Water stress responses of two Mediterranean tree species influenced by native soil microorganisms and inoculation with a plant growth promoting rhizobacterium

Soil microorganisms, such as plant growth-promoting rhizobacteria (PGPR), play crucial roles in plant growth, but their influence on plant water relations remains poorly explored. We studied the effects of native soil microorganisms and inoculation with the PGPR strain Aur6 of Pseudomonas fluorescens on water stress responses of seedlings of the drought-avoiding Pinus halepensis Mill. and the drought-tolerant Quercus coccifera L. Plant growth, nutrient concentrations and physiology (maximum photochemical efficiency of photosystem II (PSII; F(v)/F(m)), electron transport rate (ETR), stomatal conductance (g(s)) and predawn shoot water potential (Psi(PD))) were measured in well-watered plants, and in plants under moderate or severe water stress. Inoculation with PGPR and native soil microorganisms improved tree growth, and their interactions had either additive or synergistic effects. Both F(v)/F(m) and ETR were significantly affected by PGPR and native soil microorganisms. Marked differences in g(s) and Psi(PD) were found between species, confirming that they differ in mechanisms of response to water stress. A complex tree species x treatment interactive response to drought was observed. In P. halepensis, F(v)/F(m) and ETR were enhanced by PGPR and native soil microorganisms under well-watered conditions, but the effects of PGPR on Psi(PD) and g(s) were negative during a period of water stress. In Q. coccifera, F(v)/F(m) and ETR were unaffected or even reduced by inoculation under well-watered conditions, whereas Psi(PD) and g(s) were increased by PGPR during a period of water stress. Our results indicate that microbial associates of roots can significantly influence the response of tree seedlings to drought, but the magnitude and sign of this effect seems to depend on the water-use strategy of the species.     

Winter photoinhibition in the field involves different processes in four co-occurring Mediterranean tree species

Photoinhibition was examined in four co-occurring Mediterranean evergreen tree species during two consecutive winters. In response to low temperatures and saturating light, Juniperus phoenicea L., Pinus halepensis Mill., Quercus coccifera L. and Q. ilex ssp. ballota (Desf.) Samp. exhibited marked chronic photoinhibition, indicated by low predawn maximal photochemical efficiency of photosystem II (PSII) (Fv/Fm). Low Fv/Fm values were correlated with high concentrations of xanthophyll cycle components (VAZ) and with the maintenance of high concentrations of zeaxanthin overnight (DPSpd). In all species, however, chronic photoinhibition was enhanced as the winter progressed in the absence of changes in DPSpd, suggesting cumulative damage toward the end of winter. Photoinhibition differed among species: P. halepensis always displayed significantly higher Fv/Fm values; and Q. coccifera had the lowest Fv/Fm values, showing a high sensitivity to the combination of high light and low temperatures. Differences among species were not fully explained by differences in the xanthophyll pool or its de-epoxidation state. Chronic photoinhibition overlapped with a dynamic photoinhibition as shown by the low values of photochemical efficiency of the open reaction centers of PSII at midday. Winter photoprotective strategies differed among species and may involve photoprotective mechanisms in addition to those associated with xanthophylls. The observed species-specific differences matched results obtained for the same species in summer; however, comparison of the two seasons suggests that the higher VAZ concentration observed in winter has an additional structural photoprotective role.     

Water relations of seedlings of three Quercus species: variations across and within species grown in contrasting light and water regimes

