Regulated deficit irrigation and the recovery of water relations in pistachio trees

Recovery of water status in water-stressed pistachio trees (Pistacia vera L. cv. Kerman) was investigated by subjecting trees to regulated deficit irrigation (RDI) (60% of crop evapotranspiration rate, ET(c)) during stages I and II of fruit development (FD) followed by full irrigation during FD stage III (kernel-filling). Trees irrigated at 100% ET(c) throughout FD stages I, II and III served as controls. Water-stress severity was characterized by changes in soil water content and midday stem water potential (Psi(md)). Midday leaf conductance (g(1)) and trunk diameter variation (TDV) were also measured. In RDI trees, the lowest Psi(md) value, -1.8 MPa, occurred at the end of the RDI period. The corresponding value for the control trees was around -1.1 MPa. Although the RDI treatment affected gas exchange later than Psi(md), the greatest reductions in gas exchange (60% of control values) also appeared at the end of the RDI period. There were significant differences in TDV between control and RDI trees at the end of the RDI period. Although plant water status recovered within 20 days of resuming irrigation, the TDV values indicated a longer period might be necessary for complete recovery. Recovery of g(1) was faster than that of Psi(md), although differences in TDV between control and RDI trees indicated that gas exchange recovered later than Psi(md). The slow recovery of pistachio trees during FD stage III from water stress imposed during FD stages I and II suggests that irrigation should exceed 100% ET(c) during FD stage III or that more extensive irrigation should commence before the end of FD stage II.     

Effects of irrigation deprivation during the harvest period on leaf persistence and function in mature almond trees

In nut tree orchards in California, irrigation is typically withheld during the harvest period to reduce the likelihood of bark damage during mechanical shaking of the trees. The ensuing water stress, however, may result in premature defoliation and subsequent yield declines. Our objective was to establish and quantify the water stress resulting from irrigation deprivation and determine its impact on leaf function and persistence in mature almond trees (Prunus dulcis (Mill.) D.A. Webb cv. Nonpareil) during a 3-year field experiment. The severity of the water stress was characterized by measurements of predawn leaf (Psi(pd)) and midday stem (Psi(ms)) water potentials, stomatal conductance (gs), net CO2 assimilation rate (A) and leaf abscission. During 1995, Psi(ms) of fully irrigated (FI) trees was maintained above -1.0 MPa. In trees in the moderate- (MS) and severe-stress (SS) treatments, Psi(ms) was reduced to -1.4 to -2.0 MPa and -2.0 to -2.6 MPa, respectively. After 18 days of irrigation deprivation, A was reduced by 32 and 58% at midday and early afternoon, respectively, compared with morning values. A significant decrease in morning values of A only occurred after 30 days of irrigation deprivation. Water-use efficiency and A declined as evaporative demand increased from morning to afternoon. Assimilation also declined seasonally as leaves aged. Midday stem water potential was highly correlated with A, but less so with gs. The coefficient of determination between Psi(ms) and gs improved considerably when vapor pressure deficit and wind were multiply regressed with Psi(ms). Although A recovered rapidly when MS trees were irrigated, recovery in SS trees was slower and incomplete. Integrating the MS and SS effects for an extended period during 1995 resulted in 14 and 30% declines in A, and 6 and 20% declines in gs, respectively. The apparent Psi(ms) threshold for leaf abscission was -1.8 MPa. Daily canopy light interception declined with decreasing Psi(ms) as a result of premature defoliation (and perhaps altered leaf angles) from 67.9% in FI trees to 61.4 and 60.7% in MS and SS trees, respectively.     

