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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 (Fv /Fm ) in the leaves of 10 diverse angiosperm species, and tested when these occurred relative to turgor loss, declines of stomatal conductance gs , and hydraulic conductance Kleaf , 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 gs , Kleaf and turgor loss point. Losses of Fv /Fm 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.

The persistence of college students in STEM majors after their first-year of college is approximately 50%, with underrepresented populations displaying even higher rates of departure. For many undergraduates, their first-year in college is defined by large class sizes, poor access to research faculty, and minimal standing in communities of scholars. Pepperdine University and Whittier College, funded by a National Science Foundation award to Improve Undergraduate Stem Education (NSF IUSE), partnered in the development of first-year classes specifically geared to improve student persistence in STEM and academic success. This Students as Scholars Program (SAS) engaged first-year undergraduates in scholarly efforts during their first semester in college with a careful approach to original research design and mentoring by both faculty and upperclassmen experienced in research. Courses began by introducing hypothesis formulation and experimental design partnered with the scientific focus of each course (ecological, biochemical, microbiological). Students split into research teams, explored the primary literature, designed research projects, and executed experiments over a 6–7 week period, collecting, analyzing, and interpreting data. Microbiology-specific projects included partnerships with local park managers to assess water quality and microbial coliform contamination at specified locations in a coastal watershed. In addition, students explored the impact of soil salinity on microbial community structure. Analysis of these samples included next-generation sequencing and microbiome compositional analysis via collaboration with students from an upper division microbiology course. This cross-course collaboration facilitated additional student mentoring opportunities between upperclassmen and first-year students. This approach provided first-year students an introduction to the analysis of complex data sets using bioinformatics and statistically reliable gas-exchange replicates. Assessment of the impact of this program revealed students to view the research as challenging, but confidence building as they take their first steps as biology majors. In addition, the direct mentorship of first-year students by upperclassmen and faculty was viewed positively by students. Ongoing assessments have revealed SAS participants to display a 15% increased persistence rate in STEM fields when compared to non-SAS biology majors.

The purpose of this report is to provide detailed guidance on the dosimetry of the INTRABEAM® (Carl Zeiss Medical AG, Jena, Germany) electronic brachytherapy (eBT) system as it stands at the present time. This report has been developed by the members of American Association of Physicists in Medicine (AAPM) Task Group 292 and endorsed by the AAPM. Members of AAPM Task Group 292 on Electronic-Brachytherapy Dosimetry have reviewed pertinent publications and user manuals regarding the INTRABEAM system dosimetry and manufacturer-supplied dose calculation protocols. Formal written correspondence with Zeiss has also provided further clarification. Dose-rate calculations for the INTRABEAM system are highly dependent on choice of dosimetry protocol. Even with careful protocol selection, large uncertainties remain due to the incomplete characterization of the ionization chambers used for verification with respect to their energy dependence as well as manufacturing variations. There are two distinct sets of dose-rate data provided by Zeiss for the INTRABEAM system. One dataset (Calibration V4.0) is representative of the physical dose surrounding the source and the other dataset (TARGIT) has been adjusted to be consistent with a clinical trial named TARGIT (TARGeted Intraoperative RadioTherapy). The adjusted TARGIT doses are quite dissimilar to the physical doses, with differences ranging from 14% to 30% at the surface of a spherical applicator, depending on its diameter, and up to a factor of two at closer distances with the smaller needle applicators. In addition, ion chamber selection and associated manufacturing tolerances contribute to significant additional uncertainties. With these substantial differences in dose rates and their associated uncertainties, it is important for users to be aware of how each value is calculated and whether it is appropriate to be used for the intended treatment. If users intend to deliver doses that are the same as they were in 1998 at the onset of the TARGIT trial, then the TARGIT dose-rate tables should be used. The Calibration V4.0 dose rates may be more appropriate to use for applications other than TARGIT trial treatments, since they more closely represent the physical doses being delivered. Users should also be aware of the substantial uncertainties associated with the provided dose rates, which are due to beam hardening, chamber geometry, and selection of the point-of-measurement for a given ionization chamber. This report serves to describe the details and implications of the manufacturer-recommended dosimetry formalism for users of the INTRABEAM system.

