Tree growth and water-use in hyper-arid Acacia occurs during the hottest and driest season.

Drought-induced tree mortality has been recently increasing and is expected to increase further under warming climate. Conversely, tree species that survive under arid conditions might provide vital information on successful drought resistance strategies. Although Acacia (Vachellia) species dominate many of the globe's deserts, little is known about their growth dynamics and water-use in situ. Stem diameter dynamics, leaf phenology, and sap flow were monitored during 3 consecutive years in five Acacia raddiana trees and five Acacia tortilis trees in the Arid Arava Valley, southern Israel (annual precipitation 20-70 mm, restricted to October-May). We hypothesized that stem growth and other tree activities are synchronized with, and limited to single rainfall or flashflood events. Unexpectedly, cambial growth of both Acacia species was arrested during the wet season, and occurred during most of the dry season, coinciding with maximum daily temperatures as high as 45 °C and vapor pressure deficit of up to 9 kPa. Summer growth was correlated with peak sap flow in June, with almost year-round activity and foliage cover. To the best of our knowledge, these are the harshest drought conditions ever documented permitting cambial growth. These findings point to the possibility that summer cambial growth in Acacia under hyper-arid conditions relies on concurrent leaf gas exchange, which is in turn permitted by access to deep soil water. Soil water can support low-density tree populations despite heat and drought, as long as recharge is kept above a minimum threshold.

Lower leaf gas-exchange and higher photorespiration of treated wastewater irrigated Citrus trees is modulated by soil type and climate.

Water quality, soil and climate can interact to limit photosynthesis and to increase photooxidative damage in sensitive plants. This research compared diffusive and non-diffusive limitations to photosynthesis as well as photorespiration of leaves of grapefruit trees in heavy clay and sandy soils having a previous history of treated wastewater (TWW) irrigation for >10 years, with different water qualities [fresh water (FW) vs TWW and sodium amended treated wastewater (TWW + Na)] in two arid climates (summer vs winter) and in orchard and lysimeter experiments. TWW irrigation increased salts (Na(+) and Cl(-) ), membrane leakage, proline and soluble sugar content, and decreased osmotic potentials in leaves of all experiments. Reduced leaf growth and higher stomatal and non-stomatal (i.e. mesophyll) limitations were found in summer and on clay soil for TWW and TWW + Na treatments in comparison to winter, sandy soil and FW irrigation, respectively. Stomatal closure, lower chlorophyll content and altered Rubisco activity are probable causes of higher limitations. On the other hand, non-photochemical quenching, an alternative energy dissipation pathway, was only influenced by water quality, independent of soil type and season. Furthermore, light and CO2 response curves were investigated for other possible causes of higher non-stomatal limitation. A higher proportion of non-cyclic electrons were directed to the O2 dependent pathway, and a higher proportion of electrons were diverted to photorespiration in summer than in winter. In conclusion, both diffusive and non-diffusive limitations contribute to the lower photosynthetic performance of leaves following TWW irrigation, and the response depends on soil type and environmental factors.

Rationale, methods and baseline demographics of the Bhaktapur Glaucoma Study.

BACKGROUND: To describe the methodology and baseline data of a population-based study designed to determine the prevalence of glaucoma and to study the risk factors for glaucoma development in a Nepali population. DESIGN: Population-based cross-sectional study. PARTICIPANTS: Subjects 40 years and above residing in Bhaktapur District. PATIENTS AND METHODS: Power calculations suggest that a sample size of 4758 is required. Thirty clusters were randomly selected from the 2 municipalities and 16 Village Development Committees of Bhaktapur District in Nepal. A door-to-door census was conducted in the selected clusters to identify citizens 40 years of age and older. Demographic details were collected and a structured interview, regarding awareness for cataract and glaucoma was taken. All individuals fulfilling the eligibility criteria were recruited and referred to the Tilganga Institute of Ophthalmology in Kathmandu for a detailed clinical examination including glaucoma diagnostic procedures. Peripheral blood samples were taken to facilitate future genetic analysis. MAIN OUTCOME MEASURES: Prevalence of glaucoma, risk factors and genetic screening. RESULTS: A total of 4800 people were selected. The mean age of participants was 55.4 ± 12.3 years (range: 40-99) and 51.8% were female. In total, 64.8% of our cohort was aged less than 59 years and 60.5% were illiterate. Among the various ethnic races, 69.7% belonged to the Newar ethnic group. CONCLUSION: This study will determine the prevalence of glaucoma and allow for an increased understanding of the risk factors for glaucoma development in this region.