Seasonal trends in photosynthetic parameters and stomatal conductance of blue oak (Quercus douglasii) under prolonged summer drought and high temperature

Understanding seasonal changes in photosynthetic parameters and stomatal conductance is crucial for modeling long-term carbon uptake and energy fluxes of ecosystems. Gas exchange measurements of CO2 and light response curves on blue oak leaves (Quercus douglasii H. & A.) were conducted weekly throughout the growing season to study the seasonality of photosynthetic capacity (Vcmax) and Ball-Berry slope (m) under prolonged summer drought and high temperature. A leaf photosynthetic model was used to determine Vcmax. There was a pronounced seasonal pattern in Vcmax. The maximum value of Vcmax, 127 micromol m(-2) s(-1), was reached shortly after leaf expansion in early summer, when air temperature was moderate and soil water availability was high. Thereafter, Vcmax declined as the soil water profile became depleted and the trees experienced extreme air temperatures, exceeding 40 degrees C. The decline in Vcmax was gradual in midsummer, however, despite extremely low predawn leaf water potentials (Psipd, approximately -4.0 MPa). Overall, temporal changes in Vcmax were well correlated with changes in leaf nitrogen content. During spring leaf development, high rates of leaf dark respiration (Rd, 5-6 micromol m(-2) s(-1)) were observed. Once a leaf reached maturity, Rd remained low, around 0.5 micromol m(-2) s(-1). In contrast to the strong seasonality of Vcmax, m and marginal water cost per unit carbon gain (partial partial differential E/ partial partial differential A) were relatively constant over the season, even when leaf Psipd dropped to -6.8 MPa. The constancy of partial partial differential E/ partial partial differential A suggests that stomata behaved optimally under severe water-stress conditions. We discuss the implications of our findings in the context of modeling carbon and water vapor exchange between ecosystems and the atmosphere.     

Contrasting physiological responses of two co-occurring eucalypts to seasonal drought at restored bauxite mine sites

This study describes the physiological response of two co-occurring tree species (Eucalyptus marginata and Corymbia calophylla) to seasonal drought at low- and high-quality restored bauxite mine sites in south-western Australia. Seasonal changes in photosynthesis (A), stomatal conductance (g(s)), leaf water potential (ψ), leaf osmotic potential (ψ), leaf relative water content (RWC) and pressure-volume analysis were captured over an 18-month field study to (i) determine the nature and severity of physiological stress in relation to site quality and (ii) identify any physiological differences between the two species. Root system restriction at the low-quality site reduced maximum rates of gas exchange (g(s) and A) and increased water stress (midday ψ and daily RWC) in both species during drought. Both species showed high stomatal sensitivity during drought; however, E. marginata demonstrated a higher dehydration tolerance where ψ and RWC fell to -3.2 MPa and 73% compared with -2.4 MPa and 80% for C. calophylla. Corymbia calophylla showed lower g(s) and higher ψ and RWC during drought, indicating higher drought tolerance. Pressure-volume curves showed that cell-wall elasticity of E. marginata leaves increased in response to drought, while C. calophylla leaves showed lower osmotic potential at zero turgor in summer than in winter, indicating osmotic adjustment. Both species are clearly able to tolerate seasonal drought at hostile sites; however, by C. calophylla closing stomata earlier in the drought cycle, maintaining a higher water status during drought and having the additional mechanism of osmotic adjustment, it may have a greater capacity to survive extended periods of drought.     

Mechanisms of drought response in Thuja occidentalis L. II. Post-conditioning water stress and stress relief

We examined the extent of osmotic adjustment and the changes in relative water content (RWC) and transpiration rate (i.e., relative stomatal function) that occur in water-deficit-conditioned 6-year-old Thuja occidentalis L. (eastern white cedar) trees in response to a severe drought. Trees conditioned by successive cycles of mild or moderate nonlethal water stress (conditioning) and nonconditioned trees were exposed to drought (i.e., -2.0 MPa predawn water potential) to determine if water deficit conditioning enhanced tolerance to further drought stress. Following drought, all trees were well watered for 11 days to evaluate how quickly osmotic potential, RWC and transpiration rate returned to preconditioning values. Both nonconditioned trees and mildly conditioned trees exhibited similar responses to drought, whereas moderately conditioned trees maintained higher water potentials and transpiration rates were 38% lower. Both conditioned and nonconditioned trees exhibited a similar degree of osmotic adjustment (-0.39 MPa) in response to drought relative to the well-watered control trees. The well-watered control trees, nonconditioned trees and mildly conditioned trees had similar leaf RWCs that were about 3% lower than those of the moderately conditioned trees. Following the 11-day stress relief, there were no significant differences in osmotic potential between the well-watered control trees and any of the drought-treated trees. Daily transpiration rates and water potential integrals (WPI) of all drought-treated trees approached those of the well-watered control trees during the stress relief period. However, the relationship between cumulative transpiration and WPI showed that previous exposure to drought stress reduced transpiration rates. Leaf RWC of the moderately conditioned trees remained slightly higher than that of the nonconditioned and mildly conditioned trees.     

