The poorly-explored stomatal response to temperature at constant evaporative demand.

Changes in leaf temperature are known to drive stomatal responses, because the leaf-to-air water vapour gradient (Δw) increases with temperature if ambient vapour pressure is held constant, and stomata respond to changes in Δw. However, the direct response of stomata to temperature (DRST; the response when Δw is held constant by adjusting ambient humidity) has been examined far less extensively. Though the meagre available data suggest the response is usually positive, results differ widely and defy broad generalisation. As a result, little is known about the DRST. This review discusses the current state of knowledge about the DRST, including numerous hypothesised biophysical mechanisms, potential implications of the response for plant adaptation, and possible impacts of the DRST on plant-atmosphere carbon and water exchange in a changing climate.

Grape cultivars adapted to hotter, drier growing regions exhibit greater photosynthesis in hot conditions despite less drought-resistant leaves.

BACKGROUND AND AIMS: Many agricultural areas are expected to face hotter, drier conditions from climate change. Understanding the mechanisms that crops use to mitigate these stresses can guide breeding for more tolerant plant material. We tested relationships between traits, physiological function in hot conditions and historical climate associations to evaluate these mechanisms for winegrapes. We expected a more negative leaf osmotic potential at full hydration (πo), which reduces leaf turgor loss during drought, and either a metabolically cheaper or more osmoprotectant leaf chemical composition, to allow cultivars associated with hot, dry regions to maintain greater gas exchange in hot growing conditions. METHODS: We measured πo, gas exchange and leaf chemistry for seven commercially important winegrape cultivars that vary widely in historical climate associations. Vines were grown in common-garden field conditions in a hot wine-growing region (Davis, CA, USA) and measured over the hottest period of the growing season (July-September). KEY RESULTS: The value of πo varied significantly between cultivars, and all cultivars significantly reduced πo (osmotically adjusted) over the study period, although osmotic adjustment did not vary across cultivars. The value of πo was correlated with gas exchange and climate associations, but in the direction opposite to expected. Photosynthesis and πo were higher in the cultivars associated with hotter, less humid regions. Leaf chemical composition varied between cultivars but was not related to climate associations. CONCLUSIONS: These findings suggest that maintenance of leaf turgor is not a primary limitation on grapevine adaptation to hot or atmospherically dry growing conditions. Thus, selecting for a more negative πo or greater osmotic adjustment is not a promising strategy to develop more climate-resilient grape varieties, contrary to findings for other crops. Future work is needed to identify the mechanisms increasing photosynthesis in the cultivars associated with hot, dry regions.

Thresholds for persistent leaf photochemical damage predict plant drought resilience in a tropical rainforest.

Water stress can cause declines in plant function that persist after rehydration. Recent work has defined 'resilience' traits characterizing leaf resistance to persistent damage from drought, but whether these traits predict resilience in whole-plant function is unknown. It is also unknown whether the coordination between resilience and 'resistance' - the ability to maintain function during drought - observed globally occurs within ecosystems. For eight rainforest species, we dehydrated and subsequently rehydrated leaves, and measured water stress thresholds for declines in rehydration capacity and maximum quantum yield of photosystem II (Fv /Fm ). We tested correlations with embolism resistance and dry season water potentials (ΨMD ), and calculated safety margins for damage (ΨMD - thresholds) and tested correlations with drought resilience in sap flow and growth. Ψ thresholds for persistent declines in Fv /Fm , indicating resilience, were positively correlated with ΨMD and thresholds for leaf vein embolism. Safety margins for persistent declines in Fv /Fm , but not rehydration capacity, were positively correlated with drought resilience in sap flow. Correlations between resistance and resilience suggest that species' differences in performance during drought are perpetuated after drought, potentially accelerating shifts in forest composition. Resilience to photochemical damage emerged as a promising functional trait to characterize whole-plant drought resilience.

Low baseline intraspecific variation in leaf pressure-volume traits: Biophysical basis and implications for spectroscopic sensing.