We compared seedling water relations of three Mediterranean Quercus species (the evergreen shrub Q. coccifera L., the evergreen tree Q. ilex L. subsp. ballota (Desf.) Samp. and the deciduous or marcescent tree Q. faginea L.). We also explored seedling potential for acclimation to contrasting growing conditions. In March, 1-year-old seedlings of the three species were planted in pots and grown outdoors in a factorial combination of two irrigation regimes (daily (HW) and alternate day watering (LW)) and two irradiances (43 and 100% of full sunlight). At the end of July, predawn and midday water potentials (Psi(pd), Psi(md)) were measured, and pressure-volume (P-V) curves were obtained for mature current-year shoots. Species exhibited similar Psi(pd) and Psi(md) values, but differed in leaf morphology and water relations. The evergreens possessed larger leaf mass per area (LMA) and were able to maintain positive turgor pressure at lower water potentials than the deciduous species because of their lower osmotic potential at full turgor. However, the three species had similar relative water contents at the turgor loss point because Q. faginea compensated for its higher osmotic potential with greater cell wall elasticity. Values of Psi(pd) had a mean of -1.12 MPa in LW and -0.63 MPa in HW, and Psi(md) had a mean of -1.13 MPa in full sunlight and -1.64 MPa in shade, where seedlings exhibited lower LMA. However, the P-V curve traits were unaffected by the treatments. Our results suggest that Q. faginea seedlings combine the water-use characteristics of mesic deciduous oak and the drought-tolerance of xeric evergreen oak. The ability of Q. coccifera to colonize drier sites than Q. ilex was not a result of higher drought tolerance, but rather may be associated with other dehydration postponement mechanisms including drought-induced leaf shedding. The lack of treatment effects may reflect a relatively low contrast between treatment regimes, or a low inherent responsiveness of these traits in the study species, or both.     

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.     

Water relations of several hardwood species in response to throughfall manipulation in an upland oak forest during a wet year

We investigated the effects of altered precipitation on leaf osmotic potential at full turgor (Psi(pio)) of several species in an upland oak forest during the 1994 growing season as part of a Throughfall Displacement Experiment at the Walker Branch Watershed near Oak Ridge, Tennessee. The main species sampled included overstory chestnut oak (Quercus prinus L.), white oak (Q. alba L.), red maple (Acer rubrum L.); intermediates sugar maple (A. saccharum L.) and blackgum (Nyssa sylvatica Marsh.); and understory dogwood (Cornus florida L.) and red maple. The precipitation treatments were: ambient precipitation; ambient minus 33% of throughfall (dry); and ambient plus 33% of throughfall (wet). Except in late September, midday leaf water potentials (Psi(l)) were generally high in all species in all treatments, ranging from -0.31 to -1.34 MPa for C. florida, -0.58 to -1.51 MPa for A. rubrum, and -0.78 to -1.86 MPa for Q. prinus. Both treatment and species differences in Psi(pio) were evident, with oak species generally exhibiting lower Psi(pio) than A. saccharum, A. rubrum, C. florida, and N. sylvatica. The Psi(pio) of C. florida saplings declined in the dry treatment, and Q. prinus, Q. alba, and A. saccharum all exhibited a declining trend of Psi(pio) in the dry treatment, although Psi(pio) of Q. prinus leaves increased in late August, corresponding to a recovery in soil water potential. Cornus florida exhibited osmotic adjustment with the largest adjustment coinciding with the period of lowest soil water potential in June. The only other species to exhibit osmotic adjustment was Q. prinus, which also maintained a lower baseline Psi(pio) than the other species. We conclude that a 33% reduction of throughfall is sufficient both to alter the water relations of some species in the upland oak forest and to enable the identification of those species capable of osmotic adjustment to a short-term drought during a wet year.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Water storage capacitance and xylem tension in isolated branches of temperate and tropical trees

Trees of tropical semi-deciduous forests range from "drought-avoiding" stem-succulent species with low-density wood (< 0.5 g cm(-3)), which maintain high stem water potentials (psi(STEM) > -0.7 MPa) throughout the year, to "drought-tolerant" deciduous hardwood species (wood density > 0.75 g cm(-3)), which dehydrate strongly during seasonal drought (psi(STEM) < -6 MPa). In stem-succulent and other drought-avoiding species, xylem vessels are surrounded by extensive parenchyma providing intracellular water storage, whereas in deciduous species stem water storage is mainly extracellular. Thirteen tropical and two temperate tree species, representing different functional types, were studied. The contribution of stem water storage to these species' water use during water stress was determined by time-series analysis of dehydration and rehydration of excised leaf-bearing branches of these trees. During dehydration, stem water potential slowly declined 1-2 MPa in drought-avoiding species, but in deciduous species it rapidly fell 4-5 MPa, suggesting that water storage capacitance was related to xylem anatomy. After immersion of dehydrated, leafless branches in water, the decline in xylem tension and rate of water uptake during rehydration were linearly related, as predicted by application of Ohm's law to water flux. The decline of xylem tension during rehydration was biphasic, with a phase of rapid water uptake into extracellular spaces being followed by a prolonged phase of slow water uptake into living cells. The rate of water uptake during rehydration and the minima of leaf water potential observed in the field during the dry season were highly correlated with water storage capacitance, indicating that wood anatomy is a major determinant of drought adaptation.     