Deficit irrigation and rootstock: their effects on water relations, vegetative development, yield, fruit quality and mineral nutrition of Clemenules mandarin

Differences between rootstocks, 'Cleopatra' mandarin and 'Carrizo' citrange, in soil-plant water relations and the influence of these factors on vigor, crop yield, fruit quality and mineral nutrition were evaluated in field-grown Clemenules mandarin trees irrigated at 100% of potential seasonal evaporation (ET(c)) (control treatment), or irrigated at 100% ET(c), except during Phases I and III of fruit growth and post-harvest when no irrigation was applied (deficit irrigation (DI) treatment), for 3 years. Differences between rootstocks in plant-soil water relations were the primary cause of differences among trees in vegetative development and fruit yield. After 3 years of DI treatment, trees on 'Cleopatra' showed more efficient soil water extraction than trees on 'Carrizo', and maintained a higher plant water status, a higher gas exchange rate during periods of water stress and achieved faster recovery in gas exchange following irrigation after water stress. The DI treatment reduced vegetative development more in trees on 'Carrizo' than in trees on 'Cleopatra'. Cumulative fruit yield decreased more in DI trees on 'Carrizo' (40%) than on 'Cleopatra' (27%). The yield component most affected by DI in 'Cleopatra' was the number of fruit, whereas in 'Carrizo' it depended on the severity of water stress reached in each phase (severe water stress in Phase I affected mainly the number of fruit, whereas it affected fruit size the most in Phase III). In the third year of DI treatment, water-use efficiency decreased sharply in trees on 'Carrizo' (70%) compared to trees on 'Cleopatra' (30%). Thus, trees on 'Cleopatra' were able to tolerate moderate water stress, whereas trees on 'Carrizo' were more sensitive to changes in soil water content.     

Effects of irrigation deprivation during the harvest period on yield determinants in mature almond trees

Effects of irrigation deprivation during the harvest period on yield determinants in mature almond (Prunus dulcis (Mill.) D.A. Webb cv. Nonpareil) trees were investigated during a 3-year field experiment. Return bloom and fruit set were measured on 2185 individually tagged spurs. Water stress resulting from irrigation deprivation during the harvest period, which purportedly coincides with the time of flower initiation, had no effect on the percentage of spurs that flowered or set fruit during subsequent years. Although water stress had no apparent effect on spur mortality, 66% of the tagged spurs died within 3 years. In addition, many spurs were vegetative by the third year, indicating the importance of spur renewal for sustained fruit production. Reductions in nut yield were evident after two successive years of irrigation deprivation during the harvest period. Regression analysis indicated a loss in yield of 7.7 kg tree(-1) in response to each 1 MPa decrease in stem water potential below -1.2 MPa during the previous seasons. The number of fruiting positions per tree (estimated indirectly for whole trees based on weight of current-year shoots > 5 cm in length) was negatively associated with water stress. Yield reduction in response to water stress during harvest appears to be a compound, multiyear effect, associated with reduced annual growth and renewal of fruiting positions.     

