Analysis of Growth and Water Relations of Tomato Fruits in Relation to Air Vapor Pressure Deficit and Plant Fruit Load

The influence of air vapor pressure deficit (VPD) and plant fruit load on the expansion and water relations of young tomato fruits grown in a glasshouse were evaluated under summer Mediterranean conditions. The contributions of phloem, xylem and transpiration fluxes to the fruit volume increase were estimated at an hourly scale from the growth curves of intact, heat-girdled and detached fruits, measured using displacement transducers. High VPD conditions reduced the xylem influx and increased the fruit transpiration, but hardly affected the phloem influx. Net water accumulation and growth rate were reduced, and a xylem efflux even occurred during the warmest and driest hours of the day. Changes in xylem flux could be explained by variations in the gradient of water potential between stem and fruit, due to changes in stem water potential. Misting reduced air VPD and alleviated the reduction in fruit volume increase through an increase in xylem influx and a decrease in fruit transpiration. Under low fruit load, the competition for assimilates being likely reduced, the phloem flux to fruits increased, similarly to the xylem and transpiration fluxes, without any changes in the fruit water potential. However, different diurnal dynamics among treatments assume variable contributions of turgor and osmotic pressure in F3 and F6 fruits, and hypothetical short-term variations in the water potential gradient between stem and fruit, preventing xylem efflux in F3 fruits.     

Diurnal cycles of embolism formation and repair in petioles of grapevine (Vitis vinifera cv. Chasselas)

The impact of water deficit on stomatal conductance (g(s)), petiole hydraulic conductance (K(petiole)), and vulnerability to cavitation (PLC, percentage loss of hydraulic conductivity) in leaf petioles has been observed on field-grown vines (Vitis vinifera L. cv. Chasselas). Petioles were highly vulnerable to cavitation, with a 50% loss of hydraulic conductivity at a stem xylem water potential (Ψ(x)) of -0.95?MPa, and up to 90% loss of conductivity at a Ψ(x) of -1.5?MPa. K(petiole) described a daily cycle, decreasing during the day as water stress and evapotranspiration increased, then rising again in the early evening up to the previous morning's K(petiole) levels. In water-stressed vines, PLC increased sharply during the daytime and reached maximum values (70-90%) in the middle of the afternoon. Embolism repair occurred in petioles from the end of the day through the night. Indeed, PLC decreased in darkness in water-stressed vines. PLC variation in irrigated plants showed the same tendency, but with a smaller amplitude. The Chasselas cultivar appears to develop hydraulic segmentation, in which petiole cavitation plays an important role as a 'hydraulic fuse', thereby limiting leaf transpiration and the propagation of embolism and preserving the integrity of other organs (shoots and roots) during water stress. In the present study, progressive stomatal closure responded to a decrease in K(petiole) and an increase in cavitation events. Almost total closure of stomata (90%) was measured when PLC in petioles reached >90%.     

Field evaluation of water transport in grape berries during water deficits

The net flow in vascular and transpirational components of the grape berry water budget was evaluated during water deficits imposed at different stages of fruit development. Diurnal fluctuations in berry diameter were measured on field-grown grapevines ( Vitis vinifera L. cv. Cabernet Sauvignon) by using electronic displacement transducers. Water deficits were imposed by withholding irrigation, and water potentials of mid-shoot leaves, basal stem xylem and clusters were determined with a pressure chamber. The relative net flows through pedicel xylem and phloem and through berry transpiration were estimated pre-veraison and post-veraison. The xylem functioned nearly exclusively in providing net inflow pre-veraison, while the phloem was clearly dominant post-veraison. Accordingly, the amplitude of diurnal contraction was markedly smaller post-veraison than pre-veraison. The amplitude of diurnal contraction increased dramatically with decreasing plant water status pre-veraison, yet exhibited little sensitivity to low vine water status post-veraison. Measurements of the difference in water potential between clusters and source stems did not provide evidence of a gradient that would elicit significant water movement from the cluster to the stem at any time of the day. This was true for both irrigated and non-irrigated vines, although the non-irrigated vines exhibited a smaller gradient favoring inflow throughout much of the day. The gradient for xylem water transport to the cluster was considerably smaller post-veraison than pre-veraison. The results showed that berry transpiration functioned as the primary pathway for water loss both pre- and post-veraison.     

