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Abstract Rising global temperatures and vapor pressure deficits (VPDs) are increasing plant water demand and becoming major drivers of large-scale plant mortality. Controlling transient leaf water loss after stomatal closure (minimum stomatal conductance [gmin]) is recognized as a key trait determining how long plants survive during soil drought. Yet, substantial uncertainty remains regarding how gmin responds to elevated temperatures and VPD and the underlying mechanisms. We measured gmin in 24 Quercus species from temperate and Mediterranean climates to determine whether gmin was sensitive to a coupled temperature and VPD increase. We also explored mechanistic links to phenology, climate, evolutionary history, and leaf anatomy. We found that gmin in all species exhibited a nonlinear negative temperature and VPD dependence. At 25 °C (VPD = 2.2 kPa), gmin varied from 1.19 to 8.09 mmol m−2 s−1 across species but converged to 0.57 ± 0.06 mmol m−2 s−1 at 45 °C (VPD = 6.6 kPa). In a subset of species, the effect of temperature and VPD on gmin was reversible and linked to the degree of stomatal closure, which was greater at 45 °C than at 25 °C. Our results show that gmin is dependent on temperature and VPD, is highly conserved in Quercus species, and is linked to leaf anatomy and stomatal behavior.more » « less
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Sensitive Hydraulic and Stomatal Decline in Extreme Drought Tolerant Species of California CeanothusABSTRACT Identifying the physiological mechanisms by which plants are adapted to drought is critical to predict species responses to climate change. We measured the responses of leaf hydraulic and stomatal conductances (Kleafandgs, respectively) to dehydration, and their association with anatomy, in seven species of CaliforniaCeanothusgrown in a common garden, including some of the most drought‐tolerant species in the semi‐arid flora. We tested for matching of maximum hydraulic supply and demand and quantified the role of decline ofKleafin driving stomatal closure. AcrossCeanothusspecies, maximumKleafandgswere negatively correlated, and bothKleafandgsshowed steep declines with decreasing leaf water potential (i.e., a high sensitivity to dehydration). The leaf water potential at 50% decline ingswas linked with a low ratio of maximum hydraulic supply to demand (i.e., maximumKleaf:gs). This sensitivity ofgs, combined with low minimum epidermal conductance and water storage, could contribute to prolonged leaf survival under drought. The specialized anatomy of subg.Cerastesincludes trichomous stomatal crypts and pronounced hypodermis, and was associated with higher water use efficiency and water storage. Combining our data with comparative literature of other California species, species of subg. Cerastesshow traits associated with greater drought tolerance and reliance on leaf water storage relative to other California species. In addition to drought resistance mechanisms such as mechanical protection and resistance to embolism, drought avoidance mechanisms such as sensitive stomatal closure could contribute importantly to drought tolerance in dry‐climate adapted species.more » « less
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Diversification of plant hydraulic architecture and stomatal function coincides with radical changes in the Earth’s atmosphere over the past 400 my. Due to shared stomatal anatomy with the earliest land plants, bryophyte stomatal behavior may provide insights into the evolution of stomatal function, but significant uncertainty remains due to technical limitations of measuring guard cell turgor pressure in situ. Here, we introduce a method for monitoring cell turgor pressure by nucleating microbubbles within the guard cells of intact plant tissue and then examining microbubble growth and dissolution dynamics. First, we show that maximum microbubble radius decreases with increasing pressure as the pressure of the surrounding fluid constrains its growth according to a modified version of the Epstein–Plesset equation. We then apply this method to monitor turgor pressure in dark- vs. light-acclimated guard cells across bryophyte taxa with stomata, where their role in gas-exchange remains ambiguous, and in vascular plants with well-documented light-dependent turgor modulation. Our findings show no light-activated change in turgor in bryophyte guard cells, with pressures not significantly different than neighboring epidermal cells. In contrast, vascular plants show distinct pressure modulation in response to light that drives reversible changes in stomatal aperture. Complete guard cell turgor loss had no effect on bryophyte stomatal aperture but resulted in partial or complete closure in vascular plants. These results suggest that despite conserved stomatal morphology, the sampled bryophytes lack dynamic control over guard cell turgor that is critical for sustaining photosynthesis and inhibiting desiccation.more » « less
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Abstract 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 (Rleaf) to total soil‐to‐leaf hydraulic resistance (Rtotal), or fRleaf(=Rleaf/Rtotal), is thought to be large, but this has not been tested comprehensively. We compiled a multibiome data set of fRleafusing new and previously published measurements of pressure differences within trees in situ. Across 80 samples, fRleafaveraged 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 fRleaffor 19 tree samples, which was 0.40 (95% CI = 0.29−0.56). The in situ technique produces a more accurate descriptor of fRleafbecause it accounts for dynamic leaf hydraulic conductance. Both approaches demonstrate the outsized role of leaves in controlling tree hydrodynamics. A larger fRleafmay help stems from loss of hydraulic conductance. Thus, the decline in fRleafwith tree height would contribute to greater drought vulnerability in taller trees and potentially to their observed disproportionate drought mortality.more » « less
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null (Ed.)Different microclimates can have significant impact on the physiology of succulents that inhabit arid environments such as the Mojave Desert (California). We investigated variation in leaf physiology, morphology and anatomy of two dominant Mojave Desert monocots, Yucca brevifolia (Joshua tree) and Hesperoyucca whipplei , growing along a soil water availability gradient. Stomatal conductance ( g s ) and leaf thickness were recorded in the field at three different sites (north-western slope, south-eastern slope, and alluvial fan) in March of 2019. We sampled leaves from three individuals per site per species and measured in the lab relative water content at the time of g s measurements, saturated water content, cuticular conductance, leaf morphological traits (leaf area and length, leaf mass per area, % loss of thickness in the field and in dried leaves), and leaf venation. We found species varied in their g s : while Y. brevifolia showed significantly higher g s in the alluvial fan than in the slopes, H. whipplei was highest in the south-eastern slope. The differences in g s did not relate to differences in leaf water content, but rather to variation in number of veins per mm 2 in H. whipplei and leaf width in Y. brevifolia . Our results indicate that H. whipplei displays a higher water conservation strategy than Y. brevifolia . We discuss these differences and trends with water availability in relation to species’ plasticity in morphology and anatomy and the ecological consequences of differences in 3-dimensional venation architecture in these two species.more » « less
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