Tradeoffs between leaf cooling and hydraulic safety in a dominant arid land riparian tree species.

Leaf carbon gain optimization in hot environments requires balancing leaf thermoregulation with avoiding excessive water loss via transpiration and hydraulic failure. The tradeoffs between leaf thermoregulation and transpirational water loss can determine the ecological consequences of heat waves that are increasing in frequency and intensity. We evaluated leaf thermoregulation strategies in warm- (>40°C maximum summer temperature) and cool-adapted (<40°C maximum summer temperature) genotypes of the foundation tree species, Populus fremontii, using a common garden near the mid-elevational point of its distribution. We measured leaf temperatures and assessed three modes of leaf thermoregulation: leaf morphology, midday canopy stomatal conductance and stomatal sensitivity to vapour pressure deficit. Data were used to parameterize a leaf energy balance model to estimate contrasts in midday leaf temperature in warm- and cool-adapted genotypes. Warm-adapted genotypes had 39% smaller leaves and 38% higher midday stomatal conductance, reflecting a 3.8°C cooler mean leaf temperature than cool-adapted genotypes. Leaf temperatures modelled over the warmest months were on average 1.1°C cooler in warm- relative to cool-adapted genotypes. Results show that plants adapted to warm environments are predisposed to tightly regulate leaf temperatures during heat waves, potentially at an increased risk of hydraulic failure.

Episodic defoliation rapidly reduces starch but not soluble sugars in an invasive shrub, Tamarix spp.

PREMISE: Plants rely on pools of internal nonstructural carbohydrates (NSCs: soluble sugars plus starch) to support metabolism, growth, and regrowth of tissues damaged from disturbance such as foliage herbivory. However, impacts of foliage herbivory on the quantity and composition of NSC pools in long-lived woody plants are currently unclear. We implemented a controlled defoliation experiment on mature Tamarix spp.-a dominant riparian woody shrub/tree that has evolved with intense herbivory pressure-to test two interrelated hypotheses: (1) Repeated defoliation disproportionately impacts aboveground versus belowground NSC storage. (2) Defoliation disproportionately impacts starch versus soluble sugar storage. METHODS: Hypotheses were tested by transplanting six Tamarix seedlings into each of eight cylinder mesocosms (2 m diameter, 1 m in depth). After 2.5 years, plants in four of the eight mesocosms were mechanically defoliated repeatedly over a single growing season, and all plants were harvested in the following spring. RESULTS: Defoliation had no impact on either above- or belowground soluble sugar pools. However, starch in defoliated plants dropped to 55% and 26% in stems and roots, respectively, relative to control plants, resulting in an over 2-fold higher soluble sugar to starch ratio in defoliated plants. CONCLUSIONS: The results suggest that defoliation occurring over a single growing season does not impact immediate plant functions such as osmoregulation, but depleted starch could limit future fitness, particularly where defoliation occurs over multiple years. These results improve our understanding of how woody plants cope with episodic defoliation caused by foliage herbivory and other disturbances.

Does sexual dimorphism predispose dioecious riparian trees to sex ratio imbalances under climate change?

Environmental changes have resulted in significant declines in native riparian forests that are comprised largely of dioecious tree taxa, including boxelder and iconic cottonwood/willow gallery forests. Dioecious species may be especially vulnerable to the effects of climate change given that they often exhibit skewed sex ratios that are reinforced by physiological and morphological specialization of each sex to specific microhabitats. A comprehensive data synthesis suggests that male individuals of boxelder and cottonwood taxa have a higher representation on dry microhabitats than females and are less physiologically sensitive to increased aridity than co-occurring females. Consequently, extreme male-biased sex ratios are possible under future climate conditions that could reduce population fitness below a sustainable threshold. Riparian willows, on the other hand, generally do not express obvious sexual dimorphism in habitat preference or physiological sensitivity to aridity. Thus, it is unclear whether climate change will impact population structure of willows in ways that parallel other dioecious riparian tree taxa. Future riparian tree restoration programs should aim to maintain future sex ratio balance that maximizes population fitness under projected hydro-climatological conditions. Recent advances in genomics will likely provide the critical tools for early sex determination in pre-reproductive trees across riparian tree species such that sex ratio balance could be targeted during initial stages of restoration, along with adaptations for drought tolerance and other key traits that are essential for survival under future conditions.

Long-term urban carbon dioxide observations reveal spatial and temporal dynamics related to urban characteristics and growth.

Cities are concentrated areas of CO2 emissions and have become the foci of policies for mitigation actions. However, atmospheric measurement networks suitable for evaluating urban emissions over time are scarce. Here we present a unique long-term (decadal) record of CO2 mole fractions from five sites across Utah's metropolitan Salt Lake Valley. We examine "excess" CO2 above background conditions resulting from local emissions and meteorological conditions. We ascribe CO2 trends to changes in emissions, since we did not find long-term trends in atmospheric mixing proxies. Three contrasting CO2 trends emerged across urban types: negative trends at a residential-industrial site, positive trends at a site surrounded by rapid suburban growth, and relatively constant CO2 over time at multiple sites in the established, residential, and commercial urban core. Analysis of population within the atmospheric footprints of the different sites reveals approximately equal increases in population influencing the observed CO2, implying a nonlinear relationship with CO2 emissions: Population growth in rural areas that experienced suburban development was associated with increasing emissions while population growth in the developed urban core was associated with stable emissions. Four state-of-the-art global-scale emission inventories also have a nonlinear relationship with population density across the city; however, in contrast to our observations, they all have nearly constant emissions over time. Our results indicate that decadal scale changes in urban CO2 emissions are detectable through monitoring networks and constitute a valuable approach to evaluate emission inventories and studies of urban carbon cycles.