Diurnal water balance of the cowpea fruit

The vascular network of the cowpea (Vigna unguiculata [L.] Walp.) fruit exhibits the anatomical potential for reversible xylem flow between seeds, pod, and parent plant. Feeding of cut shoots with the apoplast marker acid fuchsin showed that fruits imported regularly via xylem at night, less frequently in early morning, and only rarely in the afternoon. The dye never entered seeds or inner dorsal pod strands connecting directly to seeds. Root feeding (early morning) of intact plants with ³²PO₄ or ³H₂O rapidly (20 min) labeled pod walls but not seeds, consistent with uptake through xylem. Weak subsequent (4 hours) labeling of seeds suggested slow secondary exchange of label with the phloem stream to the fruit. Vein flap feeding of subtending leaves with [¹⁴C]sucrose, ³H₂O, and ³²PO₄ labeled pod and seed intensely, indicating mass flow in phloem to the fruit. Over 90% of the ¹⁴C and ³H of fruit cryopuncture phloem sap was as sucrose and water, respectively. Specific ³H activities of transpired water collected from fruits and peduncles were assayed over 4 days after feeding ³H₂O to roots, via leaf flaps, or directly to fruits. The data indicated that fruits transpired relatively less xylem-derived (apoplastic) water than did peduncles, that fruit and peduncle relied more heavily on phloem-derived (symplastic) water for transpiration in the day than at night, and that water diffusing back from the fruit was utilized in peduncle transpiration, especially during the day. The data collectively support the hypothesis of a diurnally reversing xylem flow between developing fruit and plant.     

Xylem ABA controls the stomatal conductance of field-grown maize subjected to soil compaction or soil drying

Stomatal conductance of individual leaves was measured in a maize field, together with leaf water potential, leaf turgor, xylem ABA concentration and leaf ABA concentration in the same leaves. Stomatal conductance showed a tight relationship with xylem ABA, but not with the current leaf water status or with the concentration of ABA in the bulk leaf. The relationship between stomatal conductance and xylem [ABA] was common for variations in xylem [ABA] linked to the decline with time of the soil water reserve, to simultaneous differences between plants grown on compacted, non-compacted and irrigated soil, and to plant-to-plant variability. Therefore, this relationship is unlikely to be fortuitous or due to synchronous variations. These results suggest that increased concentration of ABA in the xylem sap in response to stress can control the gas exchange of plants under field conditions.     

Small fruit problem in Citrus trees

Summary Physiological causes of the small fruit problem which occurs in certain trees of orange [Citrus sinensis (L.) Osbeck cv. Valencia] were investigated in terms of water relations and gas exchange of fruits during early fruit development as well as tree carbohydrate reserves. These data from cv. Valencia trees with and without a small fruit potential were compared with those of the large fruited cv. Navel. Neither fruit water potential nor fruit transpiration nor tree carbohydrate reserves appeared to be a cause of the small fruit. Yield records showed the small fruit to be assocaited with a large number of fruit per tree. However, fruits from cv. Valencia trees with a small fruit potential respired faster than either fruits of the same cultivar and size from trees without the physiological disorder or fruits of the same size of cv. Navel and also exceeded the dark respiration of the respective leaves. Hence, the small fruit problem in cv. Valencia was partly attributed to inefficient fruit photosynthesis, causing excessive respiration of each of a larger number of fruits compared to fruits of a tree of the same cultivar but without the physiological disorder. Fruits of cv. Valencia respired more in their 2 months longer lifetime on the tree relative to those of cv. Navel. It is concluded that orchard management methods will have to be investigated to balance the fruit load of the cv. Valencia tree utilizing the carbon available for fruit growth and to minimise stress during the early fruit development.     

Hydraulic connections of leaves and fruit to the parent plant in Capsicum frutescens (hot pepper) during fruit ripening

Background and Aims

The hydraulic architecture and water relations of fruits and leaves of Capsicum frutescens were measured before and during the fruiting phase in order to estimate the eventual impact of xylem cavitation and embolism on the hydraulic isolation of fruits and leaves before maturation/abscission.

Methods

Measurements were performed at three different growth stages: (1) actively growing plants with some flowers before anthesis (GS1), (2) plants with about 50 % fully expanded leaves and immature fruits (GS2) and (3) plants with mature fruits and senescing basal leaves (GS3). Leaf conductance to water vapour as well as leaf and fruit water potential were measured. Hydraulic measurements were made using both the high-pressure flow meter (HPFM) and the vacuum chamber (VC) technique.

