Transport,storage and mobilization of nitrogen by trees and shrubs in the wet/dry tropics of northern Australia

Xylem sap from woody species in the wet/dry tropics of northern Australia was analyzed for N compounds. At the peak of the dry season, arginine was the main N compound in sap of most species of woodlands and deciduous monsoon forest. In the wet season, a marked change occurred with amides becoming the main sap N constituents of most species. Species from an evergreen monsoon forest, with a permanent water source, transported amides in the dry season. In the dry season, nitrate accounted for 7 and 12% of total xylem sap N in species of deciduous and evergreen monsoon forests, respectively. In the wet season, the proportion of N present as nitrate increased to 22% in deciduous monsoon forest species. These results suggest that N is taken up and assimilated mainly in the wet season and that this newly assimilated N is mostly transported as amide-N (woodland species, monsoon forest species) and nitrate (monsoon forest species). Arginine is the form in which stored N is remobilized and transported by woodland and deciduous monsoon forest species in the dry season. Several proteins, which may represent bark storage proteins, were detected in inner bark tissue from a range of trees in the dry season, indicating that, although N uptake appears to be limited in the dry season, the many tree and shrub species that produce flowers, fruit or leaves in the dry season use stored N to support growth. Nitrogen characteristics of the studied species are discussed in relation to the tropical environment.     

Hydraulic redistribution of soil water by neotropical savanna trees

The magnitude and direction of water transport by the roots of eight dominant Brazilian savanna (Cerrado) woody species were determined with a heat pulse system that allowed bidirectional measurements of sap flow. The patterns of sap flow observed during the dry season in species with dimorphic root systems were consistent with the occurrence of hydraulic redistribution of soil water, the movement of water from moist to drier regions of the soil profile via plant roots. In these species, shallow roots exhibited positive sap flow (from the soil into the plant) during the day and negative sap flow (from the plant into the soil) during the night. Sap flow in the taproots was positive throughout the 24-h period. Diel fluctuations in soil water potential, with maximum values occurring at night, provided evidence for partial rewetting of upper soil layers by water released from shallow roots. In other species, shallow roots exhibited negative sap flow during both the day and night, indicating that hydraulic redistribution was occurring continuously. A third sap flow pattern was observed at the end of the dry season after a heavy rainfall event when sap flow became negative in the taproot, and positive in the small roots, indicating movement of water from upper soil layers into shallow roots, and then into taproots and deeper soil layers. Experimental manipulations employed to evaluate the response of hydraulic redistribution to changes in plant and environmental conditions included watering the soil surface above shallow roots, decreasing transpiration by covering the plant and cutting roots where probes were inserted. Natural and manipulated patterns of sap flow in roots and stems were consistent with passive movement of water toward competing sinks in the soil and plant. Because dry shallow soil layers were often a stronger sink than the shoot, we suggest that the presence of a dimorphic root system in deciduous species may play a role in facilitating leaf expansion near the end of the dry season when the soil surrounding shallow lateral roots is still dry.     

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.     

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.     

Soil depth and soil water regime in a catchment where tropical dry evergreen and deciduous forests coexist

In several areas in Northeast Thailand, evergreen and deciduous forests coexist under uniform climatic conditions. To identify the factors that determine the distribution of these different forest types, we compared soil depth and soil physical properties between evergreen and deciduous forests, and monitored soil moisture conditions for a year in both forest types at the Sakaerat Environmental Research Station. The soil was significantly deeper under the evergreen forests (mean 97 cm) than under the deciduous forest (mean 64 cm). The soil under the evergreen forests retained much more water throughout the year than the soil under the deciduous forest, and there was also a clear tendency for the evergreen forests to occur in ravine areas, regardless of soil depth. It is possible that the evergreen trees can maintain transpiration during the dry season on thicker soils or in ravine areas, whereas shallower soils cannot provide enough water for these trees to maintain their evapotranspiration during the dry period. From the present study, we showed that soil water availability could be a significant factor determining the distribution of the deciduous and evergreen forests in our catchments.     

