiverse forests to oak shrublands. We investigated regeneration patterns in Pinus engelmannii and P. leiophylla during severe drought 10 years after the Horseshoe Two Megafire in the Chiricahua Mountains, Arizona-a follow-up to an assessment five years post-fire. In long-term plots, we examined changes in pine seedling and resprout recruitment. Past research demonstrated that topography and fire severity influenced pine recruitment across environmental gradients. We investigated here whether Landsat-8 normalized difference vegetation index (NDVI) and evapotranspiration estimated by the ECOsystem Spaceborne Thermal Radiometer Experiment on Space Station (ECOSTRESS) added explanatory value to our understanding of these patterns. Conversion of Madrean pine-oak forest to oak shrublands continued 6-10 years post-fire. A dense, low oak canopy continued to coalesce in sites subject to severe fire. The importance of resprouts in P. leiophylla regeneration accelerated because these plants outgrew competing oak resprouts. Topography and fire severity (dNBR) were important predictors of 2021 patterns of pine recruitment. NDVI added explanatory value to these models, suggesting its potential in tracking forest dynamics. Evapotranspiration did not add value, likely because ECOSTRESS' larger pixel sizes and moving pixel locations created excessive subpixel heterogeneity in this highly dissected landscape. These models suggest that P. engelmannii is more drought sensitive, was more negatively affected by drought and fire, and is more at risk to shifts in climate and wildfires than P. leiophylla.

Forest restoration treatments increased growth and did not change survival of ponderosa pines in severe drought, Arizona.

We report on survival and growth of ponderosa pines (Pinus ponderosa Douglas ex P. Lawson & C. Lawson) 2 decades after forest restoration treatments in the G. A. Pearson Natural Area, northern Arizona. Despite protection from harvest that conserved old trees, a dense forest susceptible to uncharacteristically severe disturbance had developed during more than a century of exclusion of the previous frequent surface-fire regime that ceased upon Euro-American settlement in approximately 1876. Trees were thinned in 1993 to emulate prefire-exclusion forest conditions, accumulated forest floor was removed, and surface fire was re-introduced at 4-years intervals (full restoration). There was also a partial restoration treatment consisting of thinning alone. Compared with untreated controls, mortality of old trees (mean age 243 years, maximum 462 years) differed by <1 tree ha-1 and old-tree survival was statistically indistinguishable between treatments (90.5% control, 92.3% full, 82.6% partial). Post-treatment growth as measured by basal area increment of both old (pre-1876) and young (post-1876) pines was significantly higher in both treatments than counterpart control trees for more than 2 decades following thinning. Drought meeting the definition of megadrought affected the region almost all the time since the onset of the experiment, including 3 years that were severely dry. Growth of all trees declined in the driest 3 years, but old and young treated trees had significantly less decline. Association of tree growth with temperature (negative correlation) and precipitation (positive correlation) was much weaker in treated trees, indicating that they may experience less growth decline from warmer, drier conditions predicted in future decades. Overall, tree responses after the first 2 decades following treatment suggest that forest restoration treatments have led to substantial, sustained improvement in the growth of old and young ponderosa pines without affecting old-tree survival, thereby improving resilience to a warming climate.

Combined drought and bark beetle attacks deplete non-structural carbohydrates and promote death of mature pine trees.

How carbohydrate reserves in conifers respond to drought and bark beetle attacks are poorly understood. We investigated changes in carbohydrate reserves and carbon-dependent diterpene defences in ponderosa pine trees that were experimentally subjected to two levels of drought stress (via root trenching) and two types of biotic challenge treatments (pheromone-induced bark beetle attacks or inoculations with crushed beetles that include beetle-associated fungi) for two consecutive years. Our results showed that trenching did not influence carbohydrates, whereas both biotic challenges reduced amounts of starch and sugars of trees. However, only the combined trenched-bark beetle attacked trees depleted carbohydrates and died during the first year of attacks. While live trees contained higher carbohydrates than dying trees, amounts of constitutive and induced diterpenes produced did not vary between live and beetle-attacked dying trees, respectively. Based on these results we propose that reallocation of carbohydrates to diterpenes during the early stages of beetle attacks is limited in drought-stricken trees, and that the combination of biotic and abiotic stress leads to tree death. The process of tree death is subsequently aggravated by beetle girdling of phloem, occlusion of vascular tissue by bark beetle-vectored fungi, and potential exploitation of host carbohydrates by bark beetle symbionts as nutrients.

Phase behavior and surface tension of soft active Brownian particles.