Scion-rootstock interaction affects the physiology and fruit quality of sweet cherry

Water relations, leaf gas exchange, chlorophyll a fluorescence, light canopy transmittance, leaf photosynthetic pigments and metabolites and fruit quality indices of cherry cultivars 'Burlat', 'Summit' and 'Van' growing on five rootstocks with differing size-controlling potentials that decrease in the order: Prunus avium L. > CAB 11E > Maxma 14 > Gisela 5 > Edabriz, were studied during 2002 and 2003. Rootstock genotype affected all physiological parameters. Cherry cultivars grafted on invigorating rootstocks had higher values of midday stem water potential (Psi(MD)), net CO(2) assimilation rate (A), stomatal conductance (g(s)), intercellular CO(2) concentration (C(i)) and maximum photochemical efficiency of photosystem II (PSII) (F(v)/F(m)) than cultivars grafted on dwarfing rootstocks. The Psi(MD) was positively correlated with A, g(s) and C(i). Moreover, A was positively correlated with g(s), and the slopes of the linear regression increased from invigorating to dwarfing rootstocks, indicating a stronger regulation of photosynthesis by stomatal aperture in trees on dwarfing Edabriz and Gisela 5. The effect of rootstock genotype was also statistically significant for leaf photosynthetic pigments, whereas metabolite concentrations and fruit physicochemical characteristics were more dependent on cultivar genotype. Among cultivars, 'Burlat' leaves had the lowest concentrations of photosynthetic pigments, but were richest in total soluble sugars, starch and total phenols. Compared with the other cultivars, 'Summit' had heavier fruits, independent of the rootstock. 'Burlat' cherries were less firm and had lower concentrations of soluble sugars and a lower titratable acidity than 'Van' cherries. Nevertheless, 'Van' cherries had lower lightness, chroma and hue angle, representing redder and darker cherries, compared with 'Summit' fruits. In general, Psi(MD) was positively correlated with fruit mass and A was negatively correlated with lightness and chroma. These results demonstrate that: (1) water relations and photosynthesis of sweet cherry tree are mainly influenced by the rootstock genotype; (2) different physicochemical characteristics observed in cherries of the three cultivars suggest that regulation of fruit quality was mainly dependent on the cultivar genotype, although the different size-controlling rootstocks also had a significant effect.     

Acclimation of leaves to contrasting irradiance in juvenile trees differing in shade tolerance

Leaves developing in different irradiances undergo structural and functional acclimation, although the extent of trait plasticity is species specific. We tested the hypothesis that irradiance-induced plasticity of photosynthetic and anatomical traits is lower in highly shade-tolerant species than in moderately shade-tolerant species. Seedlings of two evergreen conifers, shade-tolerant Abies alba Mill. and moderately shade-tolerant Picea abies Karst., and two deciduous angiosperm species, highly shade-tolerant Fagus sylvatica L. and moderately shade-tolerant Acer pseudoplatanus L., were grown in deep shade (LL, 5% of full irradiance) or in full solar irradiance (HL) during 2003 and 2004. Steady state responses of quantum yield of PSII (Phi(PSII)), apparent electron transport rate (ETR), nonphotochemical quenching (NPQ) and photochemical quenching (qP) were generally modified by the light environment, with slower declines in Phi(PSII) and qP and greater maximal ETR and NPQ values in HL plants in at least one season; however, no link between quantitative measures of plasticity of these traits and shade tolerance was found. Plasticity of nine anatomical traits (including palisade cell length, which was reduced in LL) showed no relationship with shade tolerance, but was less in conifers than in deciduous trees, suggesting that leaf life span may be a significant correlate of plasticity. When LL-acclimated plants were exposed to HL conditions, the degree and duration of photoinhibition (measured as a decline in maximum quantum yield) was greatest in F. sylvatica, much lower in P. abies and A. alba, and lowest in A. pseudoplatanus. Thus, as with the other traits studied, vulnerability to photoinhibition showed no relationship with shade tolerance.     