Variation in soil water uptake and its effect on plant water status in Juglans regia L. during dry and wet seasons

Temporal and spatial variations in the water status of walnut trees (Juglans regia L.) and the soil in which they were growing were traced by analyzing the differences in hydrogen isotopes during spring and summer in a 7-year-old walnut stand. Walnut root dynamics were measured in both dry and wet seasons. Walnut roots were mainly distributed in the upper soil (0-30 cm depth), with around 60% of the total root mass in upper soil layers and 40% in deep soil layers (30-80 cm depth). The upper soil layers contributed 68% of the total tree water requirement in the wet season, but only 47% in the dry season. In the wet season, total roots, living roots and new roots were all significantly more abundant than in the dry season. There were significant differences in pre-dawn branch percentage loss of hydraulic conductance (PLC), pre-dawn leaf water potential and transpiration between the dry and wet seasons. Water content in the upper soil layers remarkably influenced xylem water stable-hydrogen isotope (δD) values. Furthermore, there were linear relationships between the xylem water δD value and pre-dawn branch PLC, pre-dawn leaf water potential, transpiration rate and photosynthetic rate. In summary, J. regia was compelled to take a larger amount of water from the deep soil layers in the dry season, but this shift could not prevent water stress in the plant. The xylem water δD values could be used as an indicator to investigate the water stress of plants, besides probing profiles of soil water use.     

Comparison of soil depths between evergreen and deciduous forests as a determinant of their distribution, Northeast Thailand

In several areas in Northeast Thailand, evergreen and deciduous forests coexist under uniform terrain and climatic conditions. We compared depth and physical properties of soils between evergreen and deciduous forests in the Sakaerat Environmental Research Station to clarify what factor determines their distribution. The averaged soil depths were 79 ± 27 cm and 135 ± 58 cm in the deciduous and evergreen forests, respectively. The soils in the deciduous forests were relatively coarser in soil texture than those in the evergreen forests, particularly in the surface layers. The average available water capacity of the solum was lower in the deciduous forest soils (78 mm) than in the evergreen forest soils (123 mm). Compared with the evapotranspiration from the evergreen forest in the study area, the available water capacity of the evergreen forest soil was almost the same as the water deficit during the dry season (November–February), while that of the deciduous forest soil was lower and insufficient to maintain the evapotranspiration. These results suggest that the distribution of deciduous and evergreen forests in the study area was associated with soil water availability, which mainly depends on soil depth.     

Water economy of Neotropical savanna trees: six paradigms revisited

Biologists have long been puzzled by the striking morphological and anatomical characteristics of Neotropical savanna trees which have large scleromorphic leaves, allocate more than half of their total biomass to belowground structures and produce new leaves during the peak of the dry season. Based on results of ongoing interdisciplinary projects in the savannas of central Brazil (cerrado), we reassessed the validity of six paradigms to account for the water economy of savanna vegetation. (1) All savanna woody species are similar in their ability to take up water from deep soil layers where its availability is relatively constant throughout the year. (2) There is no substantial competition between grasses and trees for water resources during the dry season because grasses exclusively explore upper soil layers, whereas trees access water in deeper soil layers. (3) Tree species have access to abundant groundwater, their stomatal control is weak and they tend to transpire freely. (4) Savanna trees experience increased water deficits during the dry season despite their access to deep soil water. (5) Stomatal conductance of savanna species is low at night to prevent nocturnal transpiration, particularly during the dry season. (6) Savanna tree species can be classified into functional groups according to leaf phenology. We evaluated each paradigm and found differences in the patterns of water uptake between deciduous and evergreen tree species, as well as among evergreen tree species, that have implications for regulation of tree water balance. The absence of resource interactions between herbaceous and woody plants is refuted by our observation that herbaceous plants use water from deep soil layers that is released by deep-rooted trees into the upper soil layer. We obtained evidence of strong stomatal control of transpiration and show that most species exhibit homeostasis in maximum water deficit, with midday water potentials being almost identical in the wet and dry seasons. Although stomatal control is strong during the day, nocturnal transpiration is high during the dry season. Our comparative studies showed that the grouping of species into functional categories is somewhat arbitrary and that ranking species along continuous functional axes better represents the ecological complexity of adaptations of cerrado woody species to their seasonal environment.     

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.     

