Lethal combination for seedlings: extreme heat drives mortality of drought-exposed high-elevation pine seedlings.

BACKGROUND AND AIMS: Hotter drought- and biotically-driven tree mortality are expected to increase with climate change in much of the western United States, and species persistence will depend upon ongoing establishment under novel conditions or migration to track ecological niche requirements. High-elevation tree species may be particularly vulnerable to increasing water stress as snowpack declines, increasing the potential for adult mortality and simultaneous regeneration failures. Seedling survival will be determined by ecophysiological limitations in response to changing water availability and temperature. METHODS: We exposed seedlings from populations of Pinus longaeva, Pinus flexilis, and Pinus albicaulis to severe drought and concurrent temperature stress in common gardens testing timing of drought onset under two different temperature regimes. We monitored seedling functional traits, physiological function, and survival. KEY RESULTS: The combined stressors of water limitation and extreme heat led to conservative water use strategies and declines in physiological function, with these joint stressors ultimately exceeding species' tolerances and leading to complete episodic mortality across all species. Growing conditions were the primary determinant of seedling trait expression, with seedlings exhibiting more drought-resistant traits such as lower specific leaf area in the hottest, driest treatment conditions. Water stress-induced stomatal closure was also widely apparent. Under adequate soil moisture, seedlings endured prolonged exposure to high air and surface temperatures, suggesting broad margins for survival. CONCLUSIONS: The critical interaction between soil moisture and temperature suggests that rising temperatures will exacerbate growing season moisture stress. Our results highlight the importance of local conditions over population- and species-level influences in shaping strategies for stress tolerance and resistance to desiccation at this early life stage. By quantifying some of the physiological consequences of drought and heat that lead to seedling mortality, we can better understand the future effects of global change on the composition and distribution of high-elevation conifer forests.

Floral hemp (Cannabis sativa L.) responses to nitrogen fertilization under field conditions in the high desert.

While most studies on floral hemp (Cannabis sativa L.) concur that additions of nitrogen (N) increase plant growth, the performance of floral hemp is heavily influenced by environmental conditions, management and cultivar selection. In regions with a short growing season, the availability of soil N may determine plant developmental rates, final inflorescence biomass and cannabinoid concentrations, but no studies have addressed this for field-grown hemp under high-desert conditions. This field study evaluated the effect of no supplemental N and N fertilization at 90 kg ha-1 on three hemp cultivars (Berry Blossom, Red Bordeaux, and Tahoe Cinco) in Northern Nevada. N increased plant height, canopy cover, stem diameter and shoot biomass, but other physiological parameters were dependent on cultivar. For instance, inflorescence biomass and inflorescence-to-shoot ratio in Red Bordeaux was not affected by N fertilization. Similarly, cannabinoid concentrations were affected by timing of harvest and cultivar but not by N treatment. We evaluated the use of a SPAD meter for ease of determining leaf N deficiency, and correlations with leaf chlorophyll content showed that the SPAD meter was a reliable tool in two cultivars but not in Tahoe Cinco. N treatment increased overall CBD yield, which was driven by increases in inflorescence biomass. Tahoe Cinco was the best CBD yielding cultivar, as it maintained a high inflorescence-to-shoot ratio regardless of N treatment. Our study suggests that even though hemp may have a positive response to soil N management, adjustments based on genotype by environment interaction should be aimed at maximizing cannabinoid yield either by increasing biomass and/or CBD concentrations as long as THC levels are within the permissible <0.3% for U.S. industrial hemp cultivation.

Salt tolerance mechanisms in the Lycopersicon clade and their trade-offs.

Salt stress impairs growth and yield in tomato, which is mostly cultivated in arid and semi-arid areas of the world. A number of wild tomato relatives (Solanum pimpinellifolium, S. pennellii, S. cheesmaniae and S. peruvianum) are endemic to arid coastal areas and able to withstand higher concentration of soil salt concentrations, making them a good genetic resource for breeding efforts aimed at improving salt tolerance and overall crop improvement. However, the complexity of salt stress response makes it difficult to introgress tolerance traits from wild relatives that could effectively increase tomato productivity under high soil salt concentrations. Under commercial production, biomass accumulation is key for high fruit yields, and salt tolerance management strategies should aim to maintain a favourable plant water and nutrient status. In this review, we first compare the effects of salt stress on the physiology of the domesticated tomato and its wild relatives. We then discuss physiological and energetic trade-offs for the different salt tolerance mechanisms found within the Lycopersicon clade, with a focus on the importance of root traits to sustain crop productivity.