Key Results

The hydraulic architecture of hot pepper plants during the fruiting phase was clearly addressed to favour water supply to growing fruits. Hydraulic measurements revealed that leaves of GS1 plants as well as leaves and fruit peduncles of GS2 plants were free from significant xylem embolism. Substantial increases in leaf petiole and fruit peduncle resistivity were recorded in GS3 plants irrespective of the hydraulic technique used. The higher fraction of resistivity measured using the VC technique compared with the HPFM technique was apparently due to conduit embolism.

Conclusions

The present study is the first to look at the hydraulics of leaves and fruits during growth and maturation through direct, simultaneous measurements of water status and xylem efficiency of both plant regions at different hours of the day.     

Diurnal water relations of expanding and full-sized cotton fruits and subtending leaves

Water relations of cotton (Gossypium hirsutum L.) fruits have received less attention than those of leaves, although crop water status has an important influence on fruit physiology. This study was conducted to describe diurnal changes in the water relations of cotton fruits and subtending leaves. Young, expanding fruits and full-sized fruits were compared because of previously reported changes in xylem maturity with ontogeny. Diurnal changes in relative water content were greater in leaves than in the capsule walls of fruits. The capsule walls of young fruits had a higher relative water content than subtending leaves, and water content was lower in full-sized (87%) than in expanding (92%) fruits. Water potentials of subtending leaves were always approximately 0-3 MPa lower than those of capsule walls. Water potential gradients favoured passive water flow from young fruits to branches, but water potentials of branches and the capsule walls of full-sized fruits were similar (?0.7 MPa). Water potential gradients were consistent throughout the day. These results indicate that xylem transport to young fruits is unlikely, but may occur in older fruits.     

Reproductive water supply is prioritized during drought in tomato

Reproductive success largely defines the fitness of plant species. Understanding how heat and drought affect plant reproduction is thus key to predicting future plant fitness under rising global temperatures. Recent work suggests reproductive tissues are highly vulnerable to water stress in perennial plants where reproductive sacrifice could preserve plant survival. However, most crop species are annuals where such a strategy would theoretically reduce fitness. We examined the reproductive strategy of tomato (Solanum lycopersicum var. Rheinlands Ruhm) to determine whether water supply to fruits is prioritized above vegetative tissues during drought. Using optical methods, we mapped xylem cavitation and tissue shrinkage in vegetative and reproductive organs during dehydration to determine the priority of water flow under acute water stress. Stems and peduncles of tomato showed significantly greater xylem cavitation resistance than leaves. This maintenance of intact water supply enabled tomato fruit to continue to expand during acute water stress, utilizing xylem water made available by tissue collapse and early cavitation of leaves. Here, tomato plants prioritize water supply to reproductive tissues, maintaining fruit development under drought conditions. These results emphasize the critical role of water transport in shaping life history and suggest a broad relevance of hydraulic prioritization in plant ecology.     

Drought increases freezing tolerance of both leaves and xylem of Larrea tridentata

Drought and freezing are both known to limit desert plant distributions, but the interaction of these stressors is poorly understood. Drought may increase freezing tolerance in leaves while decreasing it in the xylem, potentially creating a mismatch between water supply and demand. To test this hypothesis, we subjected Larrea tridentata juveniles grown in a greenhouse under well‐watered or drought conditions to minimum temperatures ranging from ?8 to ?24 °C. We measured survival, leaf retention, gas exchange, cell death, freezing point depression and leaf‐specific xylem hydraulic conductance (k_l). Drought‐exposed plants exhibited smaller decreases in gas exchange after exposure to ?8 °C compared to well‐watered plants. Drought also conferred a significant positive effect on leaf, xylem and whole‐plant function following exposure to ?15 °C; drought‐exposed plants exhibited less cell death, greater leaf retention, higher k_l and higher rates of gas exchange than well‐watered plants. Both drought‐exposed and well‐watered plants experienced 100% mortality following exposure to ?24 °C. By documenting the combined effects of drought and freezing stress, our data provide insight into the mechanisms determining plant survival and performance following freezing and the potential for shifts in L. tridentata abundance and range in the face of changing temperature and precipitation regimes.     