Processes preventing nocturnal equilibration between leaf and soil water potential in tropical savanna woody species

The impact of nocturnal water loss and recharge of stem water storage on predawn disequilibrium between leaf (psiL) and soil (psiS) water potentials was studied in three dominant tropical savanna woody species in central Brazil (Cerrado). Sap flow continued throughout the night during the dry season and contributed from 13 to 28% of total daily transpiration. During the dry season, psiL was substantially less negative in covered transpiring leaves, throughout the day and night, than in exposed leaves. Before dawn, differences in psiL between covered and exposed leaves were about 0.4 MPa. When relationships between sap flow and psiL of exposed leaves were extrapolated to zero flow, the resulting values of psiL (a proxy of weighted mean soil water potential) in two of the species were similar to predawn values of covered leaves. Consistent with substantial nocturnal sap flow, stomatal conductance (gs) never dropped below 40 mmol m(-2) s(-1) at night, and in some cases, rose to as much as 100 mmol m(-2) s(-1) before the end of the dark period. Nocturnal gs decreased linearly with increasing air saturation deficit (D), but there were species-specific differences in the slopes of the relationships between nocturnal gs and D. Withdrawal and recharge of water from stem storage compartments were assessed by monitoring diel fluctuations of stem diameter with electronic dendrometers. Stem water storage compartments tended to recharge faster when nocturnal transpiration was reduced by covering the entire plant. Water potential of covered leaves did not stabilize in any of the plants before the end of the dark period, suggesting that, even in covered plants, water storage tissues were not fully rehydrated by dawn. Patterns of sap flow and expansion and contraction of stems reflected the dynamics of water movement during utilization and recharge of stem water storage tissues. This study showed that nighttime transpiration and recharge of internal water storage contribute to predawn disequilibrium in water potential between leaves and soil in neotropical savanna woody plants.     

Environmental and physiological controls over oxygen and carbon isotope composition of Tasmanian blue gum, Eucalyptus globulus

We measured oxygen isotope ratios (delta18O) of xylem sap, phloem sap, leaves, wood and bark of Eucalyptus globulus Labill. growing in southwestern Australia. Carbon isotope ratios (delta13C) were measured in the dry matter of phloem sap, leaves and wood. Results were used to test several aspects of a mechanistic model of 18O enrichment and provided insights into post-photosynthetic variations in dry matter delta13C. Xylem water delta18O varied little within the tree crown, whereas variation at the landscape-level was more pronounced, with plantations near the coast being enriched by up to 3 per thousand compared with plantations less than 100 km inland. Phloem water was significantly enriched in 18O compared with xylem water in two of three sampling campaigns; mean enrichments were 0.5 and 0.8 per thousand. Phloem sap sugars exported from E. globulus leaves closely reflected observed leaf water enrichment when diurnal variation in photosynthesis was taken into account. Photosynthetic rates were higher in the morning than in the afternoon, whereas leaf water 18O enrichment increased to maximum values in the afternoon. A non-steady-state model of leaf water 18O enrichment accurately predicted observed values through a full diel cycle. Mean estimates of the proportion of organic oxygen effectively exchanging with xylem water during cellulose synthesis were close to 0.40 for both leaves and wood. Carbon isotope ratios of nascent xylem tissues did not differ from those of phloem sap sugars collected concurrently, whereas nascent leaf tissues were depleted in 13C by 2 per thousand compared with phloem sap sugars, suggesting that, in E. globulus, 13C enrichment of sink tissues compared with source leaves does not result from an enriching process within the sink tissue.     

Hydraulic conductivity, photosynthesis and leaf water balance in six evergreen woody species from fall to winter

To confirm that freeze-thaw embolism is a primary stress for evergreen woody species in winter, hydraulic conductivity, photosynthesis and leaf water potential were measured during fall and winter in trees growing in a cool temperate zone (Nikko) and in a warm temperate zone (Tokyo). We examined two evergreen conifers that naturally occur in the cool temperate zone (Abies firma Siebold & Zucc. and Abies homolepis Siebold & Zucc.), and four evergreen broad-leaved woody species that are restricted to the warm temperate zone (Camellia japonica L., Cinnamomum camphora (L.) J. Presl, Ilex crenata Thunb. and Quercus myrsinaefolia Blume). In Tokyo, where no freeze-thaw cycles of xylem sap occurred, hydraulic conductivity, photosynthesis and water balance remained constant during the experimental period. In Nikko, where there were 38 daily freeze-thaw cycles by February, neither of the tracheid-bearing evergreen conifers showed xylem embolism or leaf water deficits. Similarly, the broad-leaved evergreen trees with small-diameter vessels did not exhibit severe embolism or water deficits and maintained CO(2) assimilation even in January. In contrast, the two broad-leaved evergreen trees with large-diameter vessels showed significantly reduced hydraulic conductivity and shoot die-back in winter. We conclude that freeze-thaw embolism restricts evergreen woody species with large-diameter vessels to the warm temperate zone, whereas other stresses limit the distribution of broad-leaved trees, that have small-diameter vessels, but which are restricted to the warm temperate zone.     

