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Dickman, L. Turin

Publications and source records attributed to Dickman, L. Turin.

Increased water availability at various timescales has different effects on stomatal closure point in isohydric piñon pine and anisohydric juniper

Abstract Stomatal Closure Point (SCP) has commonly been used to describe drought response strategies in plants, with isohydric species maintaining relatively high, constant SCP compared to anisohydric species that can lower SCP with increasing drought severity. However, there is evidence that, within these groups, SCP may respond dynamically to environmental conditions. Here, we explored how increasing water availability affects SCP in classically isohydric piñon pine and anisohydric one-seed or Utah juniper at various spatial- (i.e., from branch, to tree, to ecosystem) and temporal- (i.e., hours to decades) scales. Our results show that short-term increases in water availability decreased SCP in isohydric piñon pine, making it more anisohydric, while short-term rehydration had no effect on SCP in anisohydric juniper. Increasing mean annual precipitation, on the other hand, increased SCP in both species. Our findings are consistent with documented differences in the use of ABA to control stomata in iso- and aniso-hydric species on short timescales, and with structural acclimation in both species at long timescales. These results illustrate that the local environment plays a large role in determining SCP.

Kraklow, Vachel↗

Future climate doubles the risk of hydraulic failure in a wet tropical forest

Summary Future climate presents conflicting implications for forest biomass. We evaluate how plant hydraulic traits, elevated CO 2 levels, warming, and changes in precipitation affect forest primary productivity, evapotranspiration, and the risk of hydraulic failure. We used a dynamic vegetation model with plant hydrodynamics (FATES‐HYDRO) to simulate the stand‐level responses to future climate changes in a wet tropical forest in Barro Colorado Island, Panama. We calibrated the model by selecting plant trait assemblages that performed well against observations. These assemblages were run with temperature and precipitation changes for two greenhouse gas emission scenarios (2086–2100: SSP2‐45, SSP5‐85) and two CO 2 levels (contemporary, anticipated). The risk of hydraulic failure is projected to increase from a contemporary rate of 5.7% to 10.1–11.3% under future climate scenarios, and, crucially, elevated CO 2 provided only slight amelioration. By contrast, elevated CO 2 mitigated GPP reductions. We attribute a greater variation in hydraulic failure risk to trait assemblages than to either CO 2 or climate. Our results project forests with both faster growth (through productivity increases) and higher mortality rates (through increasing rates of hydraulic failure) in the neo‐tropics accompanied by certain trait plant assemblages becoming nonviable.

54 ENVIRONMENTAL SCIENCES↗

Benefits of symbiotic ectomycorrhizal fungi to plant water relations depend on plant genotype in pinyon pine

Rhizosphere microbes, such as root-associated fungi, can improve plant access to soil resources, affecting plant health, productivity, and stress tolerance. While mycorrhizal associations are ubiquitous, plant–microbe interactions can be species specific. Here we show that the specificity of the effects of microbial symbionts on plant function can go beyond species level: colonization of roots by ectomycorrhizal fungi (EMF) of the genus Geopora has opposite effects on water uptake, and stomatal control of desiccation in drought tolerant and intolerant genotypes of pinyon pine (Pinus edulis Engelm.). These results demonstrate, for the first time, that microorganisms can have significant and opposite effects on important plant functional traits like stomatal control of desiccation that are associated with differential mortality and growth in nature. They also highlight that appropriate pairing of plant genotypes and microbial associates will be important for mitigating climate change impacts on vegetation.

59 BASIC BIOLOGICAL SCIENCES↗

No carbon storage in growth-limited trees in a semi-arid woodland

Plant survival depends on a balance between carbon supply and demand. When carbon supply becomes limited, plants buffer demand by using stored carbohydrates (sugar and starch). During drought, NSCs (non-structural carbohydrates) may accumulate if growth stops before photosynthesis. This expectation is pervasive, yet few studies have combined simultaneous measurements of drought, photosynthesis, growth, and carbon storage to test this. Using a field experiment with mature trees in a semi-arid woodland, we show that growth and photosynthesis slow in parallel as ψ pd declines, preventing carbon storage in two species of conifer (J. monosperma and P. edulis). During experimental drought, growth and photosynthesis were frequently co-limited. Our results point to an alternative perspective on how plants use carbon that views growth and photosynthesis as independent processes both regulated by water availability.

