Potassium deficiency reduces grapevine transpiration through decreased leaf area and stomatal conductance.

Plants require potassium (K) to support growth and regulate hydraulics. Yet, K's effects on transpiration are still speculated. We hypothesized that K deficiency would limit grapevine water uptake by limiting canopy size and stomatal conductance (gs). Hence, we constructed large (2 m3) lysimeters and recorded vine transpiration for three years (2020-2022) under three fertilization application rates (8, 20, or 58 mg K L-1 in irrigation). Maximal K availability supported transpiration up to 75 L day-1, whereas K-deficient vines transpired only 60 L day-1 in midsummer. Limited vine growth and canopy size mainly accounted for reduced transpiration under low K conditions. Hence, considering K demand in addition to supply, we compared K deficiency effects on vines bearing 20 or 50 fruit clusters and found that reduced gs further limited transpiration when yields were high. Although fruits were strong K sinks, high yields did not alter K uptake because lower vegetative growth countered the additional K demands. Potassium deficiency leads to lower transpiration and productivity. Yet, internal mineral allocation compensates for fruit K uptake and masks biochemical indices or physiological proxies for K deficiency. Thus, decision support tools should integrate mineral availability, seasonal growth, and yield projections to determine grapevine water demands.

Losing ground: projections of climate-driven bloom shifts and their implications for the future of California's almond orchards.

Climate change is expected to impact the spring phenology of perennial trees, potentially altering the suitability of land for their cultivation. In this study, we investigate the effects of climate change on the bloom timing of almond orchards, focusing on California, the world's leading region for almond production. By analyzing historical climatic data, employing a model that considers hourly temperatures and fall non-structural carbohydrates to predict bloom dates, and examining various Coupled Model Intercomparison Project Phase 6 (CMIP6) scenarios, we assess the potential impacts of climate shifts on plant phenology and, consequently, on land suitability for almond farming. Our findings reveal that, within the next 30 years, the land suitable for almond production will not undergo significant changes. However, under unchanged emission scenarios, the available land to support almond orchard farming could decline between 48 to 73% by the end of the century. This reduction corresponds with an early shift in bloom time from the average Day of Year (DOY) 64 observed over the past 40 years to a projected earlier bloom between DOY 28-33 by 2100. These results emphasize the critical role climate shifts have in shaping future land use strategies for almond production in Central Valley, California. Consequently, understanding and addressing these factors is essential for the sustainable management and preservation of agricultural land, ensuring long-term food security and economic stability in the face of a rapidly changing climate.

The metabolic reserves, carbohydrate balance and nutritional status of jojoba (Simmondsia chinensis), in relation to its annual cycle and fruit load.

Jojoba (Simmondsia chinensis (Link) Schneider) holds high industrial value and an extended cultivation trend. Despite its increased importance, there is a lack of fundamental information about its metabolic reserves and development. Our objective was to characterise metabolite allocation and fluctuations in the carbohydrate and nutrient balance of jojoba plants, as affected by fruit load and the plant's annual cycle. Metabolite profiles were performed for each organ. Soluble carbohydrates (SC) and starch concentrations were surveyed in underground and aboveground organs of high-yield and fruit-removed plants. Simultaneously, nitrogen, potassium and phosphorus were determined in the leaves to evaluate the plant's nutritional status. We found that sucrose and pinitol were the most abundant sugars in all jojoba organs. Each sugar had a 'preferred' organ: glucose was accumulated mainly in the leaves, sucrose and pinitol in woody branches, and fructose in the trunk wood. We found that fruit load significantly influenced the carbohydrate levels in green branches, trunk wood and thin roots. The phenological stage strongly affected the SC-starch balance. Among the examined minerals, only the leaf potassium level was significantly influenced by fruit load. We conclude that jojoba's nutrient and carbohydrate balance is affected by fruit load and the phenological stage, and describe the organ-specific metabolic reserves.

Winding up the bloom clock-do sugar levels at senescence determine how trees respond to winter temperature?