Impact of snow cover on photoinhibition and winter dessication in evergreen Rhododendron ferrugineum leaves during subalpine winter

Effects of winter snow cover on photoinhibition and possible interactions with winter desiccation were investigated in situ in an evergreen subalpine woody species, Rhododendron ferrugineum L., at the alpine timberline (1950 m a.s.l.). Timing and duration of complete snow cover markedly influenced potential efficiency of photosystem II (PSII; F(v) /F(m)). Lack of snow cover led to severe but mostly reversible photoinhibition with F(v)/F(m) values as low as 0.05. Complete snow cover immediately stopped further reductions in PSII efficiency. Snow cover promoted recovery from photoinhibition, but only if, in addition to shading by snow, plants were exposed to nonfreezing temperatures close to 0 degrees C. The F(v)/F(m) ratio was closely related to minimum leaf temperatures because both photoinhibition and recovery from photoinhibition were strongly influenced by temperature. The period without major reductions in PSII efficiency lasted for only two months, reflecting the extremely short growing period in the subalpine environment. Compared with complete snow cover, incomplete snow cover led to significantly higher water losses as well as lower dehydration tolerance, because both osmotic adjustment and changes in turgor maintenance capacity were significantly reduced. Interactions between photoinhibition and winter desiccation were masked by the direct effects of freezing temperatures. However, both photoinhibition and winter desiccation are closely linked and occur together under field conditions in this evergreen subalpine woody species.     

Regulation of photosynthesis in interior spruce during water stress: changes in gas exchange and chlorophyll fluorescence

Photosynthetic response to water stress was analyzed in 1-year-old interior spruce (Picea glauca (Moench) Voss x P. engelmanni Parry hybrid complex) seedlings and emblings produced from somatic embryogenesis. Carbon dioxide uptake, oxygen evolution and chlorophyll fluorescence at 20 degrees C were monitored as predawn shoot water potential (Psi) decreased. Concurrently with stomatal closure, carbon assimilation declined rapidly as Psi decreased to -1.0 MPa. Oxygen evolution at 10,000 micro l CO(2) l(-1) declined continuously as Psi decreased to -1.6 MPa. At photon flux densities (PFD) above 50 micro mol m(-2) s(-1), photochemical efficiency of photosystem (PS) II observed during actinic light exposure (Phi(II), calculated as DeltaF/F(m)') decreased as Psi decreased. At the same PFDs, photochemical quenching (q(P)) declined with decreasing Psi and nonphotochemical quenching (q(N)) increased steadily. At PFDs below 50 micro mol m(-2) s(-1), major decreases in q(N) were not observed until Psi decreased below -1.6 MPa. We identified three phases of photosynthetic response to progressive water stress in interior spruce: a pronounced decline in gas exchange, subsequent photoprotective changes in chlorophyll fluorescence as primary photochemistry was down-regulated, and a decline in photochemical efficiency of dark-adapted needles.     

Seasonal patterns of leaf gas exchange and water relations in dry rain forest trees of contrasting leaf phenology

Diurnal and seasonal patterns of leaf gas exchange and water relations were examined in tree species of contrasting leaf phenology growing in a seasonally dry tropical rain forest in north-eastern Australia. Two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., and two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret. were studied. The deciduous species had higher specific leaf areas and maximum photosynthetic rates per leaf dry mass in the wet season than the evergreens. During the transition from wet season to dry season, total canopy area was reduced by 70-90% in the deciduous species and stomatal conductance (g(s)) and assimilation rate (A) were markedly lower in the remaining leaves. Deciduous species maintained daytime leaf water potentials (Psi(L)) at close to or above wet season values by a combination of stomatal regulation and reduction in leaf area. Thus, the timing of leaf drop in deciduous species was not associated with large negative values of daytime Psi(L) (greater than -1.6 MPa) or predawn Psi(L) (greater than -1.0 MPa). The deciduous species appeared sensitive to small perturbations in soil and leaf water status that signalled the onset of drought. The evergreen species were less sensitive to the onset of drought and g(s) values were not significantly lower during the transitional period. In the dry season, the evergreen species maintained their canopies despite increasing water-stress; however, unlike Eucalyptus species from northern Australian savannas, A and g(s) were significantly lower than wet season values.     

