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.

Orchard recycling improves climate change adaptation and mitigation potential of almond production systems.

There is an urgent need to develop climate smart agroecosystems capable of mitigating climate change and adapting to its effects. In California, high commodity prices and increased frequency of drought have encouraged orchard turnover, providing an opportunity to recycle tree biomass in situ prior to replanting an orchard. Whole orchard recycling (WOR) has potential as a carbon (C) negative cultural practice to build soil C storage, soil health, and orchard productivity. We tested the potential of this practice for long term C sequestration and hypothesized that associated co-benefits to soil health will enhance sustainability and resiliency of almond orchards to water-deficit conditions. We measured soil health metrics and productivity of an almond orchard following grinding and incorporation of woody biomass vs. burning of old orchard biomass 9 years after implementation. We also conducted a deficit irrigation trial with control and deficit irrigation (-20%) treatments to quantify shifts in tree water status and resilience. Biomass recycling led to higher yields and substantial improvement in soil functioning, including nutrient content, aggregation, porosity, and water retention. This practice also sequestered significantly higher levels of C in the topsoil (+5 t ha-1) compared to burning. We measured a 20% increase in irrigation water use efficiency and improved soil and tree water status under stress, suggesting that in situ biomass recycling can be considered as a climate smart practice in California irrigated almond systems.

Bees increase crop yield in an alleged pollinator-independent almond variety.

Wild pollinators are declining and the number of managed honey bee colonies is growing slower than agricultural demands for pollination. Because of these contrasting trends in pollinator demand and availability, breeding programs for many pollinator-dependent crops have focused on reducing the need for pollinators. Although numerous crop varieties are now available in the market with the label of pollinator-independent, the real dependence of these varieties on pollinators is mostly unknown. We evaluated the hypothesis of pollinator independence in the Independence almond variety, the fastest growing variety in California that is the main almond production region in the world. In this presumed pollinator-independent variety, we measured the effect of honey bees on fruit set, yield, and kernel nutritional quality at tree level. Fruit set was 60% higher in bee-pollinated than bee-isolated trees, which translated into a 20% increase in kernel yield. Despite its effect on almond production, there was no evidence that bee visitation affected almond nutritional quality. Based on these results, we recommend the use of bees, whether they are wild or managed, to maximize yield even in self-fertile almond varieties.

Pollination and resource limitation as interacting constraints on almond fruit set.

Pollination and resource availability are factors determining reproductive success of plants, and in agriculture these factors influence yield of fruit-bearing crops. Our understanding of the importance of crop pollination is fast improving, but less is known about how the interaction between pollination and resources constrains fruit production. We conducted an experiment with almond trees (Prunus dulcis) to examine how the number of flowers, light availability and competition for resources affected nut (fruit) production on individual spurs (fruit-bearing structures) exposed to open-pollination or hand-pollination. We found a positive relationship between flower number and nut number on spurs with up to four flowers, but no further benefit after four flowers, suggesting a resource threshold expressed by individual spurs. Spurs with few flowers increased the conversion rate of flowers to nuts when supplemented with hand-pollination, but spurs with more flowers were more likely to achieve the threshold number of nuts even under open-pollination. Our experiment included a further treatment involving spraying whole trees with pollen. This treatment reduced nut production by spurs with many flowers and high light availability, suggesting competition is experienced by well-resourced spurs when resources need to be shared among developing nuts across the whole tree. Our study supports the hypothesis that excess flower production in fruit trees increases the potential for fruit production when pollinator and resource availability is variable (bet-hedging). Spurs with more flowers typically produce more nuts (within a limited range), but only if both resources and pollen supply increase with flower number. For almond growers, a focus on maintaining high flower numbers, especially in high light regions of the canopy, is the foundation for high levels of production. Strategies to lift flower number and light are complicated by trade-offs inherent in tree architecture and orchard design. However, fruit set would be lifted above that achieved by current practice by an increase in the pollination rate of flowers.

Excessive nitrogen impairs hydraulics, limits photosynthesis, and alters the metabolic composition of almond trees.