We study quasi two-dimensional, monodisperse systems of active Brownian particles (ABPs) for a range of activities, stiffnesses, and densities. We develop a microscopic, analytical method for predicting the dense phase structure formed after motility-induced phase separation (MIPS) has occurred, including the dense cluster's area fraction, interparticle pressure, and radius. Our predictions are in good agreement with our Brownian dynamics simulations. We, then, derive a continuum model to investigate the relationship between the predicted interparticle pressure, the swim pressure, and the macroscopic pressure in the momentum equation. We find that formulating the point-wise macroscopic pressure as the interparticle pressure and modeling the particle activity through a spatially variant body force - as opposed to a volume-averaged swim pressure - results in consistent predictions of pressure in both the continuum model and the microscopic theory. This formulation of pressure also results in nearly zero surface tension for the phase separated domains, irrespective of activity, stiffness, and area fraction. Furthermore, using Brownian dynamics simulations and our continuum model, we showed that both the interface width and surface tension, are intrinsic characteristics of the system. On the other hand, if we were to exclude the body force induced by activity, we find that the resulting surface tension values are linearly dependent on the size of the simulation, in contrast to the statistical mechanical definition of surface tension.

Plant developmental stage influences responses of Pinus strobiformis seedlings to experimental warming.

Seedling emergence, survival, morphological and physiological traits, and oxidative stress resistance of southwestern white pine (Pinus strobiformis Engelm.) were studied in response to warming treatments applied during embryogenesis, germination, and early seedling growth. Daytime air temperature surrounding cones in tree canopies was warmed by +2.1°C during embryo development. Resulting seeds and seedlings were assigned to three thermal regimes in growth chambers, with each regime separated by 4°C to encompass the wide range of temperatures observed over space and time across the species' range, plus the effect of heat waves coupled with a high carbon emissions scenario of climate warming. The embryo warming treatment reduced percent seedling emergence in all germination and growth environments and reduced mortality of seedlings grown in the warmest environment. Warm thermal regimes during early seedling growth increased subsequent seedling resistance to oxidative stress and transpirational water use. Experimental warming during seed development, germination, and seedling growth affected seedling emergence and survival. Oxidative stress resistance, morphology, and water relations were affected only by warming imposed during germination and seedling growth. This work explores potential outcomes of climate warming on multiple dimensions of seedling performance and uniquely illustrates that plant responses to heat vary with plant developmental stage in addition to the magnitude of temperature change.

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.

The Fire and Tree Mortality Database, for empirical modeling of individual tree mortality after fire.

Wildland fires have a multitude of ecological effects in forests, woodlands, and savannas across the globe. A major focus of past research has been on tree mortality from fire, as trees provide a vast range of biological services. We assembled a database of individual-tree records from prescribed fires and wildfires in the United States. The Fire and Tree Mortality (FTM) database includes records from 164,293 individual trees with records of fire injury (crown scorch, bole char, etc.), tree diameter, and either mortality or top-kill up to ten years post-fire. Data span 142 species and 62 genera, from 409 fires occurring from 1981-2016. Additional variables such as insect attack are included when available. The FTM database can be used to evaluate individual fire-caused mortality models for pre-fire planning and post-fire decision support, to develop improved models, and to explore general patterns of individual fire-induced tree death. The database can also be used to identify knowledge gaps that could be addressed in future research.

Active binary mixtures of fast and slow hard spheres.

We computationally studied the phase behavior and dynamics of binary mixtures of active particles, where each species had distinct activities leading to distinct velocities, fast and slow. We obtained phase diagrams demonstrating motility-induced phase separation (MIPS) upon varying the activity and concentration of each species, and extended current kinetic theory of active/passive mixtures to active/active mixtures. We discovered two regimes of behavior quantified through the participation of each species in the dense phase compared to their monodisperse counterparts. In regime I (active/passive and active/weakly-active), we found that the dense phase was segregated by particle type into domains of fast and slow particles. Moreover, fast particles were suppressed from entering the dense phase while slow particles were enhanced entering the dense phase, compared to monodisperse systems of all-fast or all-slow particles. These effects decayed asymptotically as the activity of the slow species increased, approaching the activity of the fast species until they were negligible (regime II). In regime II, the dense phase was homogeneously mixed and each species participated in the dense phase as if it were it a monodisperse system (i.e. not mixed at all). Finally, we showed that a weighted average of constituent particle activities, which we term the net activity, defines a binodal for the MIPS transition in active/active binary mixtures. We examined the critical point of the transition and found a critical exponent (ß = 0.45) in agreement with similar studies on monodisperse systems, and distinct from equilibrium systems.

Drought-Mediated Changes in Tree Physiological Processes Weaken Tree Defenses to Bark Beetle Attack.