Seasonal changes in photosynthesis of eight savanna tree species

Seasonal variations in carbon assimilation of eight tree species of a north Australian tropical savanna were examined over two wet seasons and one dry season (18 months). Assimilation rates (A) in the two evergreen species, Eucalyptus tetrodonta F. Muell. and E. miniata A. Cunn. ex Schauer, were high throughout the study although there was a 10-20% decline in the dry season compared with the wet season. The three semi-deciduous species (Erythrophleum chlorostachys (F. Muell.) Baillon, Eucalyptus clavigera A. Cunn. ex Schauer, and Xanthostemon paradoxus F. Muell.) showed a 25-75% decline in A in the dry season compared with the wet season, and the deciduous species (Terminalia ferdinandiana Excell, Planchonia careya (F. Muell.) Kunth, and Cochlospermum fraseri Planchon) were leafless for several months in the dry season. Generally, the ratio of intercellular CO(2) concentration to ambient CO(2) concentration (C(i):C(a)) was larger in the wet season than in the dry season, indicating a smaller stomatal limitation of photosynthesis in the wet season compared with the dry season. In all species, the C(i):C(a) ratio and A were essentially independent of leaf-to-air vapor pressure difference (LAVPD) during the wet season, but both parameters generally declined with increasing LAVPD in the dry season. The slope of the positive correlation between A and transpiration rate (E) was less in the wet season than in the dry season. There was no evidence that high E inhibited A. Instantaneous transpiration efficiency was lowest in the wet season and highest during the dry season. Nitrogen-use efficiency (NUE) was higher in the wet season than in the dry season because the decline in A in the dry season was proportionally larger than the decline in foliar nitrogen content. In the wet season, evergreen species exhibited higher NUE than semi-deciduous and deciduous species. In all species, A was linearly correlated with specific leaf area (SLA) and foliar N content. Foliar N content increased with increasing SLA. All species showed a decline in midday leaf water potential as the dry season progressed. Dry season midday water potentials were lowest in semi-deciduous species and highest in the deciduous species, with evergreen species exhibiting intermediate values.     

Patterns of hydraulic architecture and water relations of two tropical canopy trees with contrasting leaf phenologies: Ochroma pyramidale and Pseudobombax septenatum

Many authors have attempted to explain the adaptive response of tropical plants to drought based on studies of water relations at the leaf level. Little attention has been given to the role of the xylem system in the control of plant water requirements. To evaluate this role, we studied the hydraulic architecture and water relations parameters of two tropical canopy trees with contrasting leaf phenologies: deciduous Pseudobombax septenatum (Jacq.) Dug and evergreen Ochroma pyramidale (Cav. ex lamb) Urban, both in the family Bombacaceae. The hydraulic architecture parameters studied include hydraulic conductivity, specific conductivity, leaf specific conductivity, and Huber value. Water relations parameters include leaf water potential, stem and leaf water storage capacitance, transpiration, stomatal conductance, and vulnerability of stems to cavitation and loss of hydraulic conductivity by embolisms. Compared to temperate trees, both species showed a pattern of highly vulnerable stems (50% loss of conductivity due to embolism at water potentials less than 1 MPa) with high leaf specific conductivities. The vulnerability of xylem to water-stress-induced embolism was remarkably similar for the two species but the leaf specific conductivity of petioles and leaf-bearing stems of the evergreen species, Ochroma (e.g., 9.08 and 11.4 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively), were 3.4 and 2.3 times higher, respectively, than those of the deciduous species, Pseudobombax (e.g., 2.64 and 5.15 x 10(-4) kg s(-1) m(-1) MPa(-1), respectively). A runaway embolism model was used to test the ability of Ochroma and Pseudobombax stems to maintain elevated transpiration rates during the higher evaporative demand of the dry season. The percent loss of leaf area predicted by the runaway embolism model for stems of Pseudobombax ranged from 5 to 30%, not enough to explain the deciduous phenology of this tree species without analysis of root resistance or leaf and petiole vulnerability to embolism.     