Horticulture nitrogen (N) runoffs are major environmental and health concerns, but current farming practices cannot detect ineffective N applications. Hence, we set to recognize high N conditions and characterize their effects on the physiology of almond trees grown in drainage lysimeters. Water and nutrients mass balances exhibited that N benefitted almond trees in a limited range (below 60 mg N L-1 in irrigation), while higher N conditions (over a 100 mg N L-1) reduced evapotranspiration (ET) by 50% and inherently constrained N uptake. Respectively, whole-tree hydraulic conductance reduced by 37%, and photosynthesis by 17%, which implied that high N concentrations could damage trees. Through gas-chromatography, we realized that high N conditions also affected components of the citric acid cycle (TCA) and carbohydrates availability. Such changes in the metabolic composition of roots and leaves probably interfered with N assimilation and respiration. It also determined the proportions between N and starch in almond leaves, which formed a new index (N:ST) that starts at 0.4 in N deficiency and reaches 0.6-0.8 in optimal N conditions. Importantly, this index continues to increase in higher N conditions (as starch reduces) and essentially indicates to excessive N applications when it exceeds 1.1.

Bacterial population dynamics after foliar fertilization of almond leaves.

AIMS: To describe the effects of foliar fertilizer application on the bacterial populations of almond tree leaves. METHODS AND RESULTS: We applied a commercially available foliar fertilizer or a water control onto the leaves of almond trees and collected leaves after 1, 7, 14 and 56 days and examined their bacterial populations by 16S rRNA gene sequence analysis. After 1 day, we observed significant differences in 3 of the 4 predominant bacterial phyla, and 5 of the 13 predominant bacterial families. After 7 days, we observed significant differences in all of the predominant phyla, and 8 of the 13 predominant families. After 14 days, the number of significant differences decreased, and after 56 days only 2 of the 13 predominant families differed significantly. CONCLUSIONS: Foliar fertilization significantly altered the bacterial population structure of almond leaves as compared to the water control. While most of the observed perturbation was transient, significant differences remained after 56 days. SIGNIFICANCE AND IMPACT OF THE STUDY: This is the first report describing the effects of foliar fertilization on the bacterial populations of almond leaves and provides new insights as to how this process alters the leaf bacterial population structure.

Diurnal Variation in Nonstructural Carbohydrate Storage in Trees: Remobilization and Vertical Mixing.

Nonstructural carbohydrate (NSC) storage plays a critical role in tree function and survival, but understanding and predicting local NSC storage dynamics is challenging because NSC storage pools are dispersed throughout the complex architecture of trees and continuously exchange carbon between source and sink organs at different time scales. To address these knowledge gaps, characterization and understanding of NSC diel variation are necessary. Here, we analyzed diurnal NSC dynamics in the overall architecture of almond (Prunus dulcis) trees. We also analyzed the allocation of newly assimilated carbon using isotopic labeling. We show that both components of NSC (i.e. soluble carbohydrates and starch) are highly dynamic at the diurnal time scale and that these trends are influenced by tissue type, age, and/or position within the canopy. In leaves, starch reserves can be depleted completely during the night, while woody tissue starch levels may vary by more than 50% over a daily cycle. Recently assimilated carbon showed a dispersed downward allocation across the entire tree. NSC diurnal fluctuations within the tree's structure in combination with dispersed carbon allocation patterns provide evidence for the presence of vertical mixing and suggest that the xylem acts as a secondary NSC redistribution pathway.

Climate change threatens central Tunisian nut orchards.

Temperate deciduous trees can only be productive where winters are cold enough to meet their chilling needs. In the Mediterranean region, chill has traditionally been sufficient for many species, but this may change as temperatures increase. We explored the region's present and future suitability for temperate trees by quantifying chill for the Sfax region in central Tunisia, one of the warmest regions where temperate nuts are commercially grown. We assessed climatic risk by calculating historic chill (since 1973) and using a weather generator calibrated with local weather data (1973-2015) to produce 101 years of chill estimates (computed with the Dynamic Model) and 3 past and 72 future scenarios (for 2041-2070 and 2071-2100, using two representative concentration pathways: RCP4.5 and RCP8.5). For almonds and pistachios, we compared available chill during the chilling period with the species' estimated chilling requirements, and we computed the date by which sufficient chill was expected to have accumulated. Our findings indicated severe chill losses for all future scenarios. For all species, the current chill period is no longer expected to be sufficient for meeting chilling requirements in the future. Chill needs may still be fulfilled later in the year, especially for low-chill almonds, but this would result in delayed phenology, with possible adverse effects on productivity. Temperate nut production is thus unlikely to remain viable at this site, highlighting an urgent need to identify locally appropriate adaptation options. This challenge is likely shared by other warm production regions of temperate fruits and nuts around the world.

Insight into the physiological role of water absorption via the leaf surface from a rehydration kinetics perspective.