Intraspecific plasticity in hydraulic and stomatal regulation under drought is linked to aridity at the seed source in a wild pear species.

Adaptations of fruit trees to future climate are a current research priority due to the rapid increase in air temperature and changes in precipitation patterns. This is aimed at securing sustainable food production for our growing populations. Key physiological traits in trees conferring drought tolerance are resistance to embolism and stomatal control over water loss. Recently, we have shown in the field that a native wild pear species performs better under drought than two cultivated pear species. A comparative greenhouse study was conducted to investigate traits associated with drought tolerance in four ecotypes of a wild pear species (Pyrus syriaca Boiss), compared with a wild pear species (Pyrus betulifolia Bunge) commonly used as a pear rootstock. Seed sources were collected from semi-arid, sub-humid and humid sites across northern Israel. Measurements of water relations, leaf physiology, hydraulic conductivity and percent loss of hydraulic conductivity (PLC) were conducted under standard irrigation, drought and recovery from drought. The four P. syriaca ecotypes maintained significantly higher leaf gas exchange values and water-use efficiency and had lower PLC than the rootstock species under prolonged drought as well as during recovery. Across the four ecotypes, stomatal closure occurred at stem water potential (Ψ) around -3.5 MPa; however, Ψ at 50% PLC ranged from -4.1 MPa in the humid ecotype to -5.2 MPa in one of the semi-arid ecotypes, rendering the latter with a higher hydraulic safety margin (the Ψ difference between stomatal closure and 50% PLC). Divergence of the ecotypes in xylem vulnerability to embolism closely matched the mean annual precipitation at their seed sources. Thus, selection of pear ecotypes from populations in semi-arid sites may be better than the currently used plant material for preparing our cultivated species for hotter and drier future climate.

Drought tolerance of wild versus cultivated tree species of almond and plum in the field.

Trees of the genus Prunus produce some of the most widely consumed fruits globally. The combination of climate change-related warming and increased drought stress, scarcity of freshwater resources for irrigation, and increasing demands due to population growth creates a need for increased drought tolerance in these tree species. Recently, we have shown in the field that a native wild pear species performs better under drought than two cultivated pear species. Here, a comparative field study was conducted in Israel to investigate traits associated with drought tolerance in almond (cultivated Prunus dulcis (Mill.) D. A. Webb vs wild Prunus ramonensis Danin) and plum (cultivated Prunus domestica L. vs wild Prunus ursina Kotschy). Measurements of xylem embolism and shoot and root carbon reserves were done along a year, including seasonal drought in the wild and a 35-day drought experiment in the orchards. Synchronous measurements of native xylem embolism and shoot water potential showed that cultivated and wild almond trees lost ~50% of hydraulic conductivity at -2.3 and -3.2 MPa, respectively. Micro-CT images confirmed the higher embolism ratio in cultivated versus wild almond, whereas the two plum species were similar. Dynamics of tissue concentrations of nonstructural carbohydrates were mostly similar across species, with higher levels in cultivated versus wild plum. Our results indicate an advantage for the wild almond over its cultivated relative in terms of xylem resistance to embolism, a major risk factor for trees under drought stress. This result is in line with our previous experiment on pear species. However, the opposite trends observed among the studied plum species mean that these trends cannot be generalized. It is possible that the potential for superior drought tolerance in wild tree species, relative to their cultivated relatives, is limited to wild species from dry and hot habitats.

Reductions in root hydraulic conductivity in response to clay soil and treated waste water are related to PIPs down-regulation in Citrus.