Stable annual pattern of water use by Acacia tortilis in Sahelian Africa

Water use by mature trees of Acacia tortilis (Forsk.) Hayne ssp. raddiana (Savi) Brenan var. raddiana growing in the northern Sahel was continuously recorded over 4 years. Water use was estimated from xylem sap flow measured by transient heat dissipation. Concurrently, cambial growth, canopy phenology, leaf water potential, climatic conditions and soil water availability (SWA) were monitored. In addition to the variation attributable to interannual variation in rainfall, SWA was increased by irrigation during one wet season. The wet season lasted from July to September, and annual rainfall ranged between 146 and 367 mm. The annual amount and pattern of tree water use were stable from year-to-year despite interannual and seasonal variations in SWA in the upper soil layers. Acacia tortilis transpired readily throughout the year, except for one month during the dry season when defoliation was at a maximum. Maximum water use of about 23 l (dm sapwood area)(-2) day(-1) was recorded at the end of the wet season. While trees retained foliage in the dry season, the decline in water use was modest at around 30%. Variation in predawn leaf water potential indicated that the trees were subject to soil water constraint. The rapid depletion of water in the uppermost soil layers after the wet season implies that there was extensive use of water from deep soil layers. The deep soil profile revealed (1) the existence of living roots at 25 m and (2) that the availability of soil water was low (-1.6 MPa) down to the water table at a depth of 31 m. However, transpiration was recorded at a predawn leaf water potential of -2.0 MPa, indicating that the trees used water from both intermediary soil layers and the water table. During the full canopy stage, mean values of whole-tree hydraulic conductance were similar in the wet and dry seasons. We propose that the stability of water use at the seasonal and annual scales resulted from a combination of features, including an extensive rooting habit related to deep water availability and an effective regulation of canopy conductance. Despite a limited effect on tree water use, irrigation during the wet season sharply increased predawn leaf water potential and cambial growth of trunks and branches.     

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.     

Drought tolerance,xylem sap abscisic acid and stomatal conductance during soil drying: a comparison of canopy trees of three temperate deciduous angiosperms

Patterns of water relations, xylem sap abscisic acid concentration ([ABA]) and stomatal aperture were characterized and compared in drought-sensitive black walnut (Juglans nigra L.), less drought-sensitive sugar maple (Acer saccharum Marsh.) and drought-tolerant white oak (Quercus alba L.) trees co-occurring in a second-growth forest in Missouri, USA. There were strong correlations among reduction in predawn leaf water potential, increased xylem sap [ABA] and stomatal closure in all species. Stomatal conductance was more closely correlated with xylem sap ABA concentration than with ABA flux or xylem sap pH and cation concentrations. In isohydric black walnut, increased concentrations of ABA in the xylem sap appeared to be primarily of root origin, causing stomatal closure in response to soil drying. In anisohydric sugar maple and white oak, however, there were reductions in midday leaf water potential associated with stomatal closure, making it uncertain whether drought-induced xylem sap ABA was of leaf or root origin. The role of root-originated xylem sap ABA in these species as a signal to the shoot of the water status of the roots is, therefore, less certain.     

Phenological differences in tree water use and the timing of tropical forest inventories: conclusions from patterns of dry season diameter change