Intra-specific trait variation (ITV) plays a role in processes at a wide range of scales from organs to ecosystems across climate gradients. Yet, ITV remains rarely quantified for many ecophysiological traits typically assessed for species means, such as pressure volume (PV) curve parameters including osmotic potential at full turgor and modulus of elasticity, which are important in plant water relations. We defined a baseline "reference ITV" (ITVref ) as the variation among fully exposed, mature sun leaves of replicate individuals of a given species grown in similar, well-watered conditions, representing the conservative sampling design commonly used for species-level ecophysiological traits. We hypothesized that PV parameters would show low ITVref relative to other leaf morphological traits, and that their intraspecific relationships would be similar to those previously established across species and proposed to arise from biophysical constraints. In a database of novel and published PV curves and additional leaf structural traits for 50 diverse species, we found low ITVref for PV parameters relative to other morphological traits, and strong intraspecific relationships among PV traits. Simulation modeling showed that conservative ITVref enables the use of species-mean PV parameters for scaling up from spectroscopic measurements of leaf water content to enable sensing of leaf water potential.

Measuring the Efficiency of Automation-Aided Performance in a Simulated Baggage Screening Task.

OBJECTIVE: The present study replicated and extended prior findings of suboptimal automation use in a signal detection task, benchmarking automation-aided performance to the predictions of several statistical models of collaborative decision making. BACKGROUND: Though automated decision aids can assist human operators to perform complex tasks, operators often use the aids suboptimally, achieving performance lower than statistically ideal. METHOD: Participants performed a simulated security screening task requiring them to judge whether a target (a knife) was present or absent in a series of colored X-ray images of passenger baggage. They completed the task both with and without assistance from a 93%-reliable automated decision aid that provided a binary text diagnosis. A series of three experiments varied task characteristics including the timing of the aid's judgment relative to the raw stimuli, target certainty, and target prevalence. RESULTS AND CONCLUSION: Automation-aided performance fell closest to the predictions of the most suboptimal model under consideration, one which assumes the participant defers to the aid's diagnosis with a probability of 50%. Performance was similar across experiments. APPLICATION: Results suggest that human operators' performance when undertaking a naturalistic search task falls far short of optimal and far lower than prior findings using an abstract signal detection task.

Predicting Stomatal Closure and Turgor Loss in Woody Plants Using Predawn and Midday Water Potential.

Knowledge about physiological stress thresholds provides crucial information about plant performance and survival under drought. In this study, we report on the triphasic nature of the relationship between plant water potential (Ψ) at predawn and midday and describe a method that predicts Ψ at stomatal closure and turgor loss exclusively from this water potential curve (WP curve). The method is based on a piecewise linear regression model that was developed to predict the boundaries (termed Θ1 and Θ2) separating the three phases of the curve and corresponding slope values. The method was tested for three economically important woody species. For all species, midday Ψ was much more negative than predawn Ψ during phase I (mild drought), reductions in midday Ψ were minor while predawn Ψ continued to decline during phase II (moderate drought), and midday and predawn Ψ reached similar values during phase III (severe drought). Corresponding measurement of leaf gas exchange indicated that boundary Θ1 between phases I and II coincided with Ψ at stomatal closure. Data from pressure-volume curves demonstrated that boundary Θ2 between phases II and III predicted Ψ at leaf turgor loss. The WP curve method described here is an advanced application of the Scholander-type pressure chamber to categorize plant dehydration under drought into three distinct phases and to predict Ψ thresholds of stomatal closure and turgor loss.

Temperature and evaporative demand drive variation in stomatal and hydraulic traits across grape cultivars.

Selection for crop cultivars has largely focused on reproductive traits, while the impacts of global change on crop productivity are expected to depend strongly on the vegetative physiology traits that drive plant resource use and stress tolerance. We evaluated relationships between physiology traits and growing season climate across wine grape cultivars to characterize trait variation across European growing regions. We compiled values from the literature for seven water use and drought tolerance traits and growing season climate. Cultivars with a lower maximum stomatal conductance were associated with regions with a higher mean temperature and mean and maximum vapor pressure deficit (r2=0.39-0.65, P<0.05, n=14-29). Cultivars with greater stem embolism resistance and more anisohydric stomatal behavior (i.e. a more negative water potential threshold for 50% stomatal closure) were associated with cooler regions (r2=0.48-0.72, P<0.03, n=10-29). Overall, cultivars grown in warmer, drier regions exhibited traits that would reduce transpiration and conserve soil water longer into the growing season, but potentially increase stomatal and temperature limitations on photosynthesis under future, hotter conditions.