Positive root pressure was critical for whole-plant recovery in two resurrection ferns, but foliar water uptake expedited leaf recovery., Desiccation-tolerant (DT) organisms can lose nearly all their water without dying. Desiccation tolerance allows organisms to survive in a nearly completely dehydrated, dormant state. At the cellular level, sugars and proteins stabilize cellular components and protect them from oxidative damage. However, there are few studies of the dynamics and drivers of whole-plant recovery in vascular DT plants. In vascular DT plants, whole-plant desiccation recovery (resurrection) depends not only on cellular rehydration, but also on the recovery of organs with unequal access to water. In this study, in situ natural and artificial irrigation experiments revealed the dynamics of desiccation recovery in two DT fern species. Organ-specific irrigation experiments revealed that the entire plant resurrected when water was supplied to roots, but leaf hydration alone (foliar water uptake) was insufficient to rehydrate the stele and roots. In both species, pressure applied to petioles of excised desiccated fronds resurrected distal leaf tissue, while capillarity alone was insufficient to resurrect distal pinnules. Upon rehydration, sucrose levels in the rhizome and stele dropped dramatically as starch levels rose, consistent with the role of accumulated sucrose as a desiccation protectant. These findings provide insight into traits that facilitate desiccation recovery in dryland ferns associated with chaparral vegetation of southern California.

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.

Resistance of stem xylem to water stress-induced cavitation and embolism among chaparral shrub species in California has been extensively studied, providing the opportunity to examine broad patterns in cavitation resistance. We used previously published as well as unpublished vulnerability to cavitation curve data from 16 chaparral shrub species of southern California to examine the variability of cavitation resistance across sites, regions, and seasons. Additionally, these data provided a unique opportunity to address a recent methodological debate within the field of plant hydraulics. We found that different methods, specifically a centrifuge method and a dehydration method, produced similar results (P = 0.184). Vulnerability to cavitation varied seasonally, with species exhibiting greater susceptibility to water-stress induced cavitation during the wet season (P = 0.003). Cavitation resistance did not differ among sites that were less than 10 km apart even though these sites differed in th...

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.

PREMISE OF THE STUDY California experienced severe drought between 2012 and 2016. During this period, we compared seasonal changes in tissue-water relations among eight fern species in the Santa Monica Mountains of southern California to elucidate differential mechanisms of drought survival and physiological performance during extreme water deficits. METHODS We monitored seasonal changes in water potential (Ψmd) and dark-adapted chlorophyll fluorescence (Fv/Fm), assessed tissue-water relations including osmotic potential at saturation and the turgor loss point (Ψπ, sat and Ψπ, tlp), and measured, for two evergreen species, xylem-specific and leaf-specific hydraulic conductivity (Ks and Kl) and vulnerability of stem xylem to water stress-induced embolism (water potential at 50% loss hydraulic conductivity, Ψ50). KEY RESULTS Species grew in either riparian or chaparral understory. The five chaparral species had a wider range of seasonal water potentials, root depths, and frond phenological traits, including one evergreen, two summer-deciduous, and two desiccation-tolerant (resurrection) species. Evergreen species were especially diverse, with an evergreen riparian species maintaining seasonal water potentials above -1.3 MPa, while an evergreen chaparral species had seasonal water potentials below -8 MPa. In those two species the Ψ50 values were -2.5 MPa and -4.3 MPa, respectively. CONCLUSIONS Observed differences in physiological performance among eight fern species reflected niche partitioning in water utilization and habitat preference associated with distinct phenological traits. We predict differential survival among fern species as future drought events in California intensify, with desiccation-tolerant resurrection ferns being the most resistant.

Browsing by exotic mule deer on Santa Catalina Island (SCI) off the coast of southern California may diminish the post-fire resilience of native shrublands. To assess this, deer exclosures were established following a wildfire to monitor post-fire recovery of three dominant, native shrub species (Heteromeles arbutifolia, Rhus integrifolia, and Rhamnus pirifolia). Post-fire resprout growth, mortality, and tissue water status as well as pre- and post-fire shrub density and cover were measured inside and outside of deer exclosures. We found that deer browsing significantly limited post-fire resprout growth and led to increased mortality of resprouting H. arbutifolia shrubs (88 % mortality outside compared to 11 % inside exclosures). Post-fire resprouts maintained favorable water status during the study despite drought conditions, indicating that water stress was not a proximate cause of resprout mortality. Deer browsing resulted in a >93 % reduction in canopy coverage of dominant shrub species. The dramatic reduction of native shrubs at this site may create opportunities for displacement by exotic species, resulting in eventual vegetation-type conversion. The observed link between intense browsing and post-fire shrub mortality provides much needed information concerning the environmental impact of exotic deer on SCI and illustrates the interaction between exotic herbivores and fire on an island system.