Leaf Gas Exchange Performance of Ten Quinoa Genotypes under a Simulated Heat Wave.

Quinoa (Chenopodium quinoa Willd.) is a highly nutritious crop that is resilient to a wide range of abiotic stresses; however, sensitivity to high temperatures is regarded as an impediment to adoption in regions prone to heat waves. Heat stress is usually associated with a decrease in crop reproductive capacity (e.g., pollen viability), yet little is known about how leaf physiological performance of quinoa is affected by high temperatures. Several trials were conducted to understand the effect of high temperatures, without confounding stressors such as drought, on ten selected quinoa genotypes considered to encompass heat sensitive and heat tolerant plant material. Plants were grown under favorable temperatures and exposed to two temperature treatments over four consecutive days. The heat treatment simulated heat waves with maximum and minimum temperatures higher during the day and night, while the control treatment was maintained under favorable temperatures (maximum and minimum temperatures for 'Heat': 45/30 °C and 'Control': 20/14 °C). Leaf gas exchange (day), chlorophyll fluorescence (predawn and day) and dark respiration (night) were measured. Results show that most quinoa genotypes under the heat treatment increased their photosynthetic rates and stomatal conductance, resulting in a lower intrinsic water use efficiency. This was partly corroborated by an increase in the maximum quantum yield of photosystem II (Fv/Fm). Dark respiration decreased under the heat treatment in most genotypes, and temperature treatment did not affect aboveground biomass by harvest (shoot and seeds). These results suggest that heat stress alone favors increases in leaf carbon assimilation capacity although the tradeoff is higher plant water demand, which may lead to plant water stress and lower yields under non-irrigated field conditions.

Tomato Rootstocks Mediate Plant-Water Relations and Leaf Nutrient Profiles of a Common Scion Under Suboptimal Soil Temperatures.

Environments with short growing seasons and variable climates can have soil temperatures that are suboptimal for chilling-sensitive crops. These conditions can adversely affect root growth and physiological performance thus impairing water and nutrient uptake. Four greenhouse trials and a field study were conducted to investigate if rootstocks can enhance tomato performance under suboptimal soil temperatures (SST). In a controlled greenhouse environment, we exposed four commercial rootstocks (Estamino, Maxifort, RST-04-106-T, and Supernatural) grafted with a common scion (cv. BHN-589) to optimal (mean: 24°C) and SST (mean: 13.5°C) and compared their performance with the non-grafted BHN-589 cultivar. Several root and shoot physiological traits were evaluated: root hydraulic conductivity and conductance, root anatomy, leaf gas exchange, leaf δ13C, shoot C and N, and biomass. Under field conditions, the same five phenotypes were evaluated for canopy growth, normalized difference vegetation index (NDVI), leaf nutrients, biomass, and yield. Under SST, root hydraulic conductivity (Lp) and conductance (K R), stomatal conductance (g s), and plant biomass decreased. Hydrostatic Lp decreased more than osmotic Lp (Lp ∗ hyd: 39-65%; Lp ∗ os: 14-40%) and some of the reduced conductivity was explained by the increased cortex area of primary roots observed under SST (67-140%). Under optimal soil temperatures, all rootstocks conferred higher g s than the non-grafted cultivar, but only two rootstocks maintained higher g s under SST. All phenotypes showed greater reductions in shoot biomass than root biomass resulting in greater (∼20%) root-to-shoot ratios. In the field, most grafted phenotypes increased early canopy cover, NDVI, shoot biomass, and fruit yield. Greenhouse results showed that Lp ∗ os may be less affected by SST than Lp ∗ hyd and that reductions in Lp may be offset by enhanced root-to-shoot ratios. We show that some commercial rootstocks possess traits that maintained better rates of stomatal conductance and shoot N content, which can contribute toward better plant establishment and improved performance under SST.

Modeling vegetative vigour in grapevine: unraveling underlying mechanisms.

Mechanistic modeling constitutes a powerful tool to unravel complex biological phenomena. This study describes the construction of a mechanistic, dynamic model for grapevine plant growth and canopy biomass (vigor). To parametrize and validate the model, the progeny from a cross of Ramsey (Vitis champinii) × Riparia Gloire (V. riparia) was evaluated. Plants with different vigor were grown in a greenhouse during the summer of 2014 and 2015. One set of plants was grafted with Cabernet Sauvignon. Shoot growth rate (b), leaf area (LA), dry biomass, whole plant and root specific hydraulic conductance (kH and Lpr), stomatal conductance (gs), and water potential (Ψ) were measured. Partitioning indices and specific leaf area (SLA) were calculated. The model includes an empirical fit of a purported seasonal pattern of bioactive GAs based on published seasonal evolutionary levels and reference values. The model provided a good fit of the experimental data, with R = 0.85. Simulation of single trait variations defined the individual effect of each variable on vigor determination. The model predicts, with acceptable accuracy, the vigor of a young plant through the measurement of Lpr and SLA. The model also permits further understanding of the functional traits that govern vigor, and, ultimately, could be considered useful for growers, breeders and those studying climate change.