The effect of cognitive load on horizontal and vertical spatial asymmetries.

Healthy individuals typically show a leftward attentional bias in the allocation of spatial attention along the horizontal plane, a phenomenon known as pseudoneglect, which relies on a right hemispheric dominance for visuospatial processing. Also, healthy individuals tend to overestimate the upper hemispace when orienting attention along the vertical plane, a phenomenon that may depend on asymmetric ventral and dorsal visual streams activation. Previous research has demonstrated that when attentional resources are reduced due to increased cognitive load, pseudoneglect is attenuated (or even reversed), due to decreased right-hemispheric activations. Critically, whether and how the reduction of attentional resources under load modulates vertical spatial asymmetries has not been addressed before. We asked participants to perform a line bisection task both with and without the addition of a concurrent auditory working memory task with lines oriented either horizontally or vertically. Results showed that increasing cognitive load reduced the typical leftward/upward bias with no difference between orientations. Our data suggest that the degree of cognitive load affects spatial attention not only in the horizontal but also in the vertical plane. Lastly, the similar effect of load on horizontal and vertical judgements suggests these biases may be related to only partially independent mechanisms.

Ironic efficiency in automation-aided signal detection.

Decision makers often make poor use of the information provided by an automated signal detection aid; recent studies have found that participants assisted by an automated aid fell well short of best-possible sensitivity levels. The present study tested the generalisability of this finding over varying levels of aid reliability. Participants performed a binary signal detection task either unaided or with assistance from a decision aid that was 60%, 85%, or 96%-reliable. Assistance from a highly reliable aid (85% or 96%) improved discrimination performance, while assistance from a low-reliability aid (60%) did not. Because their ideal strategy is to place less weight on less reliable cues, however, the decision makers' tendency to disuse the aid became more appropriate as the aid's reliability declined. Automation-aided efficiency was thus near to optimal when the aid was close to chance but became highly inefficient, ironically, as the aid's reliability increased. Practitioner Summary: Investigating operators' automation-aided information integration strategies allows human factors practitioners to predict the level of performance the operator will attain. Ironically, in an aided signal detection task, performance when assisted by a highly reliable aid is far less efficient than that obtained when assisted by a far less reliable aid. Abbreviations: OW: optimal weighting; UW: uniform weighting; CC: contingent criterion; BD: best decides; CF: coin flip; PM: probability matching; HDI: highest density interval; MCMC: markov chain monte carlo; HR: hit rate; FAR: false alarm rate.

Cognitive load exacerbates rightward biases during computer maze navigation.

Neurologically healthy individuals exhibit subtle attentional asymmetries, such that attention is preferentially directed leftwards for objects in near space and rightwards for objects in far space. These attentional biases also affect navigation and cause people to deviate to the right when passing through an aperture. The current study examined whether the rightward deviations observed in real-world environments translate to simulated environments. As proof of concept and to determine whether rightward biases could be further exacerbated, the degree of cognitive load imposed on participants was manipulated. Experiment 1 asked participants to navigate through the centre of a computer-based doorway. In one block of trials, participants completed the task by itself (baseline condition), while in another block of trials they also completed a simple auditory discrimination task (load condition). While analyses revealed rightward biases for both conditions, the difference between conditions was not significant. Experiment 2 therefore increased the difficulty of the auditory task. Analyses revealed a significant difference between conditions, suggesting that the degree of cognitive load further exacerbates rightward biases, demonstrating that the rightward asymmetries in navigation observed in the real world generalises to a simulated environment and that this phenomenon behaves in a way that is consistent with pseudoneglect.

Flowers in the Attic: Lateralization of the detection of meaning in visual noise.