Summary 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 nonsprouters. 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.

Plant adaptations to the environment are limited, and therefore plants in similar environments may display similar functional and physiological traits, a pattern termed functional convergence. Evidence was examined for functional convergence among 28 evergreen woody shrubs from three plant communities of the semi-arid winter rainfall region of southern California. Both leaf and water relations traits were examined, including seasonal stomatal conductance (gs), specific leaf area (SLA), leaf specific conductivity (Kl), seasonal water potential (Psi w), stem cavitation resistance (Psi 50), and xylem density. Species display community-specific suites of xylem and leaf traits consistent with different patterns of water use among communities, with coastal sage scrub species utilizing shallow pulses of water, Mojave Desert scrub species relying on deeper water reserves, and chaparral species utilizing both shallow and deep moisture reserves. Communities displayed similar degrees of water stress, with a community-level minimum Psi w (Psi wmin) of c. -4.6 Mpa, similar to other arid communities. Pooled across sites, there was a strong correlation between Psi wmin and xylem density, suggesting that these traits are broadly related and predictive of one another. This comparative community physiology approach may be useful in testing hypotheses of functional convergence across structurally similar semi-arid communities.

Water plays a central role in plant biology and the efficiency of water transport throughout the plant affects both photosynthetic rate and growth, an influence that scales up deterministically to the productivity of terrestrial ecosystems. Moreover, hydraulic traits mediate the ways in which plants interact with their abiotic and biotic environment. At landscape to global scale, plant hydraulic traits are important in describing the function of ecological communities and ecosystems. Plant hydraulics is increasingly recognized as a central hub within a network by which plant biology is connected to palaeobiology, agronomy, climatology, forestry, community and ecosystem ecology and earth-system science. Such grand challenges as anticipating and mitigating the impacts of climate change, and improving the security and sustainability of our food supply rely on our fundamental knowledge of how water behaves in the cells, tissues, organs, bodies and diverse communities of plants. A workshop, 'Emerging Frontiers in Plant Hydraulics' supported by the National Science Foundation, was held in Washington DC, 2015 to promote open discussion of new ideas, controversies regarding measurements and analyses, and especially, the potential for expansion of up-scaled and down-scaled inter-disciplinary research, and the strengthening of connections between plant hydraulic research, allied fields and global modelling efforts.

Resistance to xylem cavitation depends on the size of xylem pit membrane pores and the strength of vessels to resist collapse or, in the case of freezing-induced cavitation, conduit diameter. Altering these traits may impact plant biomechanics or water transport efficiency. The evergreen sclerophyllous shrub species, collectively referred to as chaparral, which dominate much of the mediterranean-type climate region of southern California, have been shown to display high cavitation resistance (pressure potential at 50% loss of hydraulic conductivity; P50). We examined xylem functional and structural traits associated with more negative P50 in stems of 26 chaparral species. We correlated raw-trait values, without phylogenetic consideration, to examine current relationships between P50 and these xylem traits. Additionally, correlations were examined using phylogenetic independent contrasts (PICs) to determine whether evolutionary changes in these xylem traits correlate with changes in P50. Co-occurring chaparral species widely differ in their P50 (� 0.9 to � 11.0 MPa). Species experiencing the most negative seasonal pressure potential (Pmin) had the highest resistance to xylem cavitation (lowest P50). Decreased P50 was associated with increased xylem density, stem mechanical strength (modulus of rupture), and transverse fiber wall area when both raw values and PICs were analyzed. These results support a functional and evolutionary relationship among these xylem traits and cavitation resistance. Chaparral species that do not sprout following fire but instead recruit post-fire from seed had the greatest resistance to cavitation, presumably because they rely on post-fire survival of seedlings during the summer dry period to persist in the landscape. Raw values of hydraulic vessel diameter (dh), maximum vessel length, and xylem-specific hydraulic conductivity (Ks) were correlated to P50; however, dh, maximum vessel length, and Ks were not correlated to P50 when analyzed using PICs, suggesting that these traits have not undergone correlated evolutionary change. We found no difference in xylem traits between species occurring at freezing vs. nonfreezing sites, although freezing has been shown to affect the survival and distributions of some chaparral species. Stem mechanical strength, fiber properties, and post-fire regeneration type appear to be key factors in the evolution of cavitation resistance among chaparral shrubs.