Differences in hydraulic traits of grapevine rootstocks are not conferred to a common Vitis vinifera scion.

Cultivars of grapevine are commonly grafted onto rootstocks to improve resistance against biotic and abiotic stress, however, it is not clear whether known differences in hydraulic traits are conferred from rootstocks to a common scion. We recently found that Vitis riparia and Vitis champinii differed in drought-induced embolism susceptibility and repair, which was related to differences in root pressure generation after rewatering (Knipfer et al. 2015). In the present study, we tested whether these and other physiological responses to drought are conferred to a common V. vinifera scion (Cabernet Sauvignon) grafted on V. riparia and V. champinii rootstocks. We measured xylem embolism formation/repair using in vivo microCT imaging, which was accompanied with analysis of leaf gas exchange, osmotic adjustment and root pressure. Our data indicate that differences in scion physiological behaviour for both rootstock combinations were negligible, suggesting that the sensitivity of Cabernet Sauvignon scion to xylem embolism formation/repair, leaf gas exchange and osmotic adjustment is unaffected by either V. riparia or V. champinii rootstock in response to drought stress.

Quinoa Abiotic Stress Responses: A Review.

Quinoa (Chenopodium quinoa Willd.) is a genetically diverse Andean crop that has earned special attention worldwide due to its nutritional and health benefits and its ability to adapt to contrasting environments, including nutrient-poor and saline soils and drought stressed marginal agroecosystems. Drought and salinity are the abiotic stresses most studied in quinoa; however, studies of other important stress factors, such as heat, cold, heavy metals, and UV-B light irradiance, are severely limited. In the last few decades, the incidence of abiotic stress has been accentuated by the increase in unpredictable weather patterns. Furthermore, stresses habitually occur as combinations of two or more. The goals of this review are to: (1) provide an in-depth description of the existing knowledge of quinoa's tolerance to different abiotic stressors; (2) summarize quinoa's physiological responses to these stressors; and (3) describe novel advances in molecular tools that can aid our understanding of the mechanisms underlying quinoa's abiotic stress tolerance.

Variations in xylem embolism susceptibility under drought between intact saplings of three walnut species.

A germplasm collection containing varied Juglans genotypes holds potential to improve drought resistance of plant materials for commercial production. We used X-ray computed microtomography to evaluate stem xylem embolism susceptibility/repair in relation to vessel anatomical features (size, arrangement, connectivity and pit characteristics) in 2-year-old saplings of three Juglans species. In vivo analysis revealed interspecific variations in embolism susceptibility among Juglans microcarpa, J. hindsii (both native to arid habitats) and J. ailantifolia (native to mesic habitats). Stem xylem of J. microcarpa was more resistant to drought-induced embolism as compared with J. hindsii and J. ailantifolia (differences in embolism susceptibility among older and current year xylem were not detected in any species). Variations in most vessel anatomical traits were negligible among the three species; however, we detected substantial interspecific differences in intervessel pit characteristics. As compared with J. hindsii and J. ailantifolia, low embolism susceptibility in J. microcarpa was associated with smaller pit size in larger diameter vessels, a smaller area of the shared vessel wall occupied by pits, lower pit frequency and no changes in pit characteristics as vessel diameters increased. Changes in amount of embolized vessels following 40 days of re-watering were minor in intact saplings of all three species highlighting that an embolism repair mechanism did not contribute to drought recovery. In conclusion, our data indicate that interspecific variations in drought-induced embolism susceptibility are associated with species-specific pit characteristics, and these traits may provide a future target for breeding efforts aimed at selecting walnut germplasm with improved drought resistance.

Effects of arbuscular mycorrhizae on tomato yield, nutrient uptake, water relations, and soil carbon dynamics under deficit irrigation in field conditions.