Interactions between water stress and induced defenses and their role in tree mortality due to bark beetles are poorly understood. We performed a factorial experiment on 48 mature ponderosa pines (Pinus ponderosa) in northern Arizona over three years that manipulated a) tree water stress by cutting roots and removing snow; b) bark beetle attacks by using pheromone lures; and c) phloem exposure to biota vectored by bark beetles by inoculating with dead beetles. Tree responses included resin flow from stem wounds, phloem composition of mono- and sesqui-terpenes, xylem water potential, leaf gas exchange, and survival. Phloem contained 21 mono- and sesqui-terpenes, which were dominated by (+)-α-pinene, (-)-limonene, and δ-3-carene. Bark beetle attacks (mostly Dendroctonus brevicomis) and biota carried by beetles induced a general increase in concentration of phloem mono- and sesqui-terpenes, whereas water stress did not. Bark beetle attacks induced an increase in resin flow for unstressed trees but not water-stressed trees. Mortality was highest for beetle-attacked water-stressed trees. Death of beetle-attacked trees was preceded by low resin flow, symptoms of water stress (low xylem water potential, leaf gas exchange), and an ephemeral increase in concentrations of mono- and sesqui-terpenes compared to surviving trees. These results show a) that ponderosa pine can undergo induction of both resin flow and phloem terpenes in response to bark beetle attack, and that the former is more constrained by water stress; b) experimental evidence that water stress predisposes ponderosa pines to mortality from bark beetles.

Disentangling the role of photosynthesis and stomatal conductance on rising forest water-use efficiency.

Multiple lines of evidence suggest that plant water-use efficiency (WUE)-the ratio of carbon assimilation to water loss-has increased in recent decades. Although rising atmospheric CO2 has been proposed as the principal cause, the underlying physiological mechanisms are still being debated, and implications for the global water cycle remain uncertain. Here, we addressed this gap using 30-y tree ring records of carbon and oxygen isotope measurements and basal area increment from 12 species in 8 North American mature temperate forests. Our goal was to separate the contributions of enhanced photosynthesis and reduced stomatal conductance to WUE trends and to assess consistency between multiple commonly used methods for estimating WUE. Our results show that tree ring-derived estimates of increases in WUE are consistent with estimates from atmospheric measurements and predictions based on an optimal balancing of carbon gains and water costs, but are lower than those based on ecosystem-scale flux observations. Although both physiological mechanisms contributed to rising WUE, enhanced photosynthesis was widespread, while reductions in stomatal conductance were modest and restricted to species that experienced moisture limitations. This finding challenges the hypothesis that rising WUE in forests is primarily the result of widespread, CO2-induced reductions in stomatal conductance.

Woody plants optimise stomatal behaviour relative to hydraulic risk.

Stomatal response to environmental conditions forms the backbone of all ecosystem and carbon cycle models, but is largely based on empirical relationships. Evolutionary theories of stomatal behaviour are critical for guarding against prediction errors of empirical models under future climates. Longstanding theory holds that stomata maximise fitness by acting to maintain constant marginal water use efficiency over a given time horizon, but a recent evolutionary theory proposes that stomata instead maximise carbon gain minus carbon costs/risk of hydraulic damage. Using data from 34 species that span global forest biomes, we find that the recent carbon-maximisation optimisation theory is widely supported, revealing that the evolution of stomatal regulation has not been primarily driven by attainment of constant marginal water use efficiency. Optimal control of stomata to manage hydraulic risk is likely to have significant consequences for ecosystem fluxes during drought, which is critical given projected intensification of the global hydrological cycle.

Scale-Up of Innovative Honeycomb Reactors for Power-to-Gas Applications - The Project Store&Go.

The German "Energiewende" is heavily based on electric power and, therefore, requests solutions to serve non-electric energy uses and to store electric energy in large scale. Synthetic natural gas (SNG) produced with hydrogen from water electrolysis and with CO2 from mainly renewable sources is one approach. For the catalytic SNG production efficient removal and utilization of the reaction heat is the main issue. A metallic honeycomb-like carrier-based reactor proved in laboratory scale to match this challenge. This type of reactor shows good heat conductivity and enables optimized operation. In the EU-funded project Store&Go the honeycomb methanation is scaled up to MW-scale. For this, heat transfer and kinetic data were determined experimentally and used in CFD calculations for the reactor design. Finally a SNG plant with 1 MW feed-in will be built and fully integrated operation will be shown.

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.

Plant water potential improves prediction of empirical stomatal models.

