Competition for water between walnut seedlings (Juglans regia) and rye grass (Lolium perenne) assessed by carbon isotope discrimination and delta18O enrichment

Container-grown walnut seedlings (Juglans regia L.) were subjected to competition with rye grass (Lolium perenne L.) and to a 2-week soil drying cycle. One and 2 weeks after the beginning of the drought treatment, H2 18O (delta approximately equals +100%) was added to the bottom layer of soil in the plant containers to create a vertical H2 18O gradient. Rye grass competition reduced aboveground and belowground biomass of the walnut seedlings by 60%, whereas drought had no effect. The presence of rye grass reduced the dry weight of walnut roots in the upper soil layer and caused a 50% reduction in lateral root length. Rye grass competition combined with the drought treatment reduced walnut leaf CO2 assimilation rate (A) and leaf conductance (gw) by 20 and 39%, respectively. Transpiration rates in rye grass, both at the leaf level and at the plant or tiller level, were higher than in walnut seedlings. Leaf intrinsic water-use efficiency (A/gw) of walnut seedlings increased in response to drought and no differences were observed between the single-species and mixed-species treatments, as confirmed by leaf carbon isotope discrimination measurements. Measurement of delta18O in soil and in plant xylem sap indicated that the presence of rye grass did not affect the vertical profile of soil water uptake by walnut seedlings. Walnut seedlings and rye grass withdrew water from the top and middle soil layers in well-watered conditions, whereas during the drought treatment, walnut seedlings obtained water from all soil layers, but rye grass took up water from the bottom soil layer only.     

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.     

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.     

Dry season conditions determine wet season water use in the wet-tropical savannas of northern Australia

Daily and seasonal patterns of transpiration were measured in evergreen eucalypt trees growing at a wet (Darwin), intermediate (Katherine) and dry site (Newcastle Waters) along a steep rainfall gradient in a north Australian savanna. Relationships between tree size and tree water use were also determined. Diameter at breast height (DBH) was an excellent predictor of sapwood area in the five eucalypt species sampled along the rainfall gradient. A single relationship existed for all species at all sites. Mean daily water use was also correlated to DBH in both wet and dry seasons. There were no significant differences in the relationship between DBH and tree water use at Darwin or Katherine. Among the sites, tree water use was lowest at Newcastle Waters at all DBHs. The relationship between DBH and tree leaf area was similar between species and locations, but the slope of the relationship was less at the end of the dry season than at the end of the wet season at all locations. There was a strong relationship between sapwood area and leaf area that was similar at all sites along the gradient. Transpiration rates were significantly lower in trees at the driest site than at the other sites, but there were no significant differences in transpiration rates between trees growing at Darwin and Katherine. Transpiration rates did not vary significantly between seasons at any site. At all sites, there was only a 10% decline in water use per tree between the wet and dry seasons. A monthly aridity index (pan evaporation/rainfall) and predawn leaf water potential showed strong seasonal patterns. It is proposed that dry season conditions exert control on tree water use during the wet season, possibly through an effect on xylem structure.     

Soil-plant hydrology of indigenous and exotic trees in an Ethiopian montane forest

Fast-growing exotic trees are widely planted in the tropics to counteract deforestation; however, their patterns of water use could be detrimental to overall ecosystem productivity through their impact on ecosystem water budget. In a comparative field study on seasonal soil-plant water dynamics of two exotic species (Cupressus lusitanica Mill. and Eucalyptus globulus Labill.) and the indigenous Podocarpus falcatus (Thunb.) Mirb. in south Ethiopia, we combined a 2.5-year record for climate and soil water availability, natural-abundance oxygen isotope ratios (delta(18)O) of soil and xylem water, destructive root sampling and transpiration measurements. Soil was generally driest under C. lusitanica with its dense canopy and shallow root system, particularly following a relatively low-rainfall wet season, with the wettest soil under E. globulus. Wet season transpiration of C. lusitanica was twice that of the other species. In the dry season, P. falcatus and C. lusitanica reduced transpiration by a factor of six and two, respectively, whereas E. globulus showed a fivefold increase. In all species, there was a shift in water uptake to deeper soil layers as the dry season progressed, accompanied by relocation of live fine root biomass (LFR) of C. lusitanica and P. falcatus to deeper layers. Under P. falcatus, variability in soil matric potential, narrow delta(18)O depth gradients and high LFR indicated fast water redistribution. Subsoil water uptake was important only for E. globulus, which had low topsoil LFR and tap roots exploiting deep water. Although P. falcatus appeared better adapted to varying soil water availability than the exotic species, both conifers decreased growth substantially during dry weather. Growth of E. globulus was largely independent of topsoil water content, giving it the potential to cause substantial dry-season groundwater depletion.     