Variable winter temperatures cause a year-to-year discrepancy in the phenology of deciduous trees. This implies that an intrinsic 'winter clock' synchronizes bloom with the progression of winter to spring. The carbohydrate-temperature (C-T) model established a mechanistic association between carbohydrate metabolism in dormant trees and hourly winter temperatures. Using historical winter temperature and bloom times of Prunus dulcis (Mill.) D. A. Webb (almond), Malus domestica L. (apple), Pistachia vera L. (pistachio) and Juglans regia L. (walnut) in California and Washington states, we parametrized species-specific metabolic parameters to the C-T model. There was a sound fit between actual and projected bloom dates with a deviation (root mean square error) of 4-7 days in all species. The parameterized model enabled us to study how the observed variability in soluble carbohydrate concentrations at senescence (SC0) could affect bloom time. The C-T model projected that low SC0 could advance, while high SC0 possibly delays, the bloom of the early blooming almond trees. In contrast, high SC0 would advance the bloom of apple, pistachio and walnut trees. These novel projections suggest that after experimental validation, SC0 could guide post-harvest farming applications that affect fall carbohydrate accumulation to mediate the effects of climate shifts.

Phosphorus fertilization induces nectar secretion for honeybee visitation and cross-pollination of almond trees.

Precise phosphorus (P) application requires a mechanistic understanding of mineral effects on crop biology and physiology. Photosynthate assimilation, metabolism, and transport require phosphorylation, and we postulated that P is critical for the bloom and fruit-set of almond trees that rely on stored carbohydrate reserves. Hence, we studied the growth, physiology and carbohydrate dynamics in 2-year-old almond trees irrigated with P concentrations between 1 mg l-1 and 20 mg l-1. Almond trees attained maximal photosynthesis, transpiration, and growth by 6 mg P l-1 irrigation. Nevertheless, almond trees continued to extract P in 10 mg P l-1 and 15 mg P l-1 irrigations, which corresponded to larger yields. We attributed the augmented productivity to increased fruit-set (59% between 6 mg P l-1 and 15 mg P l-1), caused by more frequent (29%) honeybee visits. High P improved pollinator visitation by enabling almond trees to utilize more of their starch reserves for nectar secretion (which increased by ~140% between 6 mg P l-1 and 15 mg P l-1). This work elucidates the benefits of P fertilization to plant-pollinator mutualism, critical to almond productivity, and reveals novel indices for optimal P application in almond orchards.

Wide vessels sustain marginal transpiration flux and do not optimize inefficient gas exchange activity under impaired hydraulic control and salinity.

Plants optimize water use and carbon assimilation via transient regulation of stomata resistance and by limiting hydraulic conductivity in a long-term response of xylem anatomy. We postulated that without effective hydraulic regulation plants would permanently restrain water loss and photosynthetic productivity under salt stress conditions. We compared wild-type tomatoes to a transgenic type (TT) with impaired stomatal control. Gas exchange activity, biomass, starch content, leaf area and root traits, mineral composition and main stems xylem anatomy and hydraulic conductivity were analyzed in plants exposed to salinities of 1 and 4 dS m-1 over 60 days. As the xylem cannot easily readjust to different environmental conditions, shifts in its anatomy and the permanent effect on plant hydraulic conductivity kept transpiration at lower levels under unstressed conditions and maintained it under salt-stress, while sustaining higher but inefficient assimilation rates, leading to starch accumulation and decreased plant biomass, leaf and root area and root length. Narrow conduits in unstressed TT plants were related to permanent restrain of hydraulic conductivity and plant transpiration. Under salinity, TT plants followed the atmospheric water demand, sustained similar transpiration rate from unstressed to salt-stressed conditions and possibly maintained hydraulic integrity, due to likely impaired hydraulic regulation, wider conduits and higher hydraulic conductivity. The accumulation of salts and starch in the TT plants was a strong evidence of salinity tolerance via osmotic regulation, also thought to help to maintain the assimilation rates and transpiration flux under salinity, although it was not translated into higher growth.

Comparison of phenological traits, growth patterns, and seasonal dynamics of non-structural carbohydrate in Mediterranean tree crop species.