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., This work was performed under the framework of the TREVOL projects (CGL2012-39938-C02-01 to J.G.P.) from the Spanish Government. A.L.J., R.B.P., A.V. and S.P. were supported by the following grants: IOS-1252232 (NSF), IOS-0845125 (NSF), CGL-2011-30531-CO2-02 (SURVIVE Project, Spain), ID-1120458 (Fondo Nacional de Desarrollo Científico y Tecnológico, FONDECYT, Chile), respectively. Fundación Centro de Estudios Ambientales del Mediterráneo (CEAM) is supported by Generalitat Valenciana.

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.

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 mm) showed no freezing-induced cavitation under modest water stress (xylem pressure 52 0.5 MPa), whereas species with larger diameter vessels (mean .40 mm) were nearly completely cavitated under the same conditions. Species with intermediate mean diameters (30‐40 mm) showed partial cavitation by freezing. These results are consistent with a critical diameter of 44 mm at or above which cavitation would occur by a freeze‐thaw cycle at 20.5 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.

Possible tradeoffs between efficiency of water transport and mechanical strength were examined in stems of two congeneric pairs of co-occurring chaparral shrubs. First, since previously published results indicated that Adenostoma sparsifolium (Rosaceae) had greater specific conductivity (k s or hydraulic conductivity per xylem transverse area) than A. fasciculatum, it was hypothesized that A. sparsifolium would have greater vessel lumen area per square millimeter of xylem area, and less mechanical strength, than A. fasciculatum. Secondly, since Ceanothus megacarpus (Rhamnaceae) is a non-sprouter (unable to sprout from the root crown following fire or other major disturbance) whereas C. spinosus is a sprouter and thus able to form new stems following disturbance, it was hypothesized that C. megacarpus would have greater mechanical strength, but lower k s, than C. spinosus. Both hypotheses were supported. Based upon computer-aided image analyses, A. sparsifolum had significantly higher mean and maximum vessel diameters (16.4, 40.5 vs. 14.6, 35.7 μm), a 34% greater percent vessel lumen area, and a two-fold greater measured and theoretical k s than A. fasciculatum. This corresponded to 14% lower stem density (wet weight/volume) and less mechanical strength, with a 37% lower modulus of elasticity (MOE) and a 30% lower modulus of rupture (MOR) than A. fasciculatum. Similarly, C.␣spinosus had a significantly higher maximum vessel diameter (52.7 vs. 41.8 μm) and a 92% higher theoretical k s (and 43% higher measured k s) than C. megacarpus. This corresponded to a 9% lower stem density and 20% lower MOR than for C. megacarpus. Thus, at least within these two congeneric pairs of chaparral shrubs growing together in the same habitat, there may be tradeoffs between mechanical strength and conductive efficiency of the stem xylem which correspond to differences in transport physiology and life history traits of sprouter versus non-sprouter species.

Leaf death due to freezing was examined for four, co-occurring species of chaparral shrubs from the Santa Monica Mountains of southern California, Rhus laurina ( 5 Malosma laurina), R. ovata, Ceanothus megacarpus , and C. spinosus. Measurements were made on seedlings vs. adults for all species, and for Rhus spp. in winter vs. summer, and at a warm vs. a cold site. We used four methods to determine the temperature for 50% change in activity or cell death (LT50) of leaves: (1) electrical conductivity (electrolyte leakage into a bathing solution), (2) photosynthetic fluorescent capacity (Fv/Fm), (3) percentage of palisade mesophyll cells stained by fluorescein diacetate vital stain, and (4) visual score of leaf color (Munsell color chart). In all four species seedlings were found to be more sensitive to freezing temperatures than were adults by 1 8‐ 38C. For adults the LT50 ranged from 258C for Rhus laurina in the summer to 2168C for Rhus ovata in the winter. The LT50 of R. ovata located at a colder inland site was 4 8 C lower than R. ovata at the warmer coastal site just 4 km apart, suggesting ecotypic differences between R. ovata at the two sites. Both R. laurina and R. ovata underwent significant winter hardening. At the cold site, R. ovata acclimated by 68C on average, while R. laurina acclimated by only 38C. These results were consistent with species distributions and with field observations of differential shoot dieback between these two congeneric species after a natural freeze‐thaw event in the Santa Monica Mountains.