The brain is a slave to sense; we see and hear things that are not there and engage in ongoing correction of these illusory experiences, commonly termed pareidolia. The current study investigates whether the predisposition to see meaning in noise is lateralized to one hemisphere or the other and how this predisposition to visual false-alarms is related to personality. Stimuli consisted of images of faces or flowers embedded in pink (1/f) noise generated through a novel process and presented in a divided-field paradigm. Right-handed undergraduates participated in a forced-choice signal-detection task where they determined whether a face or flower signal was present in a single-interval trial. Experiment 1 involved an equal ratio of signal-to-noise trials; experiment 2 provided more potential for illusionary perception with 25% signal and 75% noise trials. There was no asymmetry in the ability to discriminate signal from noise trials (measured using d') for either faces and flowers, although the response criterion (c) suggested a stronger predisposition to visual false alarms in the right visual field, and this was negatively correlated to the unusual experiences dimension of schizotypy. Counter to expectations, changing the signal-image to noise-image proportion in Experiment 2 did not change the number of false alarms for either faces and flowers, although a stronger bias was seen to the right visual field; sensitivity remained the same in both hemifields but there was a moderate positive correlation between cognitive disorganization and the bias (c) for "flower" judgements. Overall, these results were consistent with a rapid evidence-accumulation process of the kind described by a diffusion decision model mediating the task lateralized to the left-hemisphere.

Predicting shifts in the functional composition of tropical forests under increased drought and CO2 from trade-offs among plant hydraulic traits.

Tropical forest responses are an important feedback on global change, but changes in forest composition with projected increases in CO2 and drought are highly uncertain. Here we determine shifts in the most competitive plant hydraulic strategy (the evolutionary stable strategy or ESS) from changes in CO2 and drought frequency and intensity. Hydraulic strategies were defined along a spectrum from drought avoidance to tolerance by physiology traits. Drought impacted competition more than CO2 , with elevated CO2 reducing but not reversing drought-induced shifts in the ESS towards more tolerant strategies. Trait plasticity and/or adaptation intensified these shifts by increasing the competitive ability of the drought tolerant relative to the avoidant strategies. These findings predict losses of drought avoidant evergreens from tropical forests under global change, and point to the importance of changes in precipitation during the dry season and constraints on plasticity and adaptation in xylem traits to forest responses.

Seedling response to water stress in valley oak (Quercus lobata) is shaped by different gene networks across populations.

Drought is a major stress for plants, creating a strong selection pressure for traits that enable plant growth and survival in dry environments. Many drought responses are conserved species-wide responses, while others vary among populations distributed across heterogeneous environments. We tested how six populations of the widely distributed California valley oak (Quercus lobata) sampled from contrasting climates would differ in their response to soil drying relative to well-watered controls in a common environment by measuring ecophysiological traits in 93 individuals and gene expression (RNA-seq) in 42 individuals. Populations did not differ in their adjustment of turgor loss point during soil drying, suggesting a generalized species-wide response. Differential expression analysis identified 689 genes with a common response to treatment across populations and 470 genes with population-specific responses. Weighted gene co-expression network analysis (WGCNA) identified groups of genes with similar expression patterns that may be regulated together (gene modules). Several gene modules responded differently to water stress among populations, suggesting regional differences in gene network regulation. Populations from sites with a high mean annual temperature responded to the imposed water stress with significantly greater changes in gene module expression, indicating that these populations may be locally adapted to respond to drought. We propose that this variation among valley oak populations provides a mechanism for differential tolerance to the increasingly frequent and severe droughts in California.

The Causes of Leaf Hydraulic Vulnerability and Its Influence on Gas Exchange in Arabidopsis thaliana.