Despite non-structural carbohydrate (NSC) importance for tree productivity and resilience, little is known about their seasonal regulations and trade-off with growth and reproduction. We characterize the seasonal dynamics of NSC in relation to the aboveground phenology and temporal growth patterns of three deciduous Mediterranean species: almond (Prunus dulcis (Mill.) D. A. Webb), walnut (Juglans regia L.) and pistachio (Pistacia vera L.). Seasonal dynamics of NSC were synchronous between wood tissues from trunk, branches and twigs. Almond had almost identical levels and patterns of NSC variation in twigs, branches and trunks whereas pistachio and walnut exhibited clear concentration differences among plant parts whereby twigs had the highest and most variable NSC concentration, followed by branches and then trunk. While phenology had a significant influence on NSC seasonal trends, there was no clear trade-off between NSC storage and growth suggesting that both were similarly strong sinks for NSC. A temporal trade-off observed at the seasonal scale was influenced by the phenology of the species. We propose that late senescing species experience C allocation trade-off at the end of the growing season because of C-limiting thermal conditions and priority allocation to storage in order to survive winter.

High Nitrogen Availability Limits Photosynthesis and Compromises Carbohydrate Allocation to Storage in Roots of Manihot esculenta Crantz.

Cassava (M. esculenta Crantz), feeding countless people and attracting markets worldwide, is a model for traditional crops that need physiology-based fertigation (fertilization through irrigation) standards in intensive cultivation. Hence, we studied the effects of 10 to 200 mg L-1 nitrogen (N) fertigation on growth and yields of cassava and targeted alterations in their photosynthetic, transpiration, and carbohydrate management. We found that increasing irrigation N from 10 to 70 mg L-1 increased cassava's photosynthesis and transpiration but supported only the canopy's growth. At 100 mg N L-1 cassava reached a threshold of sugar in leaves (∼47 mg g-1), began to accumulate starch and supported higher yields. Yet, at 200 mg N L-1, the canopy became too demanding and plants had to restrain transpiration, reduce photosynthesis, decrease carbohydrates, and finally lower yields. We concluded that the phases of cassava response to nitrogen are: 1) growth that does not support yields at low N, 2) productive N application, and 3) excessive use of N. Yet traditional leaf mineral analyses fail to exhibit these responses, and therefore we propose a simple and inexpensive carbohydrate measurement to guide a precise use of N.

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.

Sodium interception by xylem parenchyma and chloride recirculation in phloem may augment exclusion in the salt tolerant Pistacia genus: context for salinity studies on tree crops.

Working in tandem with root exclusion, stems may provide salt-tolerant woody perennials with some additional capacity to restrict sodium (Na) and chloride (Cl) accumulation in leaves. The Pistacia genus, falling at the nexus of salt tolerance and human intervention, provided an ideal set of organisms for studying the influences of both variable root exclusion and potentially variable discontinuities at the bud union on stem processes. In three experiments covering a wide range of salt concentrations (0 to 150 mM NaCl) and tree ages (1, 2 and 10 years) as well as nine rootstock-scion combinations we show that proportional exclusion of both Na and Cl reached up to ~85% efficacy, but efficacy varied by both rootstock and budding treatment. Effective Na exclusion was augmented by significant retrieval of Na from the xylem sap, as evidenced by declines in the Na concentrations of both sap and wood tissue along the transpiration stream. However, while we observed little to no differences between the concentrations of the two ions in leaves, analogous declines in sap concentrations of Cl were not observed. We conclude that some parallel but separate mechanism must be acting on Cl to provide leaf protection from toxicity specific to this ion and suggest that this mechanism is recirculation of Cl in the phloem. The presented findings underline the importance of holistic assessments of salt tolerance in woody perennials. In particular, greater emphasis might be placed on the dynamics of salt sequestration in the significant storage volumes offered by the stems of woody perennials and on the potential for phloem discontinuity introduced with a bud/graft union.

Extreme mid-winter drought weakens tree hydraulic-carbohydrate systems and slows growth.