Usefulness of diurnal trunk shrinkage as a water stress indicator in plum trees

We compared seasonal changes in maximum diurnal trunk shrinkage (MDS) with seasonal changes in midday stem water potential (Psi(s)) over three years in plum trees grown in differing drip-irrigated regimes. In well-irrigated trees, day-to-day variations in Psi(s) and MDS were related to evaporative demand. Reference equations were obtained to predict MDS and Psi(s) values for well-irrigated trees as functions of environmental conditions. A decrease in plant water status toward the end of the growing season occurred even in the well-irrigated trees, probably reflecting a reduced volume of soil wetted by the drip irrigation system. Thus, for the prediction of Psi(s), different reference equations are required for the fruit-growth and after-harvest phenological periods. A seasonal change in the relationship between MDS and Psi(s) was observed, which compensated for the decrease in plant water status such that well-irrigated trees had similar MDS values during both the fruit-growth and after-harvest periods. The influence of tree size on the relationship between MDS and Psi(s) was also investigated. For tree trunk diameters ranging between 8 and 13 cm, MDS increased 13% for each cm of increase in trunk diameter, as a result of the thicker phloem tissues of the larger trees. This finding may allow extrapolation of Psi(s) predictions based on empirical relationships with MDS to plum trees of different sizes.     

Stomatal responses to drought at a Mediterranean site: a comparative study of co-occurring woody species differing in leaf longevity

We studied stomatal responses to decreasing predawn water potential (Psipd) and increasing leaf-to-air water vapor pressure difference (VPD) of co-occurring woody Mediterranean species with contrasting leaf habits and growth form. The species included two evergreen oaks (Quercus ilex subsp. ballota (Desf.) Samp. and Q. suber L.), two deciduous oaks (Q. faginea Lam. and Q. pyrenaica Willd.) and two deciduous shrubs (Pyrus bourgaeana Decne. and Crataegus monogyna Jacq.). Our main objective was to determine if stomatal sensitivity is related to differences in leaf life span and leaf habit. The deciduous shrubs had the least conservative water-use characteristics, with relatively high stomatal conductance and low stomatal sensitivity to soil and atmospheric drought. As a result, Psipd decreased greatly in both species during the growing season, resulting in early leaf abscission in the summer. The deciduous oaks showed intermediate water-use characteristics, having maximum stomatal conductances and CO2 assimilation rates similar to or even higher than those of the deciduous shrubs. However, they had greater stomatal sensitivity to soil drying and showed less negative Psipd values than the deciduous shrubs. The evergreen oaks, and especially the species with the greatest leaf longevity, Q. ilex, exhibited the most conservative water-use behavior, having lower maximum stomatal conductances and greater sensitivity to VPD than the deciduous species. As a result, Psipd decreased less during the growing season in the evergreens than in the deciduous species, which may contribute to greater leaf longevity by avoiding irreversible damage during the summer drought. However, the combination of low maximum CO2 assimilation rates and high stomatal sensitivity to drought must have a negative impact on the final carbon budget of leaves with a long life span.     

Effects of soil water deficit on gas exchange characteristics and water relations of orchard lychee (Litchi chinensis Sonn.) trees

Eight-year-old lychee (Litchi chinensis Sonn.) trees, cv. 'Bengal,' growing in krasnozem soil were subjected to soil water deficit from one month before flowering until harvest by covering the ground with polyethylene sheeting and withholding irrigation. The ratio of daytime stomatal conductance of unirrigated to irrigated trees decreased 20% during the three months of increasing water deficit. Predawn leaf water potentials of irrigated trees averaged about -0.3 MPa throughout the period, whereas they declined progressively to -0.9 MPa in unirrigated trees. Minimum daytime leaf water potential in the unirrigated trees decreased from -1.0 to -1.1 MPa at the beginning of the drought period to -2.2 to -2.4 MPa after three months, and calculated whole-plant conductance did not change with decreasing availability of water. The calculated soil-root water potential declined to less than -1.0 MPa in unirrigated trees. Capacitance effects on the relationship between leaf water potential and transpiration were significant only at low transpiration rates. Although unirrigated trees reduced soil water content at 0-30 cm depths to an equivalent water potential of -1.0 MPa, fruit shedding was significantly less than in irrigated trees. Water deficit had no effect on the fresh weight of pericarp, but caused increased seed size and decreased fresh weight of flesh, resulting in fruit from unirrigated trees being 16% lower in total fresh weight per fruit than fruit from irrigated trees.     