Partitioning of soil water among tree species in a Brazilian Cerrado ecosystem

Source water used by woody perennials in a Brazilian savanna (Cerrado) was determined by comparing the stable hydrogen isotope composition (deltaD) of xylem sap and soil water at different depths during two consecutive dry seasons (1995 and 1996). Plant water status and rates of water use were also determined and compared with xylem water deltaD values. Overall, soil water deltaD decreased with increasing depth in the soil profile. Mean deltaD values were -35 per thousand for the upper 170 cm of soil and -55 per thousand between 230 and 400 cm depth at the end of the 1995 dry season. Soil water content increased with depth, from 18% near the surface to about 28% at 400 cm. A similar pattern of decreasing soil water deltaD with increasing depth was observed at the end of the 1996 dry season. Patterns consistent with hydraulic lift were observed in soil profiles sampled in 1995 and 1997. Concurrent analyses of xylem and soil water deltaD values indicated a distinct partitioning of water resources among 10 representative woody species (five deciduous and five evergreen). Among these species, four evergreen and one deciduous species acquired water primarily in the upper soil layers (above 200 cm), whereas three deciduous and one evergreen species tapped deep sources of soil water (below 200 cm). One deciduous species exhibited intermediate behavior. Total daily sap flow was negatively correlated with xylem sap deltaD values indicating that species with higher rates of water use during the dry season tended to rely on deeper soil water sources. Among evergreen species, minimum leaf water potentials were also negatively correlated with xylem water deltaD values, suggesting that access to more readily available water at greater depth permitted maintenance of a more favorable plant water status. No significant relationship between xylem water deltaD and plant size was observed in two evergreen species, suggesting a strong selective pressure for small plants to rapidly develop a deep root system. The degree of variation in soil water partitioning, leaf phenology and leaf longevity was consistent with the high diversity of woody species in the Cerrado.     

Daily and seasonal patterns of carbon and water fluxes above a north Australian savanna

Daily and seasonal fluxes of carbon dioxide and water vapor above a north Australian savanna were recorded over a complete dry season-wet season annual cycle using the eddy covariance technique. Wet season rates of photosynthesis and transpiration were larger than those measured in the dry season and were dominated by the presence of the grassy understory. As the dry season progressed and the grass understory died, ecosystem rates of assimilation and water vapor flux declined substantially. By the end of the dry season, canopy assimilation and evapotranspiration rates were 20-25% of wet season values. Assimilation was light saturated in the dry season but not in the wet season. Stomatal control of transpiration increased between the wet and dry season. This was revealed by the decline in the slope of E with increasing leaf-to-air vapor pressure difference (D) between wet and dry seasons, and also by the significant decrease in the ratio of boundary to canopy conductance observed between the wet and dry seasons. A simple pan-tropical modeling of leaf area index or wet season canopy CO2 flux was undertaken. It was shown that with readily available data for foliar N content and the ratio of rainfall to potential evaporation, leaf index and wet season canopy CO2 flux can be successfully estimated for a number of tropical ecosystems, including north Australian savannas.     

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.     

Winter embolism,mechanisms of xylem hydraulic conductivity recovery and springtime growth patterns in walnut and peach trees

Xylem vessels of Prunus persica Batsch (peach) and Juglans regia L. (walnut) are vulnerable to frost-induced embolism. In peach, xylem embolism increased progressively over the winter, reaching a maximum of 85% loss of hydraulic conductivity (PLC) in early March. Over winter, PLC in walnut approached 100%, but the degree of xylem embolism varied during the winter, reflecting the ability of walnut to generate positive xylem pressures in winter and spring. In contrast, positive xylem pressures were not observed in peach. Controlled freeze-thaw experiments showed that frost alone is insufficient to increase embolism in peach; evaporative conditions during thawing are also required. However, when both species were protected from frost, PLC was zero. At bud break, there was complete recovery from embolism in walnut, whereas PLC remained high in peach. Three mechanisms responsible for the restoration of branch hydraulic conductivity were identified in walnut: the development of stem pressure, the development of root pressure and the formation of a new ring of functional xylem, whereas only one mechanism was observed in peach (new functional ring). The climatic conditions necessary for the manifestation of these mechanisms were investigated.     