Rising temperatures and extended periods of drought compromise tree hydraulic and carbohydrate systems, threatening forest health globally. Despite winter's biological significance to many forests, the effects of warmer and dryer winters on tree hydraulic and carbohydrate status have largely been overlooked. Here we report a sharp and previously unknown decline in stem water content of three conifer species during California's anomalous 2015 mid-winter drought that was followed by dampened spring starch accumulation. Recent precipitation and seasonal vapor pressure deficit (VPD) anomaly, not absolute VPD, best predicted the hydraulic patterns observed. By linking relative water content and hydraulic conductivity (Kh ), we estimated that stand-level Kh declined by 52% during California's 2015 mid-winter drought, followed by a 50% reduction in spring starch accumulation. Further examination of tree increment records indicated a concurrent decline of growth with rising mid-winter, but not summer, VPD anomaly. Thus, our findings suggest a seasonality to tree hydraulic and carbohydrate declines, with consequences for annual growth rates, raising novel physiological and ecological questions about how rising winter temperatures will affect forest vitality as climate changes.

Temperature gradients assist carbohydrate allocation within trees.

Trees experience two distinct environments: thermally-variable air and thermally-buffered soil. This generates intra-tree temperature gradients, which can affect carbon metabolism and water transport. In this study, we investigated whether carbohydrate allocation within trees is assisted by temperature gradients. We studied pistachio (Pistacia integerrima) to determine: (1) temperature-induced variation in xylem sugar concentration in excised branches; (2) changes in carbon allocation in young trees under simulated spring and fall conditions; and (3) seasonal variability of starch levels in mature orchard trees under field conditions. We found that warm branches had less sugar in perfused sap than cold branches due to increasing parenchyma storage. Simulated spring conditions promoted allocation of carbohydrates from cold roots to warm canopy and explained why starch levels surged in canopies of orchard trees during early spring. This driving force of sugar transport is interrupted in fall when canopies are colder than roots and carbohydrate redistribution is compartmentalized. On the basis of these findings, we propose a new mechanistic model of temperature-assisted carbohydrate allocation that links environmental cues and tree phenology. This data-enabled model provides insights into thermal "fine-tuning" of carbohydrate metabolism and a warning that the physiological performance of trees might be impaired by climatic changes.

Spring bud growth depends on sugar delivery by xylem and water recirculation by phloem Münch flow in Juglans regia.

MAIN CONCLUSION: During spring, bud growth relies on long-distance transport of remotely stored carbohydrates. A new hypothesis suggests this transport is achieved by the interplay of xylem and phloem. During the spring, carbohydrate demand of developing buds often exceeds locally available storage, thus requiring the translocation of sugars from distant locations like limbs, stems and roots. Both the phloem and xylem have the capacity for such long-distance transport, but their functional contribution is unclear. To address this ambiguity, the spatial and temporal dynamics of carbohydrate availability in extension shoots of Juglans regia L. were analyzed. A significant loss of extension shoot carbohydrates in remote locations was observed while carbohydrate availability near the buds remained unaffected. This pattern of depletion of carbohydrate reserves supports the notion of long-distance translocation. Girdling and dye perfusion experiments were performed to assess the role of phloem and xylem in the transport of carbohydrate and water towards the buds. Girdling caused a decrease in non-structural carbohydrate concentration above the point of girdling and an unexpected concurrent increase in water content associated with impeded xylem transport. Based on experimental observations and modeling, we propose a novel mechanism for maintenance of spring carbohydrate translocation in trees where xylem transports carbohydrates and this transport is maintained with the recirculation of water by phloem Münch flow. Phloem Münch flow acts as a pump for generating water flux in xylem and allows for transport and mobilization of sugars from distal locations prior to leaves photosynthetic independence and in the absence of transpiration.

Acclimation of Pistacia integerrima trees to frost in semi-arid environments depends on autumn's drought.