Interspecific variation in water-induced fluctuations in stem girth demonstrates the mechanisms promoting coexistence in seasonally dry tropical forest. In addition, these fluctuations are a potential, but unevaluated, source of bias in measurements of annual tree growth rates. To examine diurnal and seasonal patterns of stem diameter change, tree girth was measured over 2 years (1997–1999), using dendrometer bands, for three species (Celtis mildbraedii, C. zenkeri and Strombosia glaucescens) in semi-deciduous forest in Ghana. Soil matric potential was measured concurrently at 15 cm depth. In addition, measurements of all trees >20 cm dbh on three, 1 ha plots were made at the beginning and middle of the 1998/1999 dry season. During the severe 1997/1998 dry season, soil matric potential declined below −1.5 MPa and two species showed significant stem shrinkage. For the evergreen species, C. mildbraedii, there was a significant positive effect of tree diameter on stem shrinkage, and shrinkage was greater in the second, compared to the first, half of the dry season. For the deciduous species, C. zenkeri, shrinkage was reduced during the second half of the dry season, following leaf fall. During 1998/1999, soil matric potential, did not decline below −1.5 MPa, and rates of girth change remained positive for all species. There were no significant effects of size or phenology on the rate of girth change in the plot-based study. Deviations in annual increment calculated over successive monthly intervals indicate that a 10-fold difference in soil water availability between measurement occasions can lead to a 4% bias in estimates of annual growth. Measurements of forest plots should be made when inter-annual variation in soil water availability is low. In this forest, measurements should, therefore, be made during the wet season, contrary to published recommendations.     

Functional convergence in hydraulic architecture and water relations of tropical savanna trees: from leaf to whole plant

Functional convergence in hydraulic architecture and water relations, and potential trade-offs in resource allocation were investigated in six dominant neotropical savanna tree species from central Brazil during the peak of the dry season. Common relationships between wood density and several aspects of plant water relations and hydraulic architecture were observed. All species and individuals shared the same negative exponential relationship between sapwood saturated water content and wood density. Wood density was a good predictor of minimum (midday) leaf water potential and total daily transpiration, both of which decreased linearly with increasing wood density for all individuals and species. With respect to hydraulic architecture, specific and leaf-specific hydraulic conductivity decreased and the leaf:sapwood area ratio increased more than 5-fold as wood density increased from 0.37 to 0.71 g cm(-3) for all individuals and species. Wood density was also a good predictor of the temporal dynamics of water flow in stems, with the time of onset of sap flow in the morning and the maximum sap flow tending to occur progressively earlier in the day as wood density increased. Leaf properties associated with wood density included stomatal conductance, specific leaf area, and osmotic potential at the turgor loss point, which decreased linearly with increasing wood density. Wood density increased linearly with decreasing bulk soil water potential experienced by individual plants during the dry season, suggesting that wood density was greatest in individuals with mostly shallow roots, and therefore limited access to more abundant soil water at greater depths. Despite their taxonomic diversity and large intrapopulation differences in architectural traits, the six co-occurring species and their individuals shared similar functional relationships between all pairs of variables studied. Thus, rather than differing intrinsically in physiological responsiveness, the species and the individuals appeared to have distinct operating ranges along common physiological response curves dictated by plant architectural and structural features. The patterns of water uptake and access to soil water during the dry season appeared to be the main determinant of wood density, which constrained evolutionary options related to plant water economy and hydraulic architecture, leading to functional convergence in the neotropical savanna trees studied.     

Comparative water use by the riparian trees Melaleuca argentea and Corymbia bella in the wet-dry tropics of northern Australia

We examined sources of water and daily and seasonal water use patterns in two riparian tree species occupying contrasting niches within riparian zones throughout the wet-dry tropics of northern Australia: Corymbia bella Hill and Johnson is found along the top of the levee banks and Melaleuca argentea W. Fitzg. is restricted to riversides. Patterns of tree water use (sap flow) and leaf water potential were examined in four trees of each species at three locations along the Daly River in the Northern Territory. Predawn leaf water potential was higher than -0.5 MPa throughout the dry season in both species, but was lower at the end of the dry season than at the beginning of the dry season. Contrary to expectations, predawn leaf water potential was lower in M. argentea trees along the river than in C. bella trees along the levees. In contrast, midday leaf water potential was lower in the C. bella trees than in M. argentea trees. There were no seasonal differences in tree water use in either species. Daily water use was lower in M. argentea trees than in C. bella trees. Whole-tree hydraulic conductance, estimated from the slope of the relationship between leaf water potential and sap flow, did not differ between species. Xylem deuterium concentrations indicated that M. argentea trees along the riverbank were principally reliant on river water or shallow groundwater, whereas C. bella trees along the levee were reliant solely on soil water reserves. This study demonstrated strong gradients of tree water use within tropical riparian communities, with implications for estimating riparian water use requirements and for the management of groundwater resources.