Transpiration increases during the dry season: patterns of tree water use in eucalypt open-forests of northern Australia

Australian savannas exhibit marked seasonality in precipitation, with more than 90% of the annual total falling between October and May. The dry season is characterized by declining soil water availability and high vapor pressure deficits (up to 2.5 kPa). We used heat pulse technology to measure whole-tree transpiration rates on a daily and seasonal basis for the two dominant eucalypts at a site near Darwin, Australia. Contrary to expectations, transpiration rates were higher during the dry season than during the wet season, largely because of increased evaporative demand and the exploitation of groundwater reserves by the trees. Transpiration rates exhibited a marked hysteresis in relation to vapor pressure deficit, which was more marked in the dry season than in the wet season. This result may be attributable to low soil hydraulic conductivity, or the use of stored stem water, or both. Tree water use was strongly correlated with leaf area and diameter at breast height and there were no differences in transpiration between the species studied. These results are discussed in relation to scaling tree water use to stand water use.     

Growth and transpiration of Japanese cedar (Cryptomeria japonica) and Hinoki cypress (Chamaecyparis obtusa) seedlings in response to soil water content

To investigate the effects of soil water content on growth and transpiration of Japanese cedar (Cryptomeria japonica D. Don) and Hinoki cypress (Chamaecyparis obtusa (Siebold et Zucc.) Endl.), potted seedlings were grown in well-watered soil (wet treatment) or in drying soil (dry treatment) for 12 weeks. Seedlings in the wet treatment were watered once every 2 or 3 days, whereas seedlings in the dry treatment were watered when soil water content (Theta; m3 m(-3)) reached 0.30, equivalent to a soil matric potential of -0.06 MPa. From Weeks 7 to 12 after the onset of the treatments, seedling transpiration was measured by weighing the potted seedlings. After the last watering, changes in transpiration rate during soil drying were monitored intensely. The dry treatment restricted aboveground growth but increased biomass allocation to the roots in both species, resulting in no significant treatment difference in whole-plant biomass production. The species showed similar responses in relative growth rate (RGR), net assimilation rate (NAR) and shoot mass ratio (SMR) to the dry treatment. Although NAR did not change significantly in either C. japonica or C. obtusa as the soil dried, the two species responded differently to the dry treatment in terms of mean transpiration rate (E) and water-use efficiency (WUE), which are parameters that relate to NAR. In the dry treatment, both E and WUE of C. japonica were stable, whereas in C. obtusa, E decreased and WUE increased (E and WUE counterbalanced to maintain a constant NAR). Transpiration rates were lower in C. obtusa seedlings than in C. japonica seedlings, even in well-watered conditions. During soil drying, the transpiration rate decreased after Theta reached about 0.38 (-0.003 MPa) in C. obtusa and 0.32 (-0.028 MPa) in C. japonica. We conclude that C. obtusa has more water-saving characteristics than C. japonica, particularly when water supply is limited.     

Changes in whole-tree water relations during ontogeny of Pinus flexilis and Pinus ponderosa in a high-elevation meadow

We measured sap flux in Pinus ponderosa Laws. and Pinus flexilis James trees in a high-elevation meadow in northern Arizona that has been invaded by conifers over the last 150 years. Sap flux and environmental data were collected from July 1 to September 1, 2000, and used to estimate leaf specific transpiration rate (El), canopy conductance (Gc) and whole-plant hydraulic conductance (Kh). Leaf area to sapwood area ratio (LA/SA) increased with increasing tree size in P. flexilis, but decreased with increasing tree size in P. ponderosa. Both Gc and Kh decreased with increasing tree size in P. flexilis, and showed no clear trends with tree size in P. ponderosa. For both species, Gc was lower in the summer dry season than in the summer rainy season, but El did not change between wet and dry summer seasons. Midday water potential (Psi(mid)) did not change across seasons for either species, whereas predawn water potential (Psi(pre)) tracked variation in soil water content across seasons. Pinus flexilis showed greater stomatal response to vapor pressure deficit (VPD) and maintained higher Psi(mid) than P. ponderosa. Both species showed greater sensitivity to VPD at high photosynthetically active radiation (PAR; > 2500 micromol m-2 s-1) than at low PAR (< 2500 micromol m-2 s-1). We conclude that the direction of change in Gc and Kh with increasing tree size differed between co-occurring Pinus species, and was influenced by changes in LA/SA. Whole-tree water use and El were similar between wet and dry summer seasons, possibly because of tight stomatal control over water loss.     