Application of chlorophyll fluorescence to evaluate Mn tolerance of deciduous broad-leaved tree seedlings native to northern Japan

We used chlorophyll fluorescence to examine photosynthetic responses to excess Mn accumulation in leaves of four tree species differing in successional traits. Betula ermanii Cham. (Be) and Alnus hirsuta Turcz. (Ah) were studied as representatives of early-successional species. Ulmus davidiana Planch. var. japonica (Rehder) Nakai (Ud) was selected as a mid-successional species, and Acer mono Maxim. var. glabrum (Lév. et Van't.) Hara (Am) was chosen as a late-successional species. In Be, Ah and Am, high foliar concentrations of Mn had little effect on maximum photochemical efficiency of photosystem II (PSII), as indicated by the values of dark-adapted F(v)/F(m), whereas a significant decrease was observed in Ud. Photochemical quenching (qP) and the excitation capture efficiency of open PSII (F'(v)/F'(m)) decreased with increasing leaf Mn concentration at photosynthetic steady state after a 15-min exposure to 430 &mgr;mol m(-2) s(-1) PPFD. Compared with early-successional species, these decreases were greater in mid- and late-successional species leading to lower effective quantum efficiencies of PSII (DeltaF/F'(m) = qP x F'(v)/F'(m) = (F'(m) - F)/F'(m)). To determine the extent of photoinhibition, F(v)/F(m) of the illuminated leaves was remeasured after a 15-min dark period. Compared with the dark-adapted F(v)/F(m), we observed a significant decrease in F(v)/F(m) in Am leaves containing high concentrations of Mn. These chlorophyll fluorescence studies indicate that the early-successional species Be and Ah have a higher tolerance to excessive accumulations of Mn in leaves than the mid- and late-successional species Ud and Am.     

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.     

Gas exchange, leaf structure and nitrogen in contrasting successional tree species growing in open and understory sites during a drought

Seasonal ecophysiology, leaf structure and nitrogen were measured in saplings of early (Populus grandidentata Michx. and Prunus serotina J.F. Ehrh.), middle (Fraxinus americana L. and Carya tomentosa Nutt.) and late (Acer rubrum L. and Cornus florida L.) successional tree species during severe drought on adjacent open and understory sites in central Pennsylvania, USA. Area-based net photosynthesis (A) and leaf conductance to water vapor diffusion (g(wv)) varied by site and species and were highest in open growing plants and early successional species at both the open and understory sites. In response to the period of maximum drought, both sunfleck and sun leaves of the early successional species exhibited smaller decreases in A than leaves of the other species. Shaded understory leaves of all species were more susceptible to drought than sun leaves and had negative midday A values during the middle and later growing season. Shaded understory leaves also displayed a reduced photosynthetic light response during the peak drought period. Sun leaves were thicker and had a greater mass per area (LMA) and nitrogen (N) content than shaded leaves, and early and middle successional species had higher N contents and concentrations than late successional species. In both sunfleck and sun leaves, seasonal A was positively related to predawn leaf Psi, g(wv), LMA and N, and was negatively related to vapor pressure deficit, midday leaf Psi and internal CO(2). Although a significant amount of plasticity occurred in all species for most gas exchange and leaf structural parameters, middle successional species exhibited the largest degree of phenotypic plasticity between open and understory plants.     