54 ENVIRONMENTAL SCIENCES↗

Integrating plant physiology into simulation of fire behavior and effects

Summary Wildfires are a global crisis, but current fire models fail to capture vegetation response to changing climate. With drought and elevated temperature increasing the importance of vegetation dynamics to fire behavior, and the advent of next generation models capable of capturing increasingly complex physical processes, we provide a renewed focus on representation of woody vegetation in fire models. Currently, the most advanced representations of fire behavior and biophysical fire effects are found in distinct classes of fine‐scale models and do not capture variation in live fuel (i.e. living plant) properties. We demonstrate that plant water and carbon dynamics, which influence combustion and heat transfer into the plant and often dictate plant survival, provide the mechanistic linkage between fire behavior and effects. Our conceptual framework linking remotely sensed estimates of plant water and carbon to fine‐scale models of fire behavior and effects could be a critical first step toward improving the fidelity of the coarse scale models that are now relied upon for global fire forecasting. This process‐based approach will be essential to capturing the influence of physiological responses to drought and warming on live fuel conditions, strengthening the science needed to guide fire managers in an uncertain future.

54 ENVIRONMENTAL SCIENCES↗

Leaves as bottlenecks: The contribution of tree leaves to hydraulic resistance within the soil–plant–atmosphere continuum

Within vascular plants, the partitioning of hydraulic resistance along the soil-to-leaf continuum affects transpiration and its response to environmental conditions. In trees, the fractional contribution of leaf hydraulic resistance (R leaf ) to total soil-to-leaf hydraulic resistance (R total ), or fR leaf (= R leaf /R total ), is thought to be large, but this has not been tested comprehensively. Here, we compiled a multi-biome dataset of fR leaf using new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fR leaf averaged 0.51 (95% confidence interval [CI] = 0.46-0.57) and it declined with tree height. We also used the allometric relationship between field-based measurements of soil-to-leaf hydraulic conductance and laboratory-based measurements of leaf hydraulic conductance to compute the average fR leaf for 19 tree samples, which was 0.40 (95% CI = 0.29-0.56). The in-situ technique produces a more accurate descriptor of fR leaf because it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fR leaf may play a role in protecting stems from loss of hydraulic conductance. Thus, the decline in fR leaf with tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality.

54 ENVIRONMENTAL SCIENCES↗

Hydraulic architecture explains species moisture dependency but not mortality rates across a tropical rainfall gradient

Aim Intensified droughts are affecting tropical forests across the globe. However, the underlying mechanisms of tree drought response and mortality are poorly understood. Hydraulic traits and especially hydraulic safety margins (HSMs), i.e. the extent to which plants buffer themselves from thresholds of water stress, provide insights into species-specific drought vulnerability. Methods We investigated hydraulic traits during an intense drought triggered by the 2015-2016 El Niño on 27 canopy trees across three tropical forest sites with differing precipitation. We capitalized on the drought event as a time when plant water status might approach or exceed thresholds of water stress. We investigated the degree to which these traits varied across the rainfall gradient, as well as relationships amongst hydraulic traits and species-specific optimal moisture and mortality rates. Results There were no differences among sites for any measured trait. There was strong coordination among traits, with a network analysis revealing two major groups of coordinated traits. In one group there were water potentials, turgor loss point, sapwood capacitance and density, HSMs, and mortality rate. In the second group there was leaf mass per area, leaf dry matter content, hydraulic architecture (leaf area to sapwood area ratio), and species-specific optimal moisture. Conclusion These results demonstrated that while species with greater safety from turgor loss had lower mortality rates, hydraulic architecture was the only trait that explained species’ moisture dependency. Species with a greater leaf area to sapwood area ratio were associated with drier sites and reduced their transpirational demand during the dry season via deciduousness.

Hydraulic Saftey Margins, drought, tropical forest↗

The response of stomatal conductance to seasonal drought in tropical forests

Stomata regulate CO2 uptake for photosynthesis and water loss through transpiration. The approaches used to represent stomatal conductance (gs) in models vary. In particular, current understanding of drivers of the variation in a key parameter in those models, the slope parameter (i.e. a measure of intrinsic plant water-use-efficiency), is still limited, particularly in the tropics. Here we collected diurnal measurements of leaf gas exchange and water potential (?leaf), and a suite of plant traits from the upper canopy of 15 tropical trees in two contrasting Panamanian forests throughout the dry season of the 2016 El Niño. The plant traits included wood density, leaf-mass-per-area (LMA), leaf carboxylation capacity (Vc,max25), leaf water content, the degree of isohydry, and predawn ?leaf. We first investigated how the choice of four commonly used leaf-level gs models with and without the inclusion of ?leaf as an additional predictor variable influence the ability to predict gs, and then explored the abiotic (i.e. month, site-month interaction) and biotic (i.e. tree-species-specific characteristics) drivers of slope parameter variation. Our results show that the inclusion of ?leaf did not improve model performance and that the models that represent the response of gs to vapor pressure deficit performed better than corresponding models that respond to relative humidity. Within each gs model, we found large variation in the slope parameter, and this variation was attributable to the biotic driver, rather than abiotic drivers. We further investigated potential relationships between the slope parameter and the six available plant traits mentioned above, and found that only one trait, LMA, had a significant correlation with the slope parameter (R2=0.66, n=15), highlighting a potential path towards improved model parameterization. This study advances understanding of gs dynamics over seasonal drought, and identifies a practical, trait-based approach to improve modeling of carbon and water exchange in tropical forests.