Shade mitigates photoinhibition and enhances water use efficiency in kiwifruit under drought

We tested the hypotheses that a reduction of incident light of 50 % over sun-acclimated leaves of water stressed kiwifruit (Actinidia deliciosa var. deliciosa) would (1) reduce stomatal limitations to carbon supply and (2) mitigate the inactivation of the primary photochemistry associated with photosystem (PS) II, thereby this increases carbon gain and water-use efficiency (WUE). Groups of field-grown vines were either shaded or left naturally exposed and subjected to progressive water stress in order to study moderately and severely droughted vines, while other groups were well irrigated. Daily variations in leaf gas exchange and midday chlorophyll (Chl) a fluorescence were determined once plants had −0.6 MPa (moderate stress) and −1.0 MPa pre-down leaf water potential (severe stress). Variations in Chl pigment content and specific leaf area (SLA) are also discussed. Results reveal that 50 % shade application maintained efficiency of PSII close to 0.8 even under severe drought so that to prevent its large decline (0.65) recorded in sunlit leaves. Under moderate stress level stomata behaviour dominated upon metabolic impairments of PSII. Reduction of irradiance increased WUE (15–20 %) in droughted vines, representing a valuable intervention to save photosynthetic apparatus and improve WUE in vines experiencing typical Mediterranean summer stresses.     

How do salinity and water stress affect transport of water,assimilates and ions to tomato fruits?

The study was conducted in order to determine whether water stress affects the accumulation of dry matter in tomato fruits similarly to salinity, and whether the increase in fruit dry matter content is solely a result of the decrease in water content. Although the rate of water transport to tomato fruits decreased throughout the entire season in saline water irrigated plants, accumulation rates of dry matter increased significantly. Phloem water transport contributed 80–85% of the total water transport in the control and water-stressed plants, and over 90% under salinity. The concentration of organic compounds in the phloem sap was increased by 40% by salinity. The rate of ions transported via the xylem was also significantly increased by salinity, but their contribution to fruit osmotic adjustment was less. The rate of fruit transpiration was also markedly reduced by salinity. Water stress also decreased the rate of water transport to the tomato fruit and increased the rate of dry matter accumulation, but much less than salinity. The similar changes, 10–15%, indicate that the rise in dry matter accumulation was a result of the decrease in water transport. Other parameters such as fruit transpiration rates, phloem and xylem sap concentration, relative transport via phloem and xylem, solutes contributing to osmotic adjustment of fruits and leaves, were only slightly affected by water stress. The smaller response of these parameters to water stress as compared to salinity could not be attributed to milder stress intensity, as leaf water potential was found to be more negative. Measuring fruit growth of girdled trusses, in which phloem flow was inactive, and comparing it with ungirdled trusses validated the mechanistic model. The relative transport of girdled as compared to ungirdled fruits resembled the calculated values of xylem transport.     

Day-night changes in leaf water relations associated with the rhythm of crassulacean acid metabolism in Kalanchoë daigremontiana

A study was made of the day-night changes under controlled environmental conditions in the bulk-leaf water relations of Kalanchoë daigremontiana, a plant showing Crassulacean acid metabolism. In addition to nocturnal stomatal opening and net CO₂ uptake, the leaves of well-watered plants showed high rates of gas exchange during the whole of the second part of the light period. Measurements with the pressure chamber showed that xylem tension increased during the night and then decreased towards a minimum at about midday; a significant increase in xylem tension was also seen in the late afternoon. Cell-sap osmotic pressure paralleled leaf malate content and was maximum at dawn and minimum at dusk. The relationship between these two variables indicated that the nocturnally synthesized malate was apparently behaving as an ideal osmoticum. To estimate bulk-leaf turgor pressure, values for water potential were derived by correcting the pressurechamber readings for the osmotic pressure of the xylem sap. This itself was found to depend on the malate content of the leaves. Bulk-leaf turgor pressure changed rhythmically during the day-night cycle; turgor was low during the late afternoon and for most of the night, but increased quickly to a maximum of 0.20 MPa around midday. In water-stressed plants, where net CO₂ uptake was restricted to the dark period, there was also an increase in bulk-leaf turgor pressure at the start of the light period, but of reduced magnitude. Such changes in turgor pressure are likely to be of considerable ecological importance for the water economy of crassulacean-acid-metabolism plants growing in their natural habitats.Abbreviation and symbols CAM Crassulacean acid metabolism - P turgor pressure - osmotic pressure - water potential Dedicated to Professor Dr. H. Ziegler on the occasion of his 60th birthday     