Soil water transported via the petiole is a primary rehydration pathway for leaves of water-stressed plants. Leaves may also rehydrate by absorbing water via their epidermal surfaces. The mechanisms and physiological relevance of this water pathway, however, remain unclear, as the associated hydraulic properties are unknown. To gain insight into the foliar water absorption process, we compared rehydration kinetics via the petiole and surface of Prunus dulcis and Quercus lobata leaves. Petiole rehydration could be described by a double exponential function suggesting that 2 partly isolated water pools exist in leaves of both species. Surface rehydration could be described by a logistic function, suggesting that leaves behave as a single water pool. Whereas full leaf rehydration via the petiole required approximately 20 min, it took over 150 and 300 min via the surface of P. dulcis and Q. lobata, respectively. Such differences were attributed to the high resistance imposed by the leaf surface and especially the cuticle. The minimum resistance to surface rehydration was estimated to be 6.6 × 102 (P. dulcis) and 2.6 × 103  MPa·m2 ·s·g-1 (Q. lobata), which is remarkably higher than estimated for petiole rehydration. These results are discussed in a physiological context.

Pollination Requirements of Almond (Prunus dulcis): Combining Laboratory and Field Experiments.

Almond (Prunus dulcis (Mill.) D. A. Webb; Rosales: Rosaceae) is a cash crop with an estimated global value of over seven billion U.S. dollars annually and commercial varieties are highly dependent on insect pollination. Therefore, the understanding of basic pollination requirements of the main varieties including pollination efficiency of honey bees (Apis mellifera, Linnaeus, Hymenoptera: Apidae) and wild pollinators is essential for almond production. We first conducted two lab experiments to examine the threshold number of pollen grains needed for successful pollination and to determine if varietal identity or diversity promotes fruit set and weight. Further, we examined stigma and ovules of flowers visited by Apis and non-Apis pollinators in the field to study the proportion of almond to non-almond pollen grains deposited, visitation time per flower visit, and tube set. Results indicate that the threshold for successful fertilization is around 60 pollen grains, but pollen can be from any compatible variety as neither pollen varietal identity nor diversity enhanced fruit set or weight. Andrena cerasifolii Cockerell (Hymenoptera: Andrenidae) was a more effective pollinator on a per single visit basis than Apis and syrphid flies. Nevertheless, Apis was more efficient than A. cerasifolii and syrphid flies as they spent less time on a flower during a single visit. Hence, planting with two compatible varieties and managing for both Apis and non-Apis pollinators is likely to be an optimal strategy for farmers to secure high and stable pollination success.

Examination of a Managed Pollinator Strategy for Almond Production Using Apis mellifera (Hymenoptera: Apidae) and Osmia lignaria (Hymenoptera: Megachilidae).

Pollination services provided by managed bees are essential for California almond (Prunus dulcis Mill.; Rosales: Rosaceae) production. Currently, pollination needs are met by rented or owned Apis mellifera L. (Hymenoptera: Apidae; honey bee) colonies. Excessive demand on a challenged A. mellifera industry to provide strong colonies in early spring has caused sharp increases in rental prices over the past decade, inviting the consideration of alternative pollinators in addition to, or in place of, A. mellifera. Osmia lignaria Say (Hymenoptera: Megachilidae; the blue orchard bee) is an excellent pollinator of fruit and nut trees, but its pollination impacts when used in tandem with A. mellifera have yet to be evaluated in commercial almond orchards. A 2-yr study was conducted in California orchards to compare almond pollination and production using A. mellifera as sole pollinator to an alternative practice of adding O. lignaria as a co-pollinator with A. mellifera. Almond orchard managerial decisions, such as for pesticide use and irrigation intensity, vary between almond growing regions because of local climates. Therefore, both north-central and southern sites of California's San Joaquin Valley are represented. We compared bee visitation, nut set, and nut yield between orchards and between tree rows within orchards. Also, O. lignaria reproductive success was recorded to assure that these bees remained in the orchards as pollinators and to assess the ability to sustain these bees under regional orchard conditions. We demonstrated that augmenting large commercial almond orchards with O. lignaria can significantly increase nut set and sometimes nut yield in both regions evaluated.

Honey bee (Apis mellifera) colony health and pathogen composition in migratory beekeeping operations involved in California almond pollination.