Impact of soil drought on sap flow and water status of evergreen trees in a tropical monsoon forest in northern Thailand

Hill evergreen forest is the dominant vegetation type in northern Thailand. In this region, there is higher atmospheric evaporative demand and lower soil moisture during the 5- to 7-month dry season than in the rainy season under influences from Asian monsoons. In an earlier study we revealed that canopy-scale transpiration is actively maintained even during the latter part of the dry season in hill evergreen forest. However, the impact of soil drought on tree water use was not investigated. To clarify the ecohydrological processes at this site, we used individual tree-scale measurements during a 2-year period to base our examination of whether limited water use in individual trees is caused by soil drought in the latter part of the dry season. Sap flow and water potential measurements were conducted in four evergreen trees, two large emergent trees 29.8 and 25.4 m high, and two smaller understory trees 4.8 and 1.4 m high.The amount of rainfall preceding the late dry season of 2004 was significantly less than that preceding the late dry season of 2003. Although a distinct decrease in sap-flow velocities in individual trees due to soil water stress was not found in the late dry season of 2003, it did become comparatively apparent in the late dry season of 2004; ranging from 10 to 40% for a given atmospheric evaporative demand. Furthermore, the reductions in sap-flow velocities and predawn stem-water potential were most significant in the smallest tree. The recovery of sap-flow velocities and water potential in the smallest tree after irrigation confirmed that the reductions in sap-flow velocity and predawn stem-water potential in the smallest tree were caused by soil drought. These results suggest that shallower roots could be reason for the significant decrease in water use in the smallest trees. The deeper roots of larger trees could be the reason for the reduced impact of soil drought on water use in larger trees, and canopy-scale transpiration might be maintained by larger trees, even in an unusually severe drought. These possibilities provide a new insight for management of evergreen forests under Asian monsoon influences.     

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.     

Hydraulic redistribution of soil water during summer drought in two contrasting Pacific Northwest coniferous forests

The magnitude of hydraulic redistribution of soil water by roots and its impact on soil water balance were estimated by monitoring time courses of soil water status at multiple depths and root sap flow under drought conditions in a dry ponderosa pine (Pinus ponderosa Dougl. ex Laws) ecosystem and in a moist Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) ecosystem. The fate of deuterated water applied to small plots to create a strong horizontal soil water potential gradient was also monitored to assess the potential for horizontal redistribution of water and utilization of redistributed water by co-occurring shallow-rooted plants. In a 20-year-old Douglas-fir stand, approximately 28% of the water removed daily from the upper 2 m of soil was replaced by nocturnal hydraulic redistribution during late August. In an old-growth ponderosa pine stand, approximately 35% of the total daily water utilization from the upper 2 m of soil appeared to be replaced by hydraulic redistribution during July and August. By late September, hydraulic redistribution in the ponderosa pine stand was no longer apparent, even though total water use from the upper 2 m of soil was nearly identical to that observed earlier. Based on these results, hydraulic redistribution would allow 21 and 16 additional days of stored water to remain in the upper soil horizons in the ponderosa pine and Douglas-fir stands, respectively, after a 60-day drought. At both sites, localized applications of deuterated water induced strong reversal of root sap flow and caused soil water content to cease declining or even temporarily increase at locations too distant from the site of water application to have been influenced by movement of water through the soil without facilitation by roots. Xylem water deuterium values of ponderosa pine seedlings suggested utilization of redistributed water. Therefore, hydraulic redistribution may enhance seedling survival and maintain overstory transpiration during summer drought. These first approximations of the extent of hydraulic redistribution in these ecosystems suggest that it is likely to be an important process in both wet and dry forests of the Pacific Northwest.     

Growth and nitrogen status of young walnuts as affected by intercropped legumes in a Mediterranean climate

Walnut trees (Juglans nigra L. and Juglans nigra × regia NG23) were intercropped with alfalfa (Medicago sativa L.) or sainfoin (Onobrychis sativa L.) forage legumes compared to a grass (Festuca arundinacea Schr., fescue), or to spontaneous weeds as a control in two Mediterranean sites (Castries and Notre-Dame de Londres) near Montpellier (France). Tree growth, soil water depletion and nitrogen content of the tree leaves were monitored to assess the impact of both water competition and possible facilitation resulting from fixed nitrogen transfer from the leguminous crops to the trees. At Castries, where alfalfa and fescue were compared, they were found to have the same impact on tree growth. At Notre-Dame de Londres where sainfoin was compared with a spontaneous grassing treatment, the sainfoin crop was more competitive to the walnuts. The nitrogen content of walnut leaves was enhanced when intercropped with nitrogen fixing species at both sites. Perennial leguminous intercrops were more competitive for soil water resources than fescue or weeds, but in the long term this may be compensated by the improved nitrogen status of the trees. This compensating effect was observed during a rather rainy year. The overall impact of leguminous intercrops on tree growth may depend on the frequency of dry (competitive) and wet (non-competitive) years. This revised version was published online in June 2006 with corrections to the Cover Date.     

