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Given increasing water deficits across numerous ecosystems world-wide, it is urgent to understand the sequence of failure of leaf function during dehydration. We assessed dehydration-induced losses of rehydration capacity and maximum quantum yield of the photosystem II (F v/F m) in the leaves of 10 diverse angiosperm species, and tested when these occurred relative to turgor loss, declines of stomatal conductance g s, and hydraulic conductance K leaf, including xylem and outside xylem pathways for the same study plants. We resolved the sequences of relative water content and leaf water potential Ψleaf thresholds of functional impairment. On average, losses of leaf rehydration capacity occurred at dehydration beyond 50% declines of g s, K leaf and turgor loss point. Losses of F v/F m occurred after much stronger dehydration and were not recovered with leaf rehydration. Across species, tissue dehydration thresholds were intercorrelated, suggesting trait co-selection. Thresholds for each type of functional decline were much less variable across species in terms of relative water content than Ψleaf. The stomatal and leaf hydraulic systems show early functional declines before cell integrity is lost. Substantial damage to the photochemical apparatus occurs at extreme dehydration, after complete stomatal closure, and seems to be irreversible.

Chaparral is the most abundant vegetation type in California and current climate change models predict more frequent and severe droughts that could impact plant community structure. Understanding the factors related to species-specific drought mortality is essential to predict such changes. We predicted that life history type, hydraulic traits, and plant size would be related to the ability of species to survive drought. We evaluated the impact of these factors in a mature chaparral stand during the drought of 2014, which has been reported as the most severe in California in the last 1,200 years. We measured tissue water potential, native xylem specific conductivity, leaf specific conductivity, percentage loss in conductivity, and chlorophyll fluorescence for 11 species in February 2014, which was exceptionally dry following protracted drought. Mortality among the 11 dominant species ranged from 0 to 93%. Total stand density was reduced 63.4% and relative dominance of species shifted after the drought. Mortality was negatively correlated with water potential, native xylem specific conductivity, and chlorophyll fluorescence, but not with percent loss in hydraulic conductivity and leaf specific conductivity. The model that best explained mortality included species and plant size as main factors and indicated that larger plants had greater survival for 2 of the species. In general, species with greater resistance to water-stress induced cavitation showed greater mortality levels. Despite adult resprouters typically being more vulnerable to cavitation, results suggest that their more extensive root systems enable them to better access soil moisture and avoid harmful levels of dehydration. These results are consistent with the hypothesis that short-term high intensity droughts have the strongest effect on mature plants of shallow-rooted dehydration tolerant species, whereas deep-rooted dehydration avoiding species fare better in the short-term. Severe droughts can drive changes in chaparral structure as a result of the differential mortality among species.

Understanding and predicting plant response to disturbance is of paramount importance in our changing world. Resprouting ability is often considered a simple qualitative trait and used in many ecological studies. Our aim is to show some of the complexities of resprouting while highlighting cautions that need be taken in using resprouting ability to predict vegetation responses across disturbance types and biomes. There are marked differences in resprouting depending on the disturbance type, and fire is often the most severe disturbance because it includes both defoliation and lethal temperatures. In the Mediterranean biome, there are differences in functional strategies to cope with water deficit between resprouters (dehydration avoiders) and nonresprouters (dehydration tolerators); however, there is little research to unambiguously extrapolate these results to other biomes. Furthermore, predictions of vegetation responses to changes in disturbance regimes require consideration not only of resprouting, but also other relevant traits (e.g. seeding, bark thickness) and the different correlations among traits observed in different biomes; models lacking these details would behave poorly at the global scale. Overall, the lessons learned from a given disturbance regime and biome (e.g. crown-fire Mediterranean ecosystems) can guide research in other ecosystems but should not be extrapolated at the global scale.