Responses of gas exchange to reversible changes in whole-plant transpiration rate in two conifer species

This study examined the autonomy of branches with respect to the control of transpiration (E) in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata Donn) seedlings. Experiments were conducted on whole seedlings in a gas exchange system with a dual-cuvette that permitted independent manipulation and measurement of E in the upper and lower cuvettes. The value of E in one cuvette was manipulated by varying vapor pressure deficit (D) between 2.2 and 0.2 kPa, whereas D in the other cuvette was held at 2.2 kPa. Reducing D, while increasing stomatal conductance (gs), resulted in an overall decrease in E. In western red cedar, this decrease was almost threefold, and in Douglas-fir, approximately fourfold. In well-watered western red cedar, a reduction of whole-plant E by 46% (brought about by reducing D in the upper cuvette) resulted in a 12% increase in gs, a 12% increase in E and a 7% increase in net assimilation (A) of untreated foliage in the lower cuvette. Responses of gs, E and A of untreated foliage were similar irrespective of whether foliage was at the top or bottom of the seedling. When D in the treatment cuvette was restored to 2.2 kPa, gs, E and A of foliage in the untreated cuvette returned to pretreatment values. In contrast, in well-watered Douglas-fir, there was almost no change in gs, E or A of untreated foliage in one cuvette when D in the other cuvette was reduced, causing a 52% reduction in whole-plant E. However, similar manipulations on drought-stressed Douglas-fir led to 7-19% increases in gs, E and A of untreated foliage. In well-watered western red cedar, daytime leaf water potential (Psil) was maintained near -0.9 MPa over a wide range of D, whereas Psil of Douglas-fir decreased from -1.2 to -1.5 MPa as D increased. The tighter (isohydric) regulation of Psil in western red cedar may partly explain its greater stomatal response to D and variation in whole-plant E compared with Douglas-fir. In response to a reduction in E, measured increases in Psil and gs of unmanipulated foliage were less than predicted by a model assuming complete hydraulic connectivity of foliage. Our results suggest the foliage of both species is partially autonomous with respect to water.     

Water deficits at anthesis reduce CO(2) assimilation and yield of lychee (Litchi chinensis Sonn.) trees

Ten-year-old 'Tai So' lychee (Litchi chinensis Sonn.) trees growing on a sandy loam soil in subtropical South Africa (latitude 25 degrees S) were watered weekly (well-watered treatment) or droughted from late July until January (drought treatment). After 16 weeks, at which time the trees obtained most of their water from below 150 cm, average soil water content at 0 to 150 cm depth was 14.5 +/- 0.1% in the well-watered treatment and reached a minimum of 7.6% in the drought treatment. At Week 7, minimum leaf water potential (Psi(L)) in the morning and early afternoon declined to -2.6 and -2.8 MPa, respectively, in droughted trees compared with -1.5 and -2.2 MPa, respectively, in well-watered trees. From Week 9, stomatal conductance and net CO(2) assimilation rate ranged from 70 to 300 mmol m(-2) s(-1) and 3 to 13 micro mol CO(2) m(-2) s(-1), respectively, in well-watered trees. The corresponding values for droughted trees were 50 to 180 mmol m(-2) s(-1) and 2 to 6 micro mol CO(2) m(-2) s(-1). Five weeks after rewatering the droughted trees, gas exchange had not recovered to the rate in well-watered trees, although tree water status recovered within a week of rewatering. In the well-watered trees, water use (E(t)) was 26 +/- 1 mm week(-1) with evaporation (E(p)) of 20 to 70 mm week(-1) indicating a crop factor (k(c) = E(t)/E(p)) of 0.4 to 1.2. Before anthesis, tree water status did not affect extension growth of floral panicles or leafy shoots. In contrast, no vegetative shoots were initiated after fruit set in the droughted trees when Psi(L) in the morning declined to -2.5 MPa. Water deficits reduced initial fruit set by 30% and final fruit set by 70% as a result of fruit splitting (41.2 +/- 4.0% versus 10.0 +/- 1.3%). Water deficits did not alter the sigmoidal pattern of fruit growth, but reduced yield from 51.4 +/- 5.5 kg tree(-1) in well-watered trees to 7.4 +/- 3.3 kg tree(-1) in droughted trees.     