Citrus hydraulic physiology and PIP transcript levels were characterized in heavy (clay) and light (sandy loam) soils with and without treated waste water (TWW) irrigation after a summer irrigation season and at the end of a winter rainy season recovery period. Consistent reductions in clay soils compared to sandy loam were found for fresh water (FW) and TWW irrigation, respectively, in root water uptake, as well as in hydraulic conductivity of whole plant (Ks plant), stem (Ks stem) and root (Ks root). Transcript levels of most PIPs down-regulated following TWW irrigation in both soils, but relative gene expression of three PIPs was significantly higher in summer for sandy soil and FW than for clay soil and TWW; their mRNA levels was significantly correlated to Ks root. A pot experiment, which compared short term influences of saline and TWW found that both treatments, compared to FW, reduced root water uptake and PIPs mRNA levels by 2-fold after 20 days, and the decreases continued with time until the end of the experiment. These latter data indicated that salinity had an important influence. Our results suggest that plant hydraulic adjustment to soil texture and water quality occurs rapidly, i.e. within days, and is modulated by PIPs expression.

Linking the Belowground Microbial Composition, Diversity and Activity to Soilborne Disease Suppression and Growth Promotion of Tomato Amended with Biochar.

Biochar, in addition to sequestering carbon, ameliorating soil, and improving plant performance, can impact foliar and soilborne plant diseases. Nevertheless, the mechanisms associated with suppression of soilborne diseases and improved plant performances are not well understood. This study is designed to establish the relationships between biochar-induced changes in rhizosphere microbial community structure, taxonomic and functional diversity, and activity with soilborne disease suppression and enhanced plant performance in a comprehensive fashion. Biochar suppressed Fusarium crown and root-rot of tomato and simultaneously improved tomato plant growth and physiological parameters. Furthermore, biochar reduced Fusarium root colonization and survival in soil, and increased the culturable counts of several biocontrol and plant growth promoting microorganisms. Illumina sequencing analyses of 16S rRNA gene revealed substantial differences in rhizosphere bacterial taxonomical composition between biochar-amended and non-amended treatments. Moreover, biochar amendment caused a significant increase in microbial taxonomic and functional diversity, microbial activities and an overall shift in carbon-source utilization. High microbial taxonomic and functional diversity and activity in the rhizosphere has been previously associated with suppression of diseases caused by soilborne pathogens and with plant growth promotion, and may collectively explain the significant reduction of disease and improvement in plant performance observed in the presence of biochar.

Impact of treated wastewater on growth, respiration and hydraulic conductivity of citrus root systems in light and heavy soils.

Roots interact with soil properties and irrigation water quality leading to changes in root growth, structure and function. We studied these interactions in an orchard and in lysimeters with clay and sandy loam soils. Minirhizotron imaging and manual sampling showed that root growth was three times lower in the clay relative to sandy loam soil. Treated wastewater (TWW) led to a large reduction in root growth with clay (45-55%) but not with sandy loam soil (<20%). Treated wastewater increased salt uptake, membrane leakage and proline content, and decreased root viability, carbohydrate content and osmotic potentials in the fine roots, especially in clay. These results provide evidence that TWW challenges and damages the root system. The phenology and physiology of root orders were studied in lysimeters. Soil type influenced diameter, specific root area, tissue density and cortex area similarly in all root orders, while TWW influenced these only in clay soil. Respiration rates were similar in both soils, and root hydraulic conductivity was severely reduced in clay soil. Treated wastewater increased respiration rate and reduced hydraulic conductivity of all root orders in clay but only of the lower root orders in sandy loam soil. Loss of hydraulic conductivity increased with root order in clay and clay irrigated with TWW. Respiration and hydraulic properties of all root orders were significantly affected by sodium-amended TWW in sandy loam soil. These changes in root order morphology, anatomy, physiology and hydraulic properties indicate rapid and major modifications of root systems in response to differences in soil type and water quality.

Indirect Evidence for Genetic Differentiation in Vulnerability to Embolism in Pinus halepensis.

Climate change is increasing mean temperatures and in the eastern Mediterranean is expected to decrease annual precipitation. The resulting increase in aridity may be too rapid for adaptation of tree species unless their gene pool already possesses variation in drought resistance. Vulnerability to embolism, estimated by the pressure inducing 50% loss of xylem hydraulic conductivity (P 50), is strongly associated with drought stress resistance in trees. Yet, previous studies on various tree species reported low intraspecific genetic variation for this trait, and therefore limited adaptive capacities to increasing aridity. Here we quantified differences in hydraulic efficiency (xylem hydraulic conductance) and safety (resistance to embolism) in four contrasting provenances of Pinus halepensis (Aleppo pine) in a provenance trial, which is indirect evidence for genetic differences. Results obtained with three techniques (bench dehydration, centrifugation and X-ray micro-CT) evidenced significant differentiation with similar ranking between provenances. Inter-provenance variation in P 50 correlated with pit anatomical properties (torus overlap and pit aperture size). These results suggest that adaptation of P. halepensis to xeric habitats has been accompanied by modifications of bordered pit function driven by variation in pit aperture. This study thus provides evidence that appropriate exploitation of provenance differences will allow continued forestry with P. halepensis in future climates of the Eastern Mediterranean.