Patterns of leaf conductance and water potential of five Himalayan tree species

We studied variations in water relations and drought response in five Himalayan tree species (Schima wallichii (DC.) Korth. (chilaune) and Castanopsis indica (Roxb.) Miq. (dhale katus) at an elevation of 1400 m, Quercus lanata Smith (banjh) and Rhododendron arboreum Smith (lali gurans) at 2020 m, and Quercus semecarpifolia Smith (khasru) at 2130 m) at Phulchowki Hill, Kathmandu, Nepal. Soil water potential at 15 (Psi(s15)) and 30 cm (Psi(s30)) depths, tree water potential at predawn (Psi(pd)) and midday (Psi(md)), and leaf conductance during the morning (g(wAM)) and afternoon (g(wPM)) were observed from December 1998 to April 2001, except during the monsoon months. There was significant variation among sites, species and months in Psi(pd), Psi(md), g(wAM) and g(wPM), and among months for all species for Psi(s15). Mean Psi(pd) and Psi(md) were lowest in Q. semecarpifolia (-0.40 and -1.18 MPa, respectively) and highest in S. wallichii (-0.20 and -0.63 MPa, respectively). The minimum Psi value for all species (-0.70 to -1.79 MPa) was observed in March 1999, after 4 months of unusually low rainfall. Some patterns of Psi(pd) were related to phenology and leaf damage. During leafing, Psi(pd) often increased. Mean g(wAM) and g(wPM) were highest in Q. semecarpifolia (172 and 190 mmol m(-2) s(-1), respectively) and lowest in C. indica (78 and 74 mmol m(-2) s(-1), respectively). Soil water potential (Psi) at 15 cm depth correlated with plant Psi in all species, but rarely with g(wAM) and not with g(wPM). Plant Psi declined with increasing elevation, whereas g(w) increased. As Psi(pd) declined, so did maximal g(w), but overall, g(w) was correlated with Psi(pd) only for R. arboreum. Schima wallichii maintained high Psi, with low stomatal conductance, as did Castanopsis indica, except that C. indica had low Psi during dry months. Rhododendron arboreum maintained high Psi(pd) and g(w), despite low soil Psi. Quercus lanata had low g(w) and low Psi(pd) in some months, but showed no correlation between tree Psi and g(w). Quercus semecarpifolia, which grows at the highest elevation, had low soil and plant Psi and high g(w).     

Limitation of leaf carbon gain by stomatal and photochemical processes in the top canopy of Macaranga conifera,a tropical pioneer tree

Diurnal changes in gas exchange and chlorophyll fluorescence were measured in the top canopy leaves of the tropical rainforest tree species, Macaranga conifera (Zoll.) Muell. Arg. during a drought year. Maximum values of net photosynthetic rate (P(n), 10 &mgr;mol m(-2) s(-1)) and stomatal conductance (g(s), 0.2 mol m(-2) s(-1)) were found in east-facing leaves in early morning. After 1000 h, both P(n) and g(s) decreased. Minimum daytime values of P(n), g(s), and photosystem II (PSII) quantum yield (DeltaF/F(m)') were found in horizontally fixed leaves. At a given electron transport rate through PSII (ETR), P(n) was higher in early morning than at midday, suggesting a high rate of photorespiration at midday. We tested the hypothesis that the effect of low leaf temperature (T(leaf)) on P(n) is significant in the early morning, whereas the effect of low g(s) on P(n) predominates at midday. In the early morning, when T(leaf) was increased from 32 to 38 degrees C by artificial heating, P(n) at a given ETR decreased 29%, suggesting that the low T(leaf) was associated with a high P(n). When T(leaf) at midday was decreased from 37 to 32 degrees C by artificial cooling, P(n) increased 22%, but P(n) at a given ETR was higher in early morning than at midday, even at the same low T(leaf) (32 degrees C). This suggests that the rate of photorespiration was higher at midday than in early morning because low g(s) at midday caused a reduction in leaf intercellular CO(2) concentration. We conclude that low P(n) at midday was the result of both a reduction in the photochemical process and an increase in stomatal limitation.     