Main conclusion Cold acclimation is revealed through induced stem respiration during pre-winter frost of native Pistacia integerrima trees in continental semi-arid environments. Semi-arid environments challenge vegetation by simultaneous abiotic stresses. In this study, we examine the combined effects of water stress and frost on the physiology of Pistacia integerrima stems. This species is native to semi-arid environments where drought and frost frequently co-occur. We quantified carbohydrates and proline in P. integerrima stems responding to frost and experiencing water potentials between -0.2 and -1.8 MPa. We report that dehydrated trees (i.e., Ψstem <=-1 MPa) had more soluble sugars and proline than the well-watered trees (-0.2 MPa). The dehydrated trees also froze at lower temperatures and were less damaged by freezing. Interestingly, we observed a significant increase in stem CO2 efflux at near-freezing temperatures that could be linked to frost protection. This novel finding challenges current paradigm of plant respiration-kinetics which predicts, according to Arrhenius equation, lower respiration rates during frost. Our results support the notion that drought and frost are analogous stresses that can independently activate corresponding physiological processes in trees and amplify protection. This inevitable stress response 'collaboration' may be the key to understanding how non-dormant perennial plants survive the highly variable weather patterns of early winters in semi-arid environments.

Biotic degradation at night, abiotic degradation at day: positive feedbacks on litter decomposition in drylands.

The arid and semi-arid drylands of the world are increasingly recognized for their role in the terrestrial net carbon dioxide (CO2 ) uptake, which depends largely on plant litter decomposition and the subsequent release of CO2 back to the atmosphere. Observed decomposition rates in drylands are higher than predictions by biogeochemical models, which are traditionally based on microbial (biotic) degradation enabled by precipitation as the main mechanism of litter decomposition. Consequently, recent research in drylands has focused on abiotic mechanisms, mainly photochemical and thermal degradation, but they only partly explain litter decomposition under dry conditions, suggesting the operation of an additional mechanism. Here we show that in the absence of precipitation, absorption of dew and water vapor by litter in the field enables microbial degradation at night. By experimentally manipulating solar irradiance and nighttime air humidity, we estimated that most of the litter CO2 efflux and decay occurring in the dry season was due to nighttime microbial degradation, with considerable additional contributions from photochemical and thermal degradation during the daytime. In a complementary study, at three sites across the Mediterranean Basin, litter CO2 efflux was largely explained by litter moisture driving microbial degradation and ultraviolet radiation driving photodegradation. We further observed mutual enhancement of microbial activity and photodegradation at a daily scale. Identifying the interplay of decay mechanisms enhances our understanding of carbon turnover in drylands, which should improve the predictions of the long-term trend of global carbon sequestration.

Bark water uptake promotes localized hydraulic recovery in coastal redwood crown.

Coastal redwood (Sequoia sempervirens), the world's tallest tree species, rehydrates leaves via foliar water uptake during fog/rain events. Here we examine if bark also permits water uptake in redwood branches, exploring potential flow mechanisms and biological significance. Using isotopic labelling and microCT imaging, we observed that water entered the xylem via bark and reduced tracheid embolization. Moreover, prolonged bark wetting (16 h) partially restored xylem hydraulic conductivity in isolated branch segments and whole branches. Partial hydraulic recovery coincided with an increase in branch water potential from about -5.5 ± 0.4 to -4.2 ± 0.3 MPa, suggesting localized recovery and possibly hydraulic isolation. As bark water uptake rate correlated with xylem osmotic potential (R(2) = 0.88), we suspect a symplastic role in transferring water from bark to xylem. Using historical weather data from typical redwood habitat, we estimated that bark and leaves are wet more than 1000 h per year on average, with over 30 events being sufficiently long (>24 h) to allow for bark-assisted hydraulic recovery. The capacity to uptake biologically meaningful volumes of water via bark and leaves for localized hydraulic recovery throughout the crown during rain/fog events might be physiologically advantageous, allowing for relatively constant transpiration.

Frost Induces Respiration and Accelerates Carbon Depletion in Trees.