Simulating nectarine tree transpiration and dynamic water storage from responses of leaf conductance to light and sap flow to stem water potential and vapor pressure deficit.

For isohydric trees mid-day water uptake is stable and depends on soil water status, reflected in pre-dawn leaf water potential (Ψpd) and mid-day stem water potential (Ψmd), tree hydraulic conductance and a more-or-less constant leaf water potential (Ψl) for much of the day, maintained by the stomata. Stabilization of Ψl can be represented by a linear relationship between canopy resistance (Rc) and vapor pressure deficit (D), and the slope (BD) is proportional to the steady-state water uptake. By analyzing sap flow (SF), meteorological and Ψmd measurements during a series of wetting and drying (D/W) cycles in a nectarine orchard, we found that for the range of Ψmd relevant for irrigated orchards the slope of the relationship of Rc to D, BD is a linear function of Ψmd. Rc was simulated using the above relationships, and its changes in the morning and evening were simulated using a rectangular hyperbolic relationship between leaf conductance and photosynthetic irradiance, fitted to leaf-level measurements. The latter was integrated with one-leaf, two-leaf and integrative radiation models, and the latter gave the best results. Simulated Rc was used in the Penman-Monteith equation to simulate tree transpiration, which was validated by comparing with SF from a separate data set. The model gave accurate estimates of diurnal and daily total tree transpiration for the range of Ψmds used in regular and deficit irrigation. Diurnal changes in tree water content were determined from the difference between simulated transpiration and measured SF. Changes in water content caused a time lag of 90-105 min between transpiration and SF for Ψmd between -0.8 and -1.55 MPa, and water depletion reached 3 l h(-1) before noon. Estimated mean diurnal changes in water content were 5.5 l day(-1) tree(-1) at Ψmd of -0.9 MPa and increased to 12.5 l day(-1) tree(-1) at -1.45 MPa, equivalent to 6.5 and 16.5% of daily tree water use, respectively. Sixteen percent of the dynamic water volume was in the leaves. Inversion of the model shows that Ψmd can be predicted from D and Rc, which may have some importance for irrigation management to maintain target values of Ψmd. That relationship will be explored in future research.

Inactive xylem can explain differences in calibration factors for thermal dissipation probe sap flow measurements.

Thermal dissipation probes (TDPs) were calibrated in three diffuse porous fruit trees and one ornamental species in the field by comparison with heat pulse probes (nectarine and persimmon), in a greenhouse on lysimeters (apple and persimmon) and in the laboratory by pushing water through cut branches (apple, Peltophorum and nectarine). Two operational methods were used: continuous (constant thermal dissipation, CTD) and discontinuous, or transient, heating (transient thermal dissipation, TTD). Correction for the radial distribution of sap flux density was with an analytical function derived from a linear decrease in flux density with depth, as measured with a multi-depth 'Tmax' heat pulse system. When analyzed with previous calibration factors, the measured sap flow was <50% of actual value. The underestimations were consistent, and calibrations for each species in the field, greenhouse and laboratory gave approximately the same factors. Reasonable values of tree water use were obtained with the new calibration factors. Evidence is provided that even though the xylem was diffuse porous, the underestimations were caused by contact of the probes with inactive xylem along their length. The average portion of probe in contact with inactive xylem, measured in stained branches following laboratory calibrations, was 0.2-0.24. Using the measured fractions to correct temperature differentials between heated and unheated probes for CTD and TTD, based on Clearwater et al. (in Potential errors in measurement of nonuniform sap flow using heat dissipation probes. Tree Physiol 1999;19:681-687) almost completely compensated for the underestimations. Calibrations are given for each species both before and after corrections of temperature differentials, along with a multispecies calibration. These results should be an important step in reconciling many reports of different calibration factors for TDP probes.