The influence of the dynamics of leaf hydraulic conductance (K leaf) diurnally and during dehydration on stomatal conductance and photosynthesis remains unclear. Using the model species Arabidopsis (Arabidopsis thaliana ecotype Columbia-0), we applied a multitiered approach including physiological measurements, high-resolution x-ray microcomputed tomography, and modeling at a range of scales to characterize (1) K leaf decline during dehydration; (2) its basis in the hydraulic conductances of leaf xylem and outside-xylem pathways (K ox); (3) the dependence of its dynamics on irradiance; (4) its impact on diurnal patterns of stomatal conductance and photosynthetic rate; and (5) its influence on gas exchange and survival under simulated drought regimes. Arabidopsis leaves showed strong vulnerability to dehydration diurnally in both gas exchange and hydraulic conductance, despite lack of xylem embolism or conduit collapse above the turgor loss point, indicating a pronounced sensitivity of K ox to dehydration. K leaf increased under higher irradiance in well-hydrated leaves across the full range of water potential, but no shift in K leaf vulnerability was observed. Modeling indicated that responses to dehydration and irradiance are likely attributable to changes in membrane permeability and that a dynamic K ox would contribute strongly to stomatal closure, improving performance, survival, and efficient water use during drought. These findings for Columbia-0 provide a baseline for assessing variation across genotypes in hydraulic traits and their influence on gas exchange during dehydration.

Disentangling the functional trait correlates of spatial aggregation in tropical forest trees.

Environmental filtering and dispersal limitation can both maintain diversity in plant communities by aggregating conspecifics, but parsing the contribution of each process has proven difficult empirically. Here, we assess the contribution of filtering and dispersal limitation to the spatial aggregation patterns of 456 tree species in a hyperdiverse Amazonian forest and find distinct functional trait correlates of interspecific variation in these processes. Spatial point process model analysis revealed that both mechanisms are important drivers of intraspecific aggregation for the majority of species. Leaf drought tolerance was correlated with species topographic distributions in this aseasonal rainforest, showing that future increases in drought severity could significantly impact community structure. In addition, seed mass was associated with the spatial scale and density of dispersal-related aggregation. Taken together, these results suggest environmental filtering and dispersal limitation act in concert to influence the spatial and functional structure of diverse forest communities.

Climate and plant trait strategies determine tree carbon allocation to leaves and mediate future forest productivity.

Forest leaf area has enormous leverage on the carbon cycle because it mediates both forest productivity and resilience to climate extremes. Despite widespread evidence that trees are capable of adjusting to changes in environment across both space and time through modifying carbon allocation to leaves, many vegetation models use fixed carbon allocation schemes independent of environment, which introduces large uncertainties into predictions of future forest responses to atmospheric CO2 fertilization and anthropogenic climate change. Here, we develop an optimization-based model, whereby tree carbon allocation to leaves is an emergent property of environment and plant hydraulic traits. Using a combination of meta-analysis, observational datasets, and model predictions, we find strong evidence that optimal hydraulic-carbon coupling explains observed patterns in leaf allocation across large environmental and CO2 concentration gradients. Furthermore, testing the sensitivity of leaf allocation strategy to a diversity in hydraulic and economic spectrum physiological traits, we show that plant hydraulic traits in particular have an enormous impact on the global change response of forest leaf area. Our results provide a rigorous theoretical underpinning for improving carbon cycle predictions through advancing model predictions of leaf area, and underscore that tree-level carbon allocation to leaves should be derived from first principles using mechanistic plant hydraulic processes in the next generation of vegetation models.

The correlations and sequence of plant stomatal, hydraulic, and wilting responses to drought.

Climate change is expected to exacerbate drought for many plants, making drought tolerance a key driver of species and ecosystem responses. Plant drought tolerance is determined by multiple traits, but the relationships among traits, either within individual plants or across species, have not been evaluated for general patterns across plant diversity. We synthesized the published data for stomatal closure, wilting, declines in hydraulic conductivity in the leaves, stems, and roots, and plant mortality for 262 woody angiosperm and 48 gymnosperm species. We evaluated the correlations among the drought tolerance traits across species, and the general sequence of water potential thresholds for these traits within individual plants. The trait correlations across species provide a framework for predicting plant responses to a wide range of water stress from one or two sampled traits, increasing the ability to rapidly characterize drought tolerance across diverse species. Analyzing these correlations also identified correlations among the leaf and stem hydraulic traits and the wilting point, or turgor loss point, beyond those expected from shared ancestry or independent associations with water stress alone. Further, on average, the angiosperm species generally exhibited a sequence of drought tolerance traits that is expected to limit severe tissue damage during drought, such as wilting and substantial stem embolism. This synthesis of the relationships among the drought tolerance traits provides crucial, empirically supported insight into representing variation in multiple traits in models of plant and ecosystem responses to drought.