Water transport from the roots to leaves in chaparral shrubs of California occurs through xylem vessels and tra-cheids. The formation of gas bubbles in xylem can block water transport (gas embolism), leading to shoot dieback. Two environmental factors that cause gas embolism formation in xylem conduits are drought and freezing air temperatures. We compared the differential vulnerabili-ties of Rhus laurina and Ceanothus megacarpus, co-domi-nant shrub species in the coastal regions of the Santa Monica Mountains of southern California, to both water stress-induced and freezing-induced embolism of their xylem. Rhus laurina has relatively large xylem vessel diameters, a deep root system, and a large basal burl from which it vigorously resprouts after wildfire or freezing injury. In contrast, Ceanothus megacarpus has small-diameter vessels, a shallow root system, no basal burl and is a non-sprouter after shoot removal by wildfire. We found that R. laurina became 50% embolized at a water stress of -3 MPa and 100% embolized by a freeze-thaw cycle at all hydration levels. In contrast, C. megacarpus became 50% embolized at a water stress of -9 MPa and 100% embolized by freeze-thaw events only at water potentials lower than -3 MPa. Reducing thaw rates from 0.8 °C min -1 to 0.08 °C min -1 (the normal thaw rate measured in situ) had no effect on embolism formation in R. laurina but significantly reduced embolism occurrence in well-hydrated C. megacarpus (embolism reduced from 74 to 35%). These results were consistent with the theory of gas bubble formation and dissolution in xylem sap. They also agree with field observations of differential shoot dieback in these two species after a natural freeze-thaw event in the Santa Monica Mountains.

 Premise of the Study: California chaparral shrub species have different life history types: Nonsprouters (NS) are killed by fi and persist through a fi re-stimulated seed bank; facultative sprouters (FS) reestablish by a combination of vegetative sprouting and seeding; and obligate sprouters (OS) reestablish exclusively by sprouting. Nonsprouters and FS establish seedlings in opencanopy postfi re environments, whereas OS establish seedlings between fi res in the shady understory. We hypothesized that allocation differences among seedlings of postfi re sprouters and nonsprouters and regeneration niche differences would lead to contrasting patterns in biomass accumulation (NS > FS > OS, in sun; OS > FS > NS, in shade).  Methods: Seedlings of three species from each life history type were grown in sun and 75% shade. We measured net carbon assimilation and biomass accumulation after one year.  Key Results: Biomass accumulation was similar in the sun except FS>OS. In the shade, NS had lower biomass than FS and OS. Assimilation rates, nitrogen relations, and allocation differences could not fully explain biomass accumulation differences. Instead, biomass accumulation was inversely related to water-stress tolerance and shade tolerance. Additionally, OS and FS differed in root/shoot allocation even though both are sprouters.  Conclusions: Seedling growth and carbon assimilation rates were divergent among three life history types and were consistent with differences in tolerance to water stress and shade or sun regeneration niches, but not tradeoffs in sprouting-related allocation differences per se.

Leaf tensile properties were compared between the mesic deciduous tree Prunus serrulata (var. "Kwanzan") and the xeric and sclerophyllous chaparral evergreen shrub Heteromeles arbutifolia (M. Roem). All values for biomechanical parameters for H. arbutifolia were significantly greater than those of P. serrulata. The fracture planes also differed between the two species with P. serrulata fracturing along the secondary veins, while H. arbutifolia most often fractured across the leaf irrespective of the vein or mesophyll position, thus yielding qualitative differences in the stress-strain curves of the two species. Anatomically, P. serrulata exhibits features typical for a deciduous mesophytic leaf such as a thin cuticle, a single layer of palisade mesophyll, isodiametric spongy mesophyll, and extensive reticulation of the laminar veins. Heteromeles arbutifolia leaves, however, are typically two- to three-fold thicker with a 35% higher dry mass/fresh mass ratio. The vascular tissue is restricted to the interface of the palisade and spongy mesophyll near the center of the leaf. Both epidermal layers have a thick cuticle. The palisade mesophyll is tightly packed and two to three layers thick. The spongy mesophyll cells are ameboid in shape and tightly interlinked both to other spongy cells as well as to the overlying palisade layer. We conclude that the qualitative and quantitative biomechanical differences between the leaves of these two species are likely due to a complex interaction of internal architectural arrangement and the physical/chemical differences in the properties of their respective cell walls. These studies illustrate the importance that morphological and anatomical correlates play with mechanical behavior in plant material and ultimately reflect adaptations present in the leaves of chaparral shrubs that are conducive to surviving in arid environments.