Climate change is expected to lead to increases in drought frequency and severity, with deleterious effects on many ecosystems. Stomatal responses to changing environmental conditions form the backbone of all ecosystem models, but are based on empirical relationships and are not well-tested during drought conditions. Here, we use a dataset of 34 woody plant species spanning global forest biomes to examine the effect of leaf water potential on stomatal conductance and test the predictive accuracy of three major stomatal models and a recently proposed model. We find that current leaf-level empirical models have consistent biases of over-prediction of stomatal conductance during dry conditions, particularly at low soil water potentials. Furthermore, the recently proposed stomatal conductance model yields increases in predictive capability compared to current models, and with particular improvement during drought conditions. Our results reveal that including stomatal sensitivity to declining water potential and consequent impairment of plant water transport will improve predictions during drought conditions and show that many biomes contain a diversity of plant stomatal strategies that range from risky to conservative stomatal regulation during water stress. Such improvements in stomatal simulation are greatly needed to help unravel and predict the response of ecosystems to future climate extremes.

Variation in species-level plant functional traits over wetland indicator status categories.

Wetland indicator status (WIS) describes the habitat affinity of plant species and is used in wetland delineations and resource inventories. Understanding how species-level functional traits vary across WIS categories may improve designations, elucidate mechanisms of adaptation, and explain habitat optima and niche. We investigated differences in species-level traits of riparian flora across WIS categories, extending their application to indicate hydrologic habitat. We measured or compiled data on specific leaf area (SLA), stem specific gravity (SSG), seed mass, and mature height of 110 plant species that occur along the Colorado River in Grand Canyon, Arizona. Additionally, we measured leaf δ13C, δ15N, % carbon, % nitrogen, and C/N ratio of 56 species with C3 photosynthesis. We asked the following: (i) How do species-level traits vary over WIS categories? (ii) Does the pattern differ between herbaceous and woody species? (iii) How well do multivariate traits define WIS categories? (iv) Which traits are correlated? The largest trait differences among WIS categories for herbaceous species occurred for SSG, seed mass, % leaf carbon and height, and for woody species occurred for height, SSG, and δ13C. SSG increased and height decreased with habitat aridity for both woody and herbaceous species. The δ13C and hence water use efficiency of woody species increased with habitat aridity. Water use efficiency of herbaceous species increased with habitat aridity via greater occurrence of C4 grasses. Multivariate trait assemblages differed among WIS categories. Over all species, SLA was correlated with height, δ13C, % leaf N, and C/N; height was correlated with SSG and % leaf C; SSG was correlated with % leaf C. Adaptations of both herbaceous and woody riparian species to wet, frequently inundated habitats include low-density stem tissue. Adaptations to drier habitats in the riparian zone include short, high-density cavitation-resistant stem tissue, and high water use efficiency. The results enhance understanding about using traits to describe plant habitat in riparian systems.

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.

Influence of pressures up to 50bar on two-stage anaerobic digestion.

The concept of pressurized two-stage anaerobic digestion integrates biogas production, purification and pressure boosting within one process. The produced methane-rich biogas can be fed into gas grids with considerably less purification effort. To investigate biogas production under high pressures up to 50bar, a lab scale two-stage anaerobic digestion system was constructed including one continuously operated pressurized methane reactor. This investigation examined the effects of different operating pressures in methane reactor (10, 25, 50bar) on biogas quantity and quality, pH value and process stability. By increasing operating pressures in methane reactor, the pH value decreased from 6.65 at 10bar to 6.55 at 50bar. Simultaneously, methane content increased from 79.08% at 10bar to 90.45% at 50bar. The results show that methane reactors can be operated up to 50bar pressure continuously representing a viable alternative to commonly used gas upgrading methods because of reduced purification effort.

Genetic variation in Pinus strobiformis growth and drought tolerance from southwestern US populations.

The persistence of some tree species is threatened by combinations of novel abiotic and biotic stressors. To examine the hypothesis that Pinus strobiformis Engelm., a tree threatened by an invasive forest pathogen and a changing climate, exhibits intraspecific genetic variation in adaptive traits, we conducted a common garden study of seedlings at one location with two watering regimes using 24 populations. Four key findings emerged: (i) growth and physiological traits were low to moderately differentiated among populations but differentiation was high for some traits in water-stressed populations; (ii) seedlings from warmer climates grew larger, had higher stomatal density and were more water-use efficient (as measured by the carbon isotope ratio) than populations from colder climates; (iii) seedlings from the northern edge of the species' distribution had lower water-use efficiency, higher stomatal conductance, slower growth and longer survival in a lethal drought experiment compared with seedlings from more southern populations; and (iv) based on non-metric multidimensional scaling analyses, populations clustered into southern and northern groups, which did not correspond to current seed transfer zones. Our discovery of a clinal geographic pattern of genetic variation in adaptive traits of P. strobiformis seedlings will be useful in developing strategies to maintain the species during ongoing climate change and in the face of an invasive pathogen.