Can local adaptation explain varying patterns of herbivory tolerance in a recently introduced woody plant in North America?

Patterns of woody-plant mortality have been linked to global-scale environmental changes, such as extreme drought, heat stress, more frequent and intense fires, and episodic outbreaks of insects and pathogens. Although many studies have focussed on survival and mortality in response to specific physiological stresses, little attention has been paid to the role of genetic heritability of traits and local adaptation in influencing patterns of plant mortality, especially in non-native species. Tamarix spp. is a dominant, non-native riparian tree in western North America that is experiencing dieback in some areas of its range due to episodic herbivory by the recently introduced northern tamarisk leaf beetle (Diorhabda carinulata). We propose that genotype × environment interactions largely underpin current and future patterns of Tamarix mortality. We anticipate that (i) despite its recent introduction, and the potential for significant gene flow, Tamarix in western North America is generally adapted to local environmental conditions across its current range in part due to hybridization of two species; (ii) local adaptation to specific climate, soil and resource availability will yield predictable responses to episodic herbivory; and (iii) the ability to cope with a combination of episodic herbivory and increased aridity associated with climate change will be largely based on functional tradeoffs in resource allocation. This review focusses on the potential heritability of plant carbon allocation patterns in Tamarix, focussing on the relative contribution of acquired carbon to non-structural carbohydrate (NSC) pools versus other sinks as the basis for surviving episodic disturbance. Where high aridity and/or poor edaphic position lead to chronic stress, NSC pools may fall below a minimum threshold because of an imbalance between the supply of carbon and its demand by various sinks. Identifying patterns of local adaptation of traits related to resource allocation will improve forecasting of Tamarix population susceptibility to episodic herbivory.

Calibration of thermal dissipation sap flow probes for ring- and diffuse-porous trees.

Thermal dissipation probes (the Granier method) are routinely used in forest ecology and water balance studies to estimate whole-tree transpiration. This method utilizes an empirically derived equation to measure sap flux density, which has been reported as independent of wood characteristics. However, errors in calculated sap flux density may occur when large gradients in sap velocity occur along the sensor length or when sensors are inserted into non-conducting wood. These may be conditions routinely associated with ring-porous species, yet there are few cases in which the original calibration has been validated for ring-porous species. We report results from laboratory calibration measurements conducted on excised stems of four ring-porous species and two diffuse-porous species. Our calibration results for ring-porous species were considerably different compared with the original calibration equation. Calibration equation coefficients obtained in this study differed by as much as two to almost three orders of magnitude when compared with the original equation of Granier. Coefficients also differed between ring-porous species across all pressure gradient conditions considered; however, no differences between calibration slopes were observed for data collected within the range of expected in situ pressure gradients. In addition, dye perfusions showed that in three of the four ring-porous species considered, active sapwood was limited to the outermost growth ring. In contrast, our calibration results for diffuse-porous species showed generally good agreement with the empirically derived Granier calibration, and dye perfusions showed that active sapwood was associated with many annual growth rings. Our results suggest that the original calibration of Granier is not universally applicable to all species and xylem types and that previous estimates of absolute rates of water use for ring-porous species obtained using the original calibration coefficients may be associated with substantial error.

Wood anatomy constrains stomatal responses to atmospheric vapor pressure deficit in irrigated, urban trees.

Plant transpiration is strongly constrained by hydraulic architecture, which determines the critical threshold for cavitation. Because species vary greatly in vulnerability to cavitation, hydraulic limits to transpiration and stomatal conductance have not generally been incorporated into ecological and climate models. We measured sap flow, leaf transpiration, and vulnerability to cavitation of a variety of tree species in a well-irrigated but semi-arid urban environment in order to evaluate the generality of stomatal responses to high atmospheric vapor pressure deficit (D). We found evidence of broad patterns of stomatal responses to humidity based on systematic differences in vulnerability to cavitation. Ring-porous taxa consistently had vulnerable xylem and showed strong regulation of transpiration in response to D, while diffuse-porous taxa were less vulnerable and transpiration increased nearly linearly with D. These results correspond well to patterns in the distribution of the taxa, such as the prevalence of diffuse-porous species in riparian ecosystems, and also provide a means of representing maximum transpiration rates at varying D in broad categories of trees.

Evaluation of centrifugal methods for measuring xylem cavitation in conifers, diffuse- and ring-porous angiosperms.

A centrifugal method is used to measure 'vulnerability curves' which show the loss of hydraulic conductivity in xylem by cavitation. Until recently, conductivity was measured between bouts of centrifugation using a gravity-induced head. Now, conductivity can be measured during centrifugation. This 'spin' method is faster than the 'gravity' technique, but correspondence between the two has not been evaluated. The two methods were compared on the same stem segments for two conifer, four diffuse-porous, and four ring-porous species. Only 17 of 60 conductivity measurements differed, with differences in the order of 10%. When different, the spin method gave higher conductivities at the beginning of the curve and lower at the end. Pressure at 50% loss of conductivity, and mean cavitation pressure, were the same in 14 of 20 comparisons. When different, the spin method averaged 0.32 MPa less negative. Ring-porous species showed a precipitous initial drop in conductivity by both techniques. This striking pattern was confirmed by the air-injection method and native embolism measurements. Close correspondence inspires confidence in both methods, each of which has unique advantages. The observation that ring-porous species operate at only a fraction of their potential conductivity at midday demands further study.