carbon and water exchange, stomatal conductance mo↗

Homeostatic maintenance of nonstructural carbohydrates during the 2015-2016 El Niño drought across a tropical forest precipitation gradient

Non-structural carbohydrates (NSCs) are essential for maintenance of plant metabolism, and may be sensitive to both short- and long-term climatic variation. The NSC variation in NSCs in moist tropical forests has only rarely been studied, however, so regulation of NSCs in these systems is poorly understood. We measured foliar and branch NSC content in 23 tree species at three sites located across a large precipitation gradient in Panama during the 2015-2016 El Niño to examine how short- and long-term climatic variation impacts carbohydrate dynamics. Across all sites, leaf NSCs increased over diurnal time-periods. There was no significant difference in total NSCs over as the progression of the drought progressed (leaf p=0.32, branch p=0.30), nor across the rainfall gradient (leaf p=0.91, branch p=0.96). The Foliar soluble sugars decreased while starch increased :starch ratio did decrease over the duration of the dry period, suggesting greater partitioning of NSCs to storage than metabolism or transport as drought progressed. There was large variation across species at all sites, but total foliar NSCs were positively correlated with leaf mass per area, while branch sugars were positively related to leaf temperature and negatively correlated with daily photosynthesis and wood density. The NSC homeostasis across a wide range of conditions suggests that NSCs are an allocation priority in moist tropical forests.

Dickman, Lee T.↗

Mechanisms of a coniferous woodland persistence under drought and heat

Predictions of warmer droughts causing increasing forest mortality are becoming abundant, yet fewer studies have investigated the mechanisms of forest persistence. To examine the resistance of forests to warmer droughts, we used a five-year precipitation reduction (~45% removal), heat (+4°C above ambient) and combined drought and heat experiment in an isolated stand of mature Pinus edulis-Juniperus monosperma . Despite severe experimental drought and heating, no trees died, and we observed only minor evidence of hydraulic failure or carbon starvation. Two mechanisms promoting survival were supported. First, access to bedrock water, or ‘hydraulic refugia’ aided trees in their resistance to the experimental conditions. Second, the isolation of this stand amongst a landscape of dead trees precluded ingress by Ips confusus , frequently the ultimate biotic mortality agent of piñon. These combined abiotic and biotic landscape-scale processes can moderate the impacts of future droughts on tree mortality by enabling tree avoidance of hydraulic failure, carbon starvation, and exposure to attacking abiotic agents.

54 ENVIRONMENTAL SCIENCES↗

Standardized protocols and procedures can precisely and accurately quantify non-structural carbohydrates

Non-structural carbohydrates (NSCs), the stored products of photosynthesis, building blocks for growth, and fuel for respiration, are central to plant metabolism, but their measurement is challenging. Differences in methods and procedures among laboratories can cause results to vary widely, limiting our ability to integrate and generalize patterns in plant C balance among studies. A recent assessment found that NSC concentrations measured for a common set of samples can vary by an order of magnitude, but sources for this variability were unclear. We measured a common set of nine plant material types, and two synthetic samples with known NSC concentrations, using a common protocol for sugar extraction and starch digestion, and three different sugar quantification methods (ion chromatography, enzyme, acid) in six laboratories. We also tested how sample handling, extraction solvent, and centralizing parts of the procedure in one laboratory affected results. NSC concentrations measured for synthetic samples were within about 11.5% of known values for all three methods. However, differences among quantification methods were the largest source of variation in NSC measurements for natural plant samples because the three methods quantify different NSCs. The enzyme method quantified only glucose, fructose, and sucrose, with ion chromatography we additionally quantified galactose, while the acid method quantified a large range of mono- and oligosaccharides. For some natural samples, acid method quantification doubled sugar relative to the other methods, demonstrating that trees allocate C to a range of sugar molecules. Sample handling had little effect on measurements, while ethanol sugar extraction improved accuracy over water extraction. Our results demonstrate that reasonable accuracy of NSC measurements can be achieved when different methods are used, as long as protocols are robust and standardized. Thus, we provide detailed protocols for the extraction, digestion and quantification of NSCs in plant samples, which should improve the comparability of NSC measurements among laboratories.

Landhäusser, Simon M.↗