Plant strategies to cope with future droughts may be enhanced by associations between roots and soil microorganisms, including arbuscular mycorrhizal (AM) fungi. But how AM fungi affect crop growth and yield, together with plant physiology and soil carbon (C) dynamics, under water stress in actual field conditions is not well understood. The well-characterized mycorrhizal tomato (Solanum lycopersicum L.) genotype 76R (referred to as MYC+) and the mutant nonmycorrhizal tomato genotype rmc were grown in an organic farm with a deficit irrigation regime and control regime that replaced evapotranspiration. AM increased marketable tomato yields by ~25% in both irrigation regimes but did not affect shoot biomass. In both irrigation regimes, MYC+ plants had higher plant nitrogen (N) and phosphorus (P) concentrations (e.g. 5 and 24% higher N and P concentrations in leaves at fruit set, respectively), 8% higher stomatal conductance (gs), 7% higher photosynthetic rates (Pn), and greater fruit set. Stem water potential and leaf relative water content were similar in both genotypes within each irrigation regime. Three-fold higher rates of root sap exudation in detopped MYC+ plants suggest greater capacity for water uptake through osmotic driven flow, especially in the deficit irrigation regime in which root sap exudation in rmc was nearly absent. Soil with MYC+ plants also had slightly higher soil extractable organic C and microbial biomass C at anthesis but no changes in soil CO2 emissions, although the latter were 23% lower under deficit irrigation. This study provides novel, field-based evidence for how indigenous AM fungi increase crop yield and crop water use efficiency during a season-long deficit irrigation and thus play an important role in coping with increasingly limited water availability in the future.

Use of introgression lines to determine the ecophysiological basis for changes in water use efficiency and yield in California processing tomatoes.

Field and greenhouse studies examined the effects of growth habit and chloroplast presence in leaf veins for their role in increasing agronomic water use efficiency and yields of California modern processing tomato (Solanum lycopersicum L.) cultivars. Five introgression lines (ILs), made with Solanum pennellii Cor. in the genetic background of cultivar M82, differ in genes that map to a region on Chromosome 5, including the SP5G gene (determinate vs. semideterminate (Det vs. SemiDet)) and the obv gene (presence (obscure) vs. absence (clear) of leaf vein chloroplasts (Obs vs. Clr)). The five ILs and M82 represented three of the four gene combinations (Det-Clr was unavailable). Det-Obs ILs had less leaf, stem and total aboveground biomass with earlier fruit set and ripening than SemiDet-Clr ILs. By harvest, total fruit biomass was not different among ILs. Photosynthetic rates and stomatal conductance were 4-7% and 13-26% higher, respectively, in Det-Obs ILs than SemiDet-Clr ILs. SemiDet-Obs ILs were intermediate for growth and gas exchange variables. The Det-Obs ILs had lower leaf N concentration and similar chlorophyll content per leaf area (but slightly higher per leaf mass) than SemiDet-Clr ILs. The Obs trait was associated with gains in leaf gas exchange-related traits. This study suggests that a more compact growth habit, less leaf biomass and higher C assimilation capacity per leaf area were relevant traits for the increased yields in cultivars with determinate growth. Developing new introgression libraries would contribute to understanding the multiple trait effects of desirable phenotypes.

Tomato root transcriptome response to a nitrogen-enriched soil patch.

BACKGROUND: Nitrogen (N), the primary limiting factor for plant growth and yield in agriculture, has a patchy distribution in soils due to fertilizer application or decomposing organic matter. Studies in solution culture over-simplify the complex soil environment where microbial competition and spatial and temporal heterogeneity challenge roots' ability to acquire adequate amounts of nutrients required for plant growth. In this study, various ammonium treatments (as 15N) were applied to a discrete volume of soil containing tomato (Solanum lycopersicum) roots to simulate encounters with a localized enriched patch of soil. Transcriptome analysis was used to identify genes differentially expressed in roots 53 hrs after treatment. RESULTS: The ammonium treatments resulted in significantly higher concentrations of both ammonium and nitrate in the patch soil. The plant roots and shoots exhibited increased levels of 15N over time, indicating a sustained response to the enriched environment. Root transcriptome analysis identified 585 genes differentially regulated 53 hrs after the treatments. Nitrogen metabolism and cell growth genes were induced by the high ammonium (65 mug NH4+-N g-1 soil), while stress response genes were repressed. The complex regulation of specific transporters following the ammonium pulse reflects a simultaneous and synergistic response to rapidly changing concentrations of both forms of inorganic N in the soil patch. Transcriptional analysis of the phosphate transporters demonstrates cross-talk between N and phosphate uptake pathways and suggests that roots increase phosphate uptake via the arbuscular mycorrhizal symbiosis in response to N. CONCLUSION: This work enhances our understanding of root function by providing a snapshot of the response of the tomato root transcriptome to a pulse of ammonium in a complex soil environment. This response includes an important role for the mycorrhizal symbiosis in the utilization of an N patch.