Desiccation-tolerant (DT) plants can dry past – 100 MPa and subsequently recover function upon rehydration. Vascular DT plants face the unique challenges of desiccating and rehydrating complex tissues without causing structural damage. However, these dynamics have not been studied in intact DT plants. We used high resolution micro-computed tomography (microCT), light microscopy, and fluorescence microscopy to characterize the dynamics of tissue desiccation and rehydration in petioles (stipes) of intact DT ferns. During desiccation, xylem conduits in stipes embolized before cellular dehydration of living tissues within the vascular cylinder. During resurrection, the chlorenchyma and phloem within the stipe vascular cylinder rehydrated before xylem refilling. We identified unique stipe traits that may facilitate desiccation and resurrection of the vascular system, including xylem conduits containing pectin (which may confer flexibility and wettability); chloroplasts within the vascular cylinder; and an endodermal layer impregnated with hydrophobic substances that impede apoplastic leakage while facilitating the upward flow of water within the vascular cylinder. Resurrection ferns are a novel system for studying extreme dehydration recovery and embolism repair in the petioles of intact plants. The unique anatomical traits identified here may contribute to the spatial and temporal dynamics of water movement observed during desiccation and resurrection.

Possible mechanical and hydraulic costs to increased cavitation resistance were examined among six co-occurring species of chaparral shrubs in southern California. We measured cavitation resistance (xylem pressure at 50% loss of hydraulic conductivity), seasonal low pressure potential (P[subscript min]), xylem conductive efficiency (specific conductivity), mechanical strength of stems (modulus of elasticity and modulus of rupture), and xylem density. At the cellular level, we measured vessel and fiber wall thickness and lumen diameter, transverse fiber wall and total lumen area, and estimated vessel implosion resistance using (t/b)[subscript h]², where t is the thickness of adjoining vessel walls and b is the vessel lumen diameter. Increased cavitation resistance was correlated with increased mechanical strength (r² = 0.74 and 0.76 for modulus of elasticity and modulus of rupture, respectively), xylem density (r² = 0.88), and P[subscript min] (r² = 0.96). In contrast, cavitation resistance and P[subscript min] were not correlated with decreased specific conductivity, suggesting no tradeoff between these traits. At the cellular level, increased cavitation resistance was correlated with increased (t/b)[subscript h]² (r² = 0.95), increased transverse fiber wall area (r² = 0.89), and decreased fiber lumen area (r² = 0.76). To our knowledge, the correlation between cavitation resistance and fiber wall area has not been shown previously and suggests a mechanical role for fibers in cavitation resistance. Fiber efficacy in prevention of vessel implosion, defined as inward bending or collapse of vessels, is discussed.

Wood density ($D_{\\text{t}}$), an excellent predictor of mechanical properties, is typically viewed in relation to support against gravity, wind, snow, and other environmental forces. In contrast, we show the surprising extent to which variation in$D_{\\text{t}}$and wood structure is linked to support against implosion by negative pressure in the xylem pipeline. The more drought-tolerant the plant, the more negative the xylem pressure can become without cavitation, and the greater the internal load on the xylem conduit walls. Accordingly,$D_{\\text{t}}$was correlated with cavitation resistance. This trend was consistent with the maintenance of a safety factor from implosion by negative pressure: conduit wall span (b) and thickness (t) scaled so that$(t/b)^{2}$was proportional to cavitation resistance as required to avoid wall collapse. Unexpectedly, trends in$D_{\\text{t}}$may be as much or more related to support of the xylem pipeline as to support of the plant.

The centrifuge method for measuring the resistance of xylem to cavitation by water stress was modified to also account for any additional cavitation that might occur from a freeze-thaw cycle. A strong correlation was found between cavitation by freezing and mean conduit diameter. On the one extreme, a tracheid-bearing conifer and diffuse-porous angiosperms with small-diameter vessels (mean diameter <30 micrometers) showed no freezing-induced cavitation under modest water stress (xylem pressure = -0.5 MPa), whereas species with larger diameter vessels (mean >40 micrometers) were nearly completely cavitated under the same conditions. Species with intermediate mean diameters (30-40 micrometers) showed partial cavitation by freezing. These results are consistent with a critical diameter of 44 micrometers at or above which cavitation would occur by a freeze-thaw cycle at -0.5 MPa. As expected, vulnerability to cavitation by freezing was correlated with the hydraulic conductivity per stem transverse area. The results confirm and extend previous reports that small-diameter conduits are relatively resistant to cavitation by freezing. It appears that the centrifuge method, modified to include freeze-thaw cycles, may be useful in separating the interactive effects of xylem pressure and freezing on cavitation.