Hydraulic adjustments underlying drought resistance of Pinus halepensis

Drought-induced tree mortality has increased over the last decades in forests around the globe. Our objective was to investigate under controlled conditions the hydraulic adjustments underlying the observed ability of Pinus halepensis to survive seasonal drought under semi-arid conditions. One hundred 18-month saplings were exposed in the greenhouse to 10 different drought treatments, simulating combinations of intensities (fraction of water supply relative to control) and durations (period with no water supply) for 30 weeks. Stomata closed at a leaf water potential (Ψ(l)) of -2.8 MPa, suggesting isohydric stomatal regulation. In trees under extreme drought treatments, stomatal closure reduced CO(2) uptake to -1 μmol m(-2) s(-1), indicating the development of carbon starvation. A narrow hydraulic safety margin of 0.3 MPa (from stomatal closure to 50% loss of hydraulic conductivity) was observed, indicating a strategy of maximization of CO2 uptake in trees otherwise adapted to water stress. A differential effect of drought intensity and duration was observed, and was explained by a strong dependence of the water stress effect on the ratio of transpiration to evapotranspiration T/ET and the larger partitioning to transpiration associated with larger irrigation doses. Under intense or prolonged drought, the root system became the main target for biomass accumulation, taking up to 100% of the added biomass, while the stem tissue biomass decreased, associated with up to 60% reduction in xylem volume.     

Ecophysiological responses of a young blue gum (Eucalyptus globulus) plantation to weed control

Early weed control may improve the growth of forest plantations by influencing soil water and nutrient availability. To understand eucalypt growth responses to weed control, we examined the temporal responses of leaf gas-exchange, leaf nitrogen concentration (N) and water status of 7-month-old Eucalyptus globulus L. trees in a paired-plot field trial. In addition, we monitored the growth, leaf N and water status of the competing vegetation in the weed treatment. By the end of the 11-month experiment, complete weed control (WF treatment) of largely woody competitors increased the basal diameter of E. globulus by 14%. As indicated by pre-dawn water potentials of >?-?0.05 MPa, interspecies competition for water resources was minimal at this site. In contrast, competition for N appeared to be the major factor limiting growth. Estimations of total plot leaf N (g m(-2) ground) showed that competing vegetation accounted for up to 70% of the total leaf N at the start of the trial. This value fell to 15% by the end of the trial. Despite increased leaf N(area) in WF trees 5 months after imposition of weed control, the photosynthetic capacity (A(1500)) of E. globulus was unaffected by treatment suggesting that the growth gains from weed control were largely unrelated to changes in leaf-level photosynthesis. Increased nutrient availability brought about by weed control enabled trees to increase investment into leaf-area production. Estimates of whole-tree carbon budget based on direct measurements of dark respiration and A(1500) allowed us to clearly demonstrate the importance of leaf area driving greater productivity following early weed control in a nutrient-limited site.     

Influence of soil texture on hydraulic properties and water relations of a dominant warm-desert phreatophyte

We investigated hydraulic constraints on water uptake by velvet mesquite (Prosopis velutina Woot.) at a site with sandy-loam soil and at a site with loamy-clay soil in southeastern Arizona, USA. We predicted that trees on sandy-loam soil have less negative xylem and soil water potentials during drought and a lower resistance to xylem cavitation, and reach E(crit) (the maximum steady-state transpiration rate without hydraulic failure) at higher soil water potentials than trees on loamy-clay soil. However, minimum predawn leaf xylem water potentials measured during the height of summer drought were significantly lower at the sandy-loam site (-3.5 +/- 0.1 MPa; all errors are 95% confidence limits) than at the loamy-clay site (-2.9 +/- 0.1 MPa). Minimum midday xylem water potentials also were lower at the sandy-loam site (-4.5 +/- 0.1 MPa) than at the loamy-clay site (-4.0 +/- 0.1 MPa). Despite the differences in leaf water potentials, there were no significant differences in either root or stem xylem embolism, mean cavitation pressure or Psi(95) (xylem water potential causing 95% cavitation) between trees at the two sites. A soil-plant hydraulic model parameterized with the field data predicted that E(crit) approaches zero at a substantially higher bulk soil water potential (Psi(s)) on sandy-loam soil than on loamy-clay soil, because of limiting rhizosphere conductance. The model predicted that transpiration at the sandy-loam site is limited by E(crit) and is tightly coupled to Psi(s) over much of the growing season, suggesting that seasonal transpiration fluxes at the sandy-loam site are strongly linked to intra-annual precipitation pulses. Conversely, the model predicted that trees on loamy-clay soil operate below E(crit) throughout the growing season, suggesting that fluxes on fine-textured soils are closely coupled to inter-annual changes in precipitation. Information on the combined importance of xylem and rhizosphere constraints to leaf water supply across soil texture gradients provides insight into processes controlling plant water balance and larger scale hydrologic processes.     