Meta-analysis reveals that hydraulic traits explain cross-species patterns of drought-induced tree mortality across the globe.

Drought-induced tree mortality has been observed globally and is expected to increase under climate change scenarios, with large potential consequences for the terrestrial carbon sink. Predicting mortality across species is crucial for assessing the effects of climate extremes on forest community biodiversity, composition, and carbon sequestration. However, the physiological traits associated with elevated risk of mortality in diverse ecosystems remain unknown, although these traits could greatly improve understanding and prediction of tree mortality in forests. We performed a meta-analysis on species' mortality rates across 475 species from 33 studies around the globe to assess which traits determine a species' mortality risk. We found that species-specific mortality anomalies from community mortality rate in a given drought were associated with plant hydraulic traits. Across all species, mortality was best predicted by a low hydraulic safety margin-the difference between typical minimum xylem water potential and that causing xylem dysfunction-and xylem vulnerability to embolism. Angiosperms and gymnosperms experienced roughly equal mortality risks. Our results provide broad support for the hypothesis that hydraulic traits capture key mechanisms determining tree death and highlight that physiological traits can improve vegetation model prediction of tree mortality during climate extremes.

No Effect of Cue Format on Automation Dependence in an Aided Signal Detection Task.

OBJECTIVE: To investigate whether manipulating the format of an automated decision aid's cues can improve participants' information integration strategies in a signal detection task. BACKGROUND: Automation-aided decision making is often suboptimal, falling well short of statistically ideal levels. The choice of format in which the cues from the aid are displayed may help users to better understand and integrate the aid's judgments with their own. METHOD: Participants performed a signal detection task that asked them to classify random dot images as either blue or orange dominant. They made their judgments either unaided or with assistance from a 93% reliable automated decision aid. The aid provided a binary judgment, along with an estimate of signal strength in the form of either a raw value, a likelihood ratio, or a confidence rating (Experiments 1 and 2) or a binary judgment along with either a verbal or verbal-visuospatial expression of confidence (Experiment 3). Aided sensitivity was benchmarked to the predictions of various statistical models of collaborative decision making. RESULTS AND CONCLUSION: Aided performance was suboptimal, matching the predictions of some of the least efficient models. Most importantly, performance was similar across cue formats. APPLICATION: Results indicate that changes to the format in which cues from a signal detection aid are rendered are unlikely to dramatically improve the efficiency of automation-aided decision making.

Outside-Xylem Vulnerability, Not Xylem Embolism, Controls Leaf Hydraulic Decline during Dehydration.

Leaf hydraulic supply is crucial to maintaining open stomata for CO2 capture and plant growth. During drought-induced dehydration, the leaf hydraulic conductance (Kleaf) declines, which contributes to stomatal closure and, eventually, to leaf death. Previous studies have tended to attribute the decline of Kleaf to embolism in the leaf vein xylem. We visualized at high resolution and quantified experimentally the hydraulic vulnerability of xylem and outside-xylem pathways and modeled their respective influences on plant water transport. Evidence from all approaches indicated that the decline of Kleaf during dehydration arose first and foremost due to the vulnerability of outside-xylem tissues. In vivo x-ray microcomputed tomography of dehydrating leaves of four diverse angiosperm species showed that, at the turgor loss point, only small fractions of leaf vein xylem conduits were embolized, and substantial xylem embolism arose only under severe dehydration. Experiments on an expanded set of eight angiosperm species showed that outside-xylem hydraulic vulnerability explained 75% to 100% of Kleaf decline across the range of dehydration from mild water stress to beyond turgor loss point. Spatially explicit modeling of leaf water transport pointed to a role for reduced membrane conductivity consistent with published data for cells and tissues. Plant-scale modeling suggested that outside-xylem hydraulic vulnerability can protect the xylem from tensions that would induce embolism and disruption of water transport under mild to moderate soil and atmospheric droughts. These findings pinpoint outside-xylem tissues as a central locus for the control of leaf and plant water transport during progressive drought.