Cellular respiration depletes stored carbohydrates during extended periods of limited photosynthesis, e.g. winter dormancy or drought. As respiration rate is largely a function of temperature, the thermal conditions during such periods may affect non-structural carbohydrate (NSC) availability and, ultimately, recovery. Here, we surveyed stem responses to temperature changes in 15 woody species. For two species with divergent respirational response to frost, P. integerrima and P. trichocarpa, we also examined corresponding changes in NSC levels. Finally, we simulated respiration-induced NSC depletion using historical temperature data for the western US. We report a novel finding that tree stems significantly increase respiration in response to near freezing temperatures. We observed this excess respiration in 13 of 15 species, deviating 10% to 170% over values predicted by the Arrhenius equation. Excess respiration persisted at temperatures above 0 °C during warming and reoccurred over multiple frost-warming cycles. A large adjustment of NSCs accompanied excess respiration in P. integerrima, whereas P. trichocarpa neither excessively respired nor adjusted NSCs. Over the course of the years included in our model, frost-induced respiration accelerated stem NSC consumption by 8.4 mg (glucose eq.) cm(-3) yr(-1) on average in the western US, a level of depletion that may continue to significantly affect spring NSC availability. This novel finding revises the current paradigm of low temperature respiration kinetics.

Direct in vivo evidence of immense stem water exploitation in irrigated date palms.

During the summer, evaporative demand at midday often exceeds the transport capacity of most desert plants. However, date palms maintain their ecological dominance with sustained and uniquely high rates of transpiration. This high rate of flow cannot be attributed to soil water supply alone. In order to quantify intra-plant water allocation in irrigated date palms, three water-sensing techniques have been incorporated: heat dissipation, gravimetric sampling, and time domain reflectrometry. Each of these methods has known limitations but their integration resulted in a quantitative in vivo accounting of the date palm diurnal and seasonal water mass balance. By incorporating these methods it was possible to determine that date palms substantially rely on the exploitation and recharge of the stem reservoir in their water budget. The stem of mature date palms can hold up to 1 m(3) of water and supply 25% of daily transpiration (i.e. 5000 l of water in 100 d of summer). The internal stem water reservoir is consistently recharged by over 50 l per night which allows for successive daytime reuse throughout the entire growing season. More broadly, these findings suggest that internal water allocation and night-time soil-water availability could provide useful information for improving date palm irrigation practices.

Estimating sap flux densities in date palm trees using the heat dissipation method and weighing lysimeters.

In a world of diminishing water reservoirs and a rising demand for food, the practice and development of water stress indicators and sensors are in rapid progress. The heat dissipation method, originally established by Granier, is herein applied and modified to enable sap flow measurements in date palm trees in the southern Arava desert of Israel. A long and tough sensor was constructed to withstand insertion into the date palm's hard exterior stem. This stem is wide and fibrous, surrounded by an even tougher external non-conducting layer of dead leaf bases. Furthermore, being a monocot species, water flow does not necessarily occur through the outer part of the palm's stem, as in most trees. Therefore, it is highly important to investigate the variations of the sap flux densities and determine the preferable location for sap flow sensing within the stem. Once installed into fully grown date palm trees stationed on weighing lysimeters, sap flow as measured by the modified sensors was compared with the actual transpiration. Sap flow was found to be well correlated with transpiration, especially when using a recent calibration equation rather than the original Granier equation. Furthermore, inducing the axial variability of the sap flux densities was found to be highly important for accurate assessments of transpiration by sap flow measurements. The sensors indicated no transpiration at night, a high increase of transpiration from 06:00 to 09:00, maximum transpiration at 12:00, followed by a moderate reduction until 08:00; when transpiration ceased. These results were reinforced by the lysimeters' output. Reduced sap flux densities were detected at the stem's mantle when compared with its center. These results were reinforced by mechanistic measurements of the stem's specific hydraulic conductivity. Variance on the vertical axis was also observed, indicating an accelerated flow towards the upper parts of the tree and raising a hypothesis concerning dehydrating mechanisms of the date palm tree. Finally, the sensors indicated reduction in flow almost immediately after irrigation of field-grown trees was withheld, at a time when no climatic or phenological conditions could have led to reduction in transpiration.