Crop load and water stress effects on daily stem growth in peach (Prunus persica)

We investigated crop load and water stress effects on diurnal stem extension growth of field-grown peach (Prunus persica (L.) Batsch) trees. Neither the presence of fruit nor reduced irrigation significantly altered the timing of diurnal fluctuations in stem growth rate. Stems with subtending fruit had significantly reduced growth compared to stems with no subtending fruit. Crop load had no significant effect on relative stem extension rates and the majority of the reduction in absolute growth was the result of a smaller zone of elongation in fruit-bearing stems than in stems with no subtending fruit. Fruit removal did not increase growth rates within 24 h. When irrigation was reduced, the length of the stem elongation zone and total daily stem growth were significantly decreased relative to well-irrigated controls and the decreases were highly correlated with stem water potential. Compared with well-irrigated controls, relative stem extension rates of water-stressed trees were reduced at several times during the 24-h period, but the degree of reduction was not proportional to the difference in stem water potentials between the treatments.     

Effects of water stress cycles on turgor maintenance processes in pear leaves (Pyrus communis)

Effects of water deficits on leaf turgor maintenance processes were analyzed for pear trees (Pyrus communis L. cv. "Barlett") grown in 120-liter containers. Four irrigation treatments were applied: a well-watered control treatment, a spring water stress cycle (Sp), a summer water stress cycle (Su), and a spring plus summer water stress cycle (Sp + Su). For the Sp treatment, water application was progressively reduced from 100 to 20% of the control dose over a period of 27 days in spring. For the Su treatment, water application was progressively reduced over 23 days in summer, from 100 to 20% of the control dose. The Sp + Su treatment comprised both the spring and summer drought stress cycles. Pressure-volume (P-V) curves were constructed and stomatal conductances were determined for pear leaves from each treatment during the spring and summer stress cycles. Leaf water potential (Psi(pi) (0)) and relative water content (R(0)) at the turgor loss point of control leaves tended to decrease from spring to summer. Changes in leaf osmotic water potential at full turgor (Psi(pi) (100)) and in symplast water fraction (R(s)) did not explain the seasonal decrease in Psi(pi) (0). The water stress treatments had no effect on Psi(pi) (100), but R(s) was reduced by the water stress treatments, particularly during the summer stress cycle of the Su and Sp + Su treatments. The decrease in R(s) was correlated with an increase in the slope of the linear region of the P-V curve. Such a coupled adjustment would lead to increased water uptake capacity of water-stressed trees only under non-turgor conditions. Furthermore, pear leaves did not actively accumulate solutes. We conclude, therefore, that changes in leaf tissue water relations as a result of leaf acclimation to water stress are unlikely to facilitate maintenance of fruit productivity under drought.     

Fruit load and canopy shading affect leaf characteristics and net gas exchange of 'Spring' navel orange trees