Increased photosynthesis and water potentials in Silphium integrifolium galled by cynipid wasps

Interactions between drought, insect herbivory, photosynthesis, and water potential play a key role in determining how plants tolerate and defend against herbivory, yet the effects of insect herbivores on photosynthesis and water potential are seldom assessed. We present evidence that cynipid wasp galls formed by Antistrophus silphii on Silphium integrifolium increase photosynthesis (A), stomatal conductance (g), and xylem water potential (). Preliminary data showed that in drought-stressed plants galled shoots had 36% greater A, and 10% greater stem than ungalled shoots, while in well-watered plants leaf gas exchange was not affected by galls. We hypothesized that 1) galled shoots have higher , g, and A than ungalled shoots, but this differences diminishes if plant drought stress is reduced, and 2) galls can reduce decreases in A and g if water availability decreases. A field experiment testing the first hypothesis found that galls increased g and , but that differences between galled and ungalled shoots did not diminish after plants were heavily watered. A laboratory test of the second hypothesis using potted Silphium found that galled plants had smaller drops in A and g over a 4-day dry-down period. A vs g and A vs intercellular CO₂ concentration relationships were consistent with the explanation that increased allows galls to increase A by reducing stomatal limitation of A, rather than by altering sink-source relationships or by removing low- limitations on non-stomatal components of A. Our working hypothesis is that galls increase and A by reducing the shoot: root ratio so that the plant is exploiting a greater soil volume per unit leaf area. We argue that increased A is an ineffective way for Silphium to compensate for negative effects of gall insect attack. Instead, increased and A may protect gall insects from variation in resource availability caused by periodic drought stress, potentially reducing negative effects of drought on plant quality and on gall insect populations.     

Plant water relations and control of cell elongation at low water potentials

Recent developments in water status measurement techniques using the psychrometer, the pressure probe, the osmometer and pressure chamber are reviewed, and the process of cell elongation from the viewpoint of plant-water relations is discussed for plants subjected to various environmental stress conditions. Under water-deficient conditions, cell elongation of higher plants can be inhibited by interruption of water flow from the xylem to the surrounding elongating cells. The process of growth inhibition at low water potentials could be reversed by increasing the xylem water potential by means of pressure application in the root region, allowing water to flow from the xylem to the surrounding cells. This finding confirmed that a water potential field associated with growth process,i.e., the growth-induced water potential, is an important regulating factor for cell elongation other than metabolic factors. The concept of the growth-induced water potential was found to be applicable for growth retardation caused by cold stress, heat stress, nutrient deficiency and salinity stress conditions. In the present review, the fact that the cell elongation rate is primarily associated with how much water can be absorbed by elongating cells under water-deficiency, nutrient deficiency, salt stress, cold stress and heat stress conditions is suggested.     

High light decreases xylem contribution to fruit growth in tomato

Recently, contradicting evidence has been reported on the contribution of xylem and phloem influx into tomato fruits, urging the need for a better understanding of the mechanisms involved in fruit growth. So far, little research has been performed on quantifying the effect of light intensity on the different contributors to the fruit water balance. However, as light intensity affects both transpiration and photosynthesis, it might be expected to induce important changes in the fruit water balance. In this study, tomato plants (Solanum lycopersicum L.) were grown in light and shade conditions and the fruit water balance was studied by measuring fruit growth of girdled and intact fruits with linear variable displacement transducers combined with a model‐based approach. Results indicated that the relative xylem contribution significantly increased when shading lowered light intensity. This resulted from both a higher xylem influx and a lower phloem influx during the daytime. Plants from the shade treatment were able to maintain a stronger gradient in total water potential between stem and fruits during daytime, thereby promoting xylem influx. It appeared that the xylem pathway was still functional at 35 days after anthesis and that relative xylem contribution was strongly affected by environmental conditions.     

Diurnal pattern of water potential in woody plants

The dynamic relationship between the rates of water loss and uptake controls plant water status. Marked diurnal variations in water potential of both leaves and fruit occurred in all plants studied. Variations in water status during the day were most clearly related to changes in evaporative demand of the air and were different for the east and west sides of a tree. At night, the plant water potential reflected the soil moisture status.