Honey bees are important pollinators of agricultural crops. Pathogens and other factors have been implicated in high annual losses of honey bee colonies in North America and some European countries. To further investigate the relationship between multiple factors, including pathogen prevalence and abundance and colony health, we monitored commercially managed migratory honey bee colonies involved in California almond pollination in 2014. At each sampling event, honey bee colony health was assessed, using colony population size as a proxy for health, and the prevalence and abundance of seven honey bee pathogens was evaluated using PCR and quantitative PCR, respectively. In this sample cohort, pathogen prevalence and abundance did not correlate with colony health, but did correlate with the date of sampling. In general, pathogen prevalence (i.e., the number of specific pathogens harbored within a colony) was lower early in the year (January-March) and was greater in the summer, with peak prevalence occurring in June. Pathogen abundance in individual honey bee colonies varied throughout the year and was strongly associated with the sampling date, and was influenced by beekeeping operation, colony health, and mite infestation level. Together, data from this and other observational cohort studies that monitor individual honey bee colonies and precisely account for sampling date (i.e., day of year) will lead to a better understanding of the influence of pathogens on colony mortality and the effects of other factors on these associations.

[Response of phenophase to meteorological conditions and flowering forecast model on Amygdalus communis in Shache County, Xinjiang, China].

Based on the phenophase data of Amygdalus communis and homochronous meteorological observation data at agrometeorological experimental station of Shache County during 2008-2013, the change characteristics of phenological period of A. communis and the effects of temperature and sunshine duration on them were analyzed. The results showed that before flowering, positive correlations existed among the first day of phenological phases, and after flowering, the correlations among the first day of phenological phases were mostly less. A significant positive correlation was observed between earlier bud flower swelling and the days of dormant period. and growth period, and a significant negative correlation existed between later bud flower swelling and the days of dormant period and growth period. Before fruit maturation, there was negative correlation between temperature and the interval days of phenological period, and after fruit maturation, the correlations were mostly positive. But the correlation between sunshine duration and the interval days of phenological period was positive before and after fruit maturation. The interval days from fruit maturation to the beginning date of leaf colour change had evident response to the average maximum temperature, and the interval days from the emergence of inflorescence to the ending data of flowering, and from the beginning date of leaf colour change to the ending date of leaf fall, had obvious response to sunshine duration. When the dormant period exceeded 30 days and the average daily temperature met the rang from -3.0 to -7.5 °C, A. communis would get into the flower swelling period after another 17-28 d. There were one-to-one correspondences between flower swelling, the beginning date of flowering, the beginning date of leaf colour change, the ending date of leaf fall, and the first pentad average temperature greater than or equal to 4 °C and pentad average maximum temperature greater than or equal to 12 °C, pentad average temperature greater than or equal to 14 °C and pentad average maximum temperature greater than or equal to 22 °C in spring, the first pentad temperature less than or equal to 10 °C and pentad average maximum temperature less than or equal to 18 °C in autumn the first pentad average temperature less than or equal to 1.9 °C in winter, respectively. By using partial least squares regression analysis, the first day of flowering forecast model of A. communis was established with good prediction.

Mongolian Almond (Prunus mongolica Maxim): The Morpho-Physiological, Biochemical and Transcriptomic Response to Drought Stress.

Prunus mongolica Maxim, which is widely established in the Gobi Desert, shows extreme tolerance to drought. However, there is a lack of available transcriptomic resources for this species related to its response to water deficiency. To investigate the mechanisms that allow P. mongolica to maintain growth in extremely arid environments, the response of P. mongolica seedlings to drought stress was analyzed using morphological, physiological, biochemical and high-throughput sequencing approaches. We generated 28,713,735 and 26,650,133 raw reads from no-stress control and drought-stressed P. mongolica seedlings, respectively. In total, we obtained 67,352 transcripts with an average length of 874.44 bp. Compared with the no-stress control, 3,365 transcripts were differentially expressed in the drought-stressed seedlings, including 55.75% (1,876 transcripts) up-regulated and 44.25% (1,489 transcripts) down-regulated transcripts. The photosynthesis response showed a decreasing tendency under drought stress, but the changes in the levels of hormones (auxins, cytokinins and abscisic acid) resulted in the closing of stomata and decreased cell enlargement and division; these changes were effective for promoting P. mongolica survival in Gobi Desert. Next, we analyzed the aquaporin and superoxide dismutase gene families due to their importance in plant resistance to drought stress. We found that all of the plasma membrane intrinsic protein transcripts were down-regulated in the drought-stressed treatment, whereas drought did not affect the expression of nodulin intrinsic protein or small basic intrinsic protein transcripts in P. mongolica seedlings. In addition, activation of iron superoxide dismutase transcription and enhanced transcription of manganese superoxide dismutase were observed in P. mongolica to promote tolerance of drought stress. This study identified drought response genes in P. mongolica seedlings. Our results provide a significant contribution to the understanding of how P. mongolica responds to drought stress at the transcriptome level, which may help to elucidate molecular mechanisms associated with the drought response of almond plants.