1. Shrubs in fire prone chaparral communities have evolved different life history types in response to fire. A key to understanding the evolution of life history type differences is to understand how physiological traits are linked to differences in life history type. Vascular adaptations are important for delivering an efficient and stable water supply to evergreen chaparral shrub leaves. This study tested for a link between vascular physiology and life history type in chaparral shrubs. 2. Chaparral shrub species along the south-western coast of North America survive wildfire by three different life histories. Non-sprouters are killed by fire and re-establish exclusively through germination of fire-stimulated seeds, facultative sprouters re-establish by a combination of vegetative sprouting and fire-stimulated seeds, and obligate sprouters re-establish exclusively by vegetative sprouting because their seeds do not survive fire. Non-sprouters and facultative sprouters establish seedlings in the open canopy post fire environment, whereas obligate sprouters establish seedlings in the shady understory of the mature chaparral canopy. 3. Seedlings of nine species (Rhamnaceae) representing three each of the different life history types were grown in deep containers in a common garden under treatments of sun and shade. Hydraulic conductance was measured using a high-pressure flow meter for all organs, and a vacuum technique was used to measure conductance of fine and woody roots. We predicted that non-sprouters would exhibit greater hydraulic efficiency than the sprouting species, and that facultative sprouters would be more efficient than the shade tolerant obligate sprouters. 4. Non-sprouters had the greatest hydraulic conductance per unit leaf and sapwood area at the whole seedling level, whereas facultative and obligate sprouters were not different. Comparing hydraulic conductance across major organs (from fine roots to leaves) showed that the hydraulic system was well coordinated. At the whole seedling level, the root system was more of a bottleneck than the shoot system. This pattern was consistent with high resistance extraxylary pathways in roots and differences in root architecture. 5. The greater hydraulic efficiency of the non-sprouter life history type is attributed to its post-fire pioneering habit and may partially explain the relatively high speciation in the non-sprouters. Lower hydraulic efficiency is associated with a sprouting life history and greater shade tolerance. The seedling root systems represent a hydraulic bottleneck that may place roots under especially intense selection.

Trees grow tall where resources are abundant, stresses are minor, and competition for light places a premium on height growth 1 , 2 . The height to which trees can grow and the biophysical determinants of maximum height are poorly understood. Some models predict heights of up to 120 m in the absence of mechanical damage 3 , 4 , but there are historical accounts of taller trees 5 . Current hypotheses of height limitation focus on increasing water transport constraints in taller trees and the resulting reductions in leaf photosynthesis 6 . We studied redwoods ( Sequoia sempervirens ), including the tallest known tree on Earth (112.7 m), in wet temperate forests of northern California. Our regression analyses of height gradients in leaf functional characteristics estimate a maximum tree height of 122–130 m barring mechanical damage, similar to the tallest recorded trees of the past. As trees grow taller, increasing leaf water stress due to gravity and path length resistance may ultimately limit leaf expansion and photosynthesis for further height growth, even with ample soil moisture.

Progressive diebacks of outer canopy branchlets of Ceanothus crassifolius were repeatedly observed after rainless periods up to 9 mo in duration in the Santa Monica Mountains of southern California. Mean xylem pressures of branchlets near the end of drought were as low as -11.2 MPa (N = 22) with a mean of about 60 dead branchlets per shrub. Inoculation (N = 15) with three species of fungi previously isolated from the same population of C. crassifolius did not promote dieback, suggesting that the observed decline was not fungal induced, as had been proposed. Further, at least 50% of healthy-appearing twigs, without symptoms of dieback, contained isolatible endophytic fungi. We used a centrifugal force method to determine the range of xylem pressure causing cavitation (vulnerability curves) for branchlets (N = 12) and roots (N = 16). We combined vulnerability curves with soil texture data (N = 6) into a water transport model that estimated the critical values ($P_{Lcrit}$) of leaf xylem pressure associated with the loss of water from soil to foliage. Maximum$P_{Lcrit}$was between -10 and -11 MPa and within the range of minimum measured xylem pressures of branchlets during drought and dieback. Branchlet dieback correlated with seasonal declines in xylem pressure in concert with declining safety margins from hydraulic failure. Symptoms of dieback were duplicated in the field by partially severing stem xylem that normally supplied branchlets with water. Taken together, these results indicate that loss of hydraulic conductance to foliage was the probable cause of the observed dieback in C. crassifolius. Partial dieback of peripheral branchlets, and its attendant reduction in evaporative surface area, may be a last-resort mechanism for whole-plant water conservation and drought survival in this species.