Changes in the water potential of pear fruit were correlated with changes in fruit diameter. Since water loss from fruit occurred mostly through the pedicel into the xylem of the tree, the fruit could be used as a crude gauge of xylem water potential, which also showed dramatic changes during the day.

     

Effects of Water Stress on Fruit Quality Attributes of Kiwifruit

Four-year-old kiwifruit vines (Actinidia deliciosa(A. Chev.)C. F. Liang et A. R. Ferguson var.deliciosacv. Hayward) werestudied to determine response of the plant and effects on fruitquality when irrigation water was withheld either early or latein the growing season. The greatest effect on fruit growth occurredwhen water was withheld early in the season. Harvest weightof fruit from early-stressed vines was approx. 25% less thanthe weight of fruit on control vines. Early season water stressresulted in a transient increase in concentrations of solublecarbohydrates in both leaves and fruit. This was accompaniedby a reduction in stomatal conductance of the leaves. Starchlevels in leaves but not fruit were reduced by both stress treatments.Concentrations of sucrose at harvest in fruit from vines stressedlate in the season were markedly higher than in other fruit,and softness of the fruit was unaffected. These differenceswere maintained through the 12 weeks in cool storage after harvest.Withholding irrigation water to kiwifruit vines late in theseason may prove a useful management tool to manipulate somequality attributes of the fruit.Copyright 1998 Annals of BotanyCompany Kiwifruit;Actinidia deliciosa; water stress; fruit quality; soluble solids.     

An abscisic acid-related reduced transpiration promotes gradual embolism repair when grapevines are rehydrated after drought

* Proposed mechanisms of embolism recovery are controversial for plants that are transpiring while undergoing cycles of dehydration and rehydration. * Here, water stress was imposed on grapevines (Vitis vinifera), and the course of embolism recovery, leaf water potential (Psi(leaf)), transpiration (E) and abscisic acid (ABA) concentration followed during the rehydration process. * As expected, Psi(leaf) and E decreased upon water stress, whereas xylem embolism and leaf ABA concentration increased. Upon rehydration, Psi(leaf) recovered in 5 h, whereas E fully recovered only after an additional 48 h. The ABA content of recovering leaves was higher than in droughted controls, both on the day of rewatering and the day after, suggesting that ABA accumulated in roots during drought was delivered to the rehydrated leaves. In recovering plants, xylem embolism in petioles, shoots, and roots decreased during the 24 h following rehydration. * A model is proposed to describe plant recovery after rehydration based on three main points: embolism repair occurs progressively in shoots and further in roots and in petioles, following an almost full recovery of Psi(leaf); hydraulic conductance recovers during diurnal transpiring hours, when formation and repair of embolisms occurs in all plant organs; an ABA residual signal in rehydrated leaves hinders stomatal opening even when water relations have recovered, suggesting that an ABA-induced transpiration control promotes gradual embolism repair in rehydrated grapevines.     

Mechanism of export of organic material from the developing fruits of pea

Pisum sativum L. fruits export a small quantity of radiolabeled substances to other plant parts after the fruits are allowed to photosynthesize in the presence of ¹⁴CO₂. Export was uninhibited by peduncle girdling suggesting an apoplastic route for transport of material, presumably by `reverse' flow in the peduncle xylem. To determine if any diurnal water potential gradient formed between pea leaves and fruit might be responsible for the observed export, the water potentials of the various organs were monitored over 24 hours. Water potential differences of up to 7.5 bars existed between leaves and fruit in long photoperiods, and up to 2.5 bars in short photoperiods. Pulses of <sup>14</sup>CO<sub>2</sub> labeling indicated that initial delivery of exported label was to `transpirational sinks,' with subsequent redistribution of label to metabolic sinks. Export to the apical bud appeared to be direct via the xylem. Application of membrane-impermeable inulin to a surgically opened seed coat `cup' resulted in export mainly to the subtending leaf with little redistribution. Simultaneous application of sucrose to the seed coat resulted in more extensive distribution of the sucrose, consistent with reloading of the sucrose into mature leaf phloem. Thus, export of material from fruits appears to occur via a xylem pathway in response to transpirationally derived water potential gradients.