Intracavitary accelerated partial breast irradiation (APBI) has become a popular treatment for early stage breast cancer in recent years due to its shortened course of treatment and simplified treatment planning compared to traditional external beam breast conservation therapy. However, the exit dose to the skin is a major concern and can be a limiting factor for these treatments. Most treatment planning systems (TPSs) currently used for high dose-rate (HDR) 192Ir brachytherapy overestimate the exit skin dose because they assume a homogeneous water medium and do not account for finite patient dimensions. The purpose of this work was to quantify the TPS overestimation of the exit skin dose for a group of patients and several phantom configurations.The TPS calculated skin dose for 59 HDR 192Ir APBI patients was compared to the skin dose measured with LiF:Mg,Ti thermoluminescent dosimeters (TLDs). Additionally, the TPS calculated dose was compared to the TLD measured dose and the Monte Carlo (MC) calculated dose for eight phantom configurations. Four of the phantom configurations simulated treatment conditions with no scattering material beyond the point of measurement and the other four configurations simulated the homogeneous scattering conditions assumed by the TPS. Since the calibration TLDs for this work were irradiated with 137Cs and the experimental irradiations were performed with 192Ir, experiments were performed to determine the intrinsic energy dependence of the TLDs. Correction factors that relate the dose at the point of measurement (center of TLD) to the dose at the point of interest (basal skin layer) were also determined and applied for each irradiation geometry.The TLD intrinsic energy dependence for 192Ir relative to 137Cs was 1.041 +/- 1.78%. The TPS overestimated the exit skin dose by an average of 16% for the group of 59 patients studied, and by 9%-15% for the four phantom setups simulating treatment conditions. For the four phantom setups simulating the conditions assumed by the TPS, the TPS calculated dose agreed well with the TLD and MC results (within 3% and 1%, respectively). The inverse square geometry correction factor ranged from 1.023 to 1.042, and an additional correction factor of 0.978 was applied to account for the lack of charged particle equilibrium in the TLD and basal skin layer.TPS calculations that assume a homogeneous water medium overestimate the exit skin dose for intracavitary APBI treatments. It is important to determine the actual skin dose received during intracavitary APBI to determine the skin dose-response relationship and establish dose limits for optimal skin sparing. This study has demonstrated that TLDs can measure the skin dose with an expanded uncertainty (k = 2) of 5.6% when the proper corrections are applied.

Trees grow tall where resources are abundant, stresses are minor, and competition for light places a premium on height growth. 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, but there are historical accounts of taller trees. Current hypotheses of height limitation focus on increasing water transport constraints in taller trees and the resulting reductions in leaf photosynthesis. 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.

Freeze/thaw stress was examined in chaparral shrubs of the genus Ceanothus to determine the interactive effects of freezing and drought and to consider which is the more vulnerable component, the living leaves (symplast) or the non-living water transport system (apoplast). We hypothesized that where Ceanothus species co-occurred, the more inland species C. crassifolius would be more tolerant of low temperatures than the coastal species C. spinosus, both in terms of leaf survival (LT(50), or the temperature at which there is 50% loss of function or viability) and in terms of resistance to freezing-induced embolism (measurements of percent loss hydraulic conductivity due to embolism following freeze/thaw). Cooling experiments on 2 m long winter-acclimated shoots resulted in LT(50) values of about -10 degrees C for C. spinosus versus -18 degrees C for C. crassifolius. Freeze-thaw cycles resulted in no change in embolism when the plants were well hydrated (-0.7 to -2.0 MPa). However, when plants were dehydrated to -5.0 MPa, C. spinosus became 96% embolized with freeze/thaw, versus only 61% embolism for C. crassifolius. Stems of C. crassifolius became 90% and 97% embolized at -6.6 and -8.0 MPa, respectively, meaning that even in this species, stems could be more vulnerable than leaves under conditions of extreme water stress combined with freeze/thaw events. The dominance of C. crassifolius at colder sites and the restriction of C. spinosus to warmer sites are consistent with both the relative tolerance of their symplasts to low temperatures and the relative tolerance of their apoplasts to freeze events in combination with drought stress.