Response of gas exchange to water stress in seedlings of woody angiosperms

Responses of net photosynthesis (A), leaf conductance to water vapor (g(wv)) and instantaneous water use efficiency (WUE) to decreasing leaf and soil water potentials (Psi(l), Psi(s)) were studied in three-month-old white oak (Quercus alba L.), post oak (Q. stellata Wangenh.), sugar maple (Acer saccharum Marsh.), and black walnut (Juglans nigra L.) seedlings. Quercus seedlings had the highest A and g(wv) when plants were well watered. As the soil was allowed to dry, both A and g(wv) decreased; however, trace amounts of A were observed at a Psi(l) as low as -2.9 MPa in Q. stellata and -2.6 MPa in Q. alba and A. saccharum. Photosynthesis was not measurable at Psi(l) lower than -2.2 MPa in J. nigra and water stress-induced leaflet senescence was observed in this species. Within each species, g(wv) showed a similar relationship to soil and leaf Psi, but the response to Psi(l) was shifted to more negative values by 1.2 to 1.6 MPa. As Psi(s) declined below -1 MPa, the difference between soil and leaf Psi diminished because of the suppression of transpiration. There was no indication that Psi(s) had a more direct influence on g(wv) than did Psi(l). Water use efficiency showed an initial increase as the soil dried, followed by a decline under severe water stress. Water use efficiency was highest in J. nigra, intermediate in Quercus species and lowest in A. saccharum. There was an evident relationship between gas exchange characteristics and natural distribution in these species, with the more xeric species showing higher A and g(wv) under both well-watered and water-stressed conditions. There was no trend toward increased efficiency of water use in the more xeric species.     

Seasonal changes in photosynthesis and photoprotection in a Quercus ilex subsp. ballota woodland located in its upper altitudinal extreme in the Iberian Peninsula

Quercus ilex L. subsp. ballota (Desf.) Samp., a Mediterranean evergreen species growing in a continental Mediterranean climate, did not experience water stress and showed greater sensitivity to winter stress than to summer stress over a 12-month period. Net CO2 assimilation rates and photosystem II (PSII) efficiency decreased markedly during the cold months and recovered completely in spring. Lutein, neoxanthin and beta-carotene to chlorophyll (Chl) molar ratios all showed the same trend throughout the year, increasing from September to March. This increase was a result of increases in carotenoid concentrations, because Chl concentration per unit leaf area remained stable, and was higher at the end than at the beginning of the first growing season. Lutein-epoxide was a minor component of the total lutein pool. Thermal energy dissipation and non-photochemical quenching (NPQ) were associated with the de-epoxidated forms of the xanthophyll cycle pigments in the warm months. Photosynthetic rates decreased slightly at midday in summer. These changes were accompanied by decreases in maximum potential PSII efficiency (which recovered during the night), actual and intrinsic PSII efficiencies, photochemical quenching and increases in NPQ. Overall, our data indicate down-regulation of photosynthesis during the summer. The diurnal de-epoxidation of violaxanthin to antheraxanthin and zeaxanthin occurred throughout the year, except in January. Antioxidant enzymatic activity increased in the winter months, especially during the coldest months, highlighting its key role in photoprotection against photo-oxidation. Structural and functional modifications protected PSII from permanent damage and allowed 1-year-old leaves to photosynthesize at high rates when temperatures increased in spring.     

Factors involved in alleviating water stress by partial crop removal in pear trees

We studied the relief of water stress associated with fruit thinning in pear (Pyrus communis L.) trees during drought to determine what mechanisms, other than stomatal adjustment, were involved. Combinations of control irrigation (equal to crop water use less effective rainfall) and deficit irrigation (equal to 20% of control irrigation), fruit load (unthinned and thinned to 40 fruits per tree) and root pruning (pruned and unpruned) treatments were applied to pear (cv. 'Conference') trees during Stage II of fruit development. Daily patterns of midday stem water potential (Psi(stem)) and leaf conductance to water vapor (g(l)) of deficit-irrigated trees differed after fruit thinning. In response to fruit thinning, gl progressively declined with water stress until 30 days after fruit thinning and then leveled off, whereas the effects of decreased fruit load on Psi(stem) peaked 30-40 days after fruit thinning and then tended to decline. Soil water depletion was significantly correlated with fruit load during drought. Our results indicate that stomatal adjustment and the resulting soil water conservation were the factors determining the Psi(stem) response to fruit thinning. However, these factors could not explain differences in daily patterns between g(l) and Psi(stem) after fruit thinning. In all cases, effects of root pruning treatments on Psi(stem) in deficit-irrigated trees were transitory (Psi(stem) recovered from root pruning in less than 30 days), but the recovery of Psi(stem) after root pruning was faster in trees with low fruit loads. This behavior is compatible with the concept that the water balance (reflected by Psi(stem) values) was better in trees with low fruit loads compared with unthinned trees, perhaps because more carbon was available for root growth. Thus, a root growth component is hypothesized as a mechanism to explain the bimodal Psi(stem) response to fruit thinning during drought.