Five-year-old 'Spring' navel (Citrus sinensis (L.) Osbeck) orange trees were completely defruited, 50% defruited or left fully laden to study effects of fruit load on concentrations of nitrogen (N) and carbohydrate, net assimilation of CO2 (Ac) and stomatal conductance (gs) of mature leaves on clear winter days just before fruit harvest. Leaves on defruited trees were larger, had higher starch concentrations and greater leaf dry mass per area (LDMa) than leaves on fruited trees. Both Ac and gs were more than 40% lower in sunlit leaves on defruited trees than in sunlit leaves on trees with fruit. Leaves immediately adjacent to fruit were smaller, had lower leaf nitrogen and carbohydrate concentrations, lower LDMa and lower Ac than leaves on non-fruiting branches of the same trees. Removing half the crop increased individual fruit mass, but reduced fruit color development. Half the trees were shaded with 50% shade cloth for 4 months before harvest to determine the effects of lower leaf temperature (Tl) and leaf-to-air vapor pressure difference on leaf responses. On relatively warm days when sunlit Tl > 25 degrees C, shade increased Ac and gs, but had no effect on the ratio of internal to ambient CO2 (Ci/Ca) concentration in leaves, implying that high mesophyll temperatures in sunlit leaves were more important than gs in limiting Ac. Sunlit leaves were more photoinhibited than shaded leaves on cooler days when Tl < 25 degrees C. Shade decreased total soluble sugar concentrations in leaves, but had no effect on leaf starch concentrations. Shading had no effects on canopy volume, yield or fruit size, but shaded fruit developed better external color than sun-exposed fruit. Overall, the presence of a normal fruit crop resulted in lower foliar carbohydrate concentrations and higher Ac compared with defruited trees, except on warm days when Ac was reduced by high leaf temperatures.     

Effects of soil drought on photosynthetic traits and antioxidant enzyme activities in Hippophae rhamnoides seedlings

Water deficit is one of the major limiting factors in vegetation recovery and reconstruction in the semi-arid area of loess hilly regions. Leaf photosynthesis in Hippophae rhamnoides Linn., a common tree grown in this region, decreases under water stress, but the mechanism responsible is not clear. The objective of this study was to investigate the effects of drought stress on photosynthesis and the relationship between photosynthetic variables and soil water contents to help us better understand the photophysiological characteristics of H. rhamnoides under water stress and guide cultivation in the loess hilly region. Here,gas exchange, chlorophyll fluorescence and antioxidant enzyme activity in leaves of 3-year-old saplings of H.rhamnoides grown in pots were tested under eight soil water conditions. When soil water content(RWC) was between 38.9 and 70.5 %, stomatal limitation was responsible for the reduced net photosynthetic rate(PN).When RWC was lower than 38.9 %, nonstomatal limitation was the main factor restricting PN. Moderate water stress improved the water use efficiency(WUE) of the leaf.Water stress significantly influenced fluorescence variables and the antioxidant enzyme system. When RWC was between 38.9 and 70.5 %, nonphotochemical quenching(NPQ) increased and then decreased, indicating that thermal energy dissipation was a significant photoprotection mechanism. Antioxidant enzymes were activated when RWC ranged from 48.3 to 70.5 %; under severe water stress(RWC \ 38.9 %), the antioxidant enzyme system was damaged, the activity of the antioxidant enzymes declined, and membranes were damaged. In the semiarid loess hilly region, RWC between 58.6 and 70.5 % was the economic water threshold value that maintained higher WUE and PN, and the maximum soil water deficit level that could sustain H. rhamnoides was RWC of 38.9 %.     

Relationship of water status to vegetative growth and leaf gas exchange of peach (Prunus persica) trees on different rootstocks

We investigated relationships between tree water status, vegetative growth and leaf gas exchange of peach trees growing on different rootstocks under field conditions. Tree water status was manipulated by partially covering (0, approximately 30 and approximately 60%) the tree canopies on individual days and then evaluating the effects of tree water status on vegetative growth and leaf gas exchange. Early morning stem water potentials were approximately -0.4 MPa for trees in all treatments, but mean midday values ranged from -1.1 to -1.7 MPa depending on rootstock and canopy coverage treatment. Relative shoot extension growth rate, leaf conductance, transpiration rate and net CO2 exchange rate differed significantly among trees in the different rootstocks and canopy coverage treatments. Shoot extension growth rate, leaf conductance, leaf transpiration rate and leaf net CO2 exchange rate were linearly correlated with midday stem water potential. These relationships were independent of the rootstock and canopy coverage treatments, indicating that tree water relations are probably directly involved in the mechanism that imparts vegetative growth control by selected peach rootstocks.