The fate of hydraulically redistributed water in a semi-arid zone eucalyptus species

Although hydraulic redistribution has been observed for a range of tree species, including Eucalyptus kochii subsp. borealis (C. Gardner) D. Nicolle, there is limited direct evidence that water taken up by deep roots in moist soil is in fact exuded by shallow roots in dry soil. This paper reports an experiment designed to test this hypothesis. Water enriched with deuterium was added to the groundwater via a slotted tube at 4.5 m depth below 5-year-old E. kochii subsp. borealis trees. Nocturnal sap flow increased markedly immediately after deep irrigation, indicating that the trees were using water from this depth. Two weeks later, samples of surface soil and xylem water were found to contain levels of deuterium up to 30% higher than soils and xylem water from a control plot upslope of the main treatment plot. This is strong evidence that trees used groundwater and that efflux of important amounts of hydraulically redistributed water occurred via the roots of E. kochii subsp. borealis.     

Persisting soil drought reduces leaf specific conductivity in Scots pine (Pinus sylvestris) and pubescent oak (Quercus pubescens)

Leaf specific conductivity (LSC; the ratio of stem conductivity (K(P)) to leaf area (A(L))), a measure of the hydraulic capacity of the stem to supply leaves with water, varies with soil water content. Empirical evidence for LSC responses to drought is ambiguous, because previously published results were subject to many confounding factors. We tested how LSC of similar-sized trees of the same population, under similar climatic conditions, responds to persistently wet or dry soil. Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) trees were compared between a dry site and a wet site in the Valais, an inner alpine valley in Switzerland. Soil water strongly influenced A(L) and K(P) and the plant components affecting K(P), such as conduit radius, conduit density and functional sapwood area. Trees at the dry site had lower LSC than trees with the same stem diameter at the wet site. Low LSC in trees at the dry site was associated with a smaller functional sapwood area and narrower conduits, resulting in a stronger reduction in K(P) than in A(L). These observations support the hypothesis that trees maintain a homeostatic water pressure gradient. An alternative hypothesis is that relatively high investments in leaves compared with sapwood contribute to carbon gain over an entire season by enabling rapid whole-plant photosynthesis during periods of high water availability (e.g., in spring, after rain events and during morning hours when leaf-to-air vapor pressure deficit is small). Dynamic data and a hydraulic plant growth model are needed to test how investments in leaves versus sapwood and roots contribute to transpiration and to maximizing carbon gain throughout entire growth seasons.     

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.     

Differential summer water use by Pinus edulis and Juniperus osteosperma reflects contrasting hydraulic characteristics

Previous studies of pinyon-juniper woodlands show that Pinus edulis Engelm. makes better use of soil water from summer precipitation pulses than does co-occurring Juniperus osteosperma (Torr.) Little. To investigate the basis of this difference, we examined seasonal variation in cavitation and hydraulic conductance. Pinus edulis remained isohydric over the growing season. Minimum water potentials never fell below -2.3 MPa, and the extent of xylem cavitation remained near constant during the dry season. In contrast, J. osteosperma was anisohydric, reaching water potentials as low as -6.9 MPa, and experiencing progressively greater xylem cavitation as the dry season progressed despite having more cavitation-resistant xylem than P. edulis. We conducted an irrigation experiment to observe the responses of the study species to a summer pulse of water. Although sap flow increased in both species in response to the 25-mm irrigation pulse, only J. osteosperma responded to the 10-mm pulse. This was inconsistent with the response of P. edulis to light rain events and may have been due to a difference in the distribution of irrigation water and rain water between the under- and between-canopy areas. Whole-plant conductance increased following the 25-mm irrigation in P. edulis but remained constant in J. osteosperma. We hypothesized that this difference was caused, in part, by differential refilling of embolized xylem. Area specific hydraulic conductivity was 66% higher in roots of irrigated P. edulis trees relative to roots of control trees 3 days after the 25-mm irrigation (t = 2.14, P = 0.02, df = 16). There was no change in hydraulic conductivity of the roots of J. osteosperma or in the stems of either species. Our results indicate that the response to an irrigation pulse in P. edulis depended on cavitation avoidance in stems and the reversal of cavitation in roots, resulting in increased whole-plant conductance and water uptake. In contrast, J. osteosperma failed to exploit light summer rain events but was able to extract deep soil water at low water potentials.