Regulation of transpirational water loss in Quercus suber trees in a Mediterranean-type ecosystem

Sap flux density in branches, leaf transpiration, stomatal conductance and leaf water potentials were measured in 16-year-old Quercus suber L. trees growing in a plantation in southern Portugal to understand how evergreen Mediterranean trees regulate water loss during summer drought. Leaf specific hydraulic conductance and leaf gas exchange were monitored during the progressive summer drought to establish how changes along the hydraulic pathway influence shoot responses. As soil water became limiting, leaf water potential, stomatal conductance and leaf transpiration declined significantly. Predawn leaf water potential reflected soil water potential measured at 1-m depth in the rhizospheres of most trees. The lowest predawn leaf water potential recorded during this period was -1.8 MPa. Mean maximum stomatal conductance declined from 300 to 50 mmol m(-2) s(-1), reducing transpiration from 6 to 2 mmol m(-2) s(-1). Changes in leaf gas exchange were attributed to reduced soil water availability, increased resistances along the hydraulic pathway and, hence, reduced leaf water supply. There was a strong coupling between changes in soil water content and stomatal conductance as well as between stomatal conductance and leaf specific hydraulic conductance. Despite significant seasonal differences among trees in predawn leaf water potential, stomatal conductance, leaf transpiration and leaf specific hydraulic conductance, there were no differences in midday leaf water potentials. The strong regulation of changes in leaf water potential in Q. suber both diurnally and seasonally is achieved through stomatal closure, which is sensitive to changes in both liquid and vapor phase conductance. This sensitivity allows for optimization of carbon and water resource use without compromising the root-shoot hydraulic link.     

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.     

Chronic cork oak decline and water status: new insights

Chronic decline and Sudden death are two syndromes of cork oak (Quercus suber) dieback. Mortality is associated with water stress, but underlying physiological mechanisms are poorly understood. Here, we investigated the physiological performance of declining trees during the summer drought. Leaf water potential, gas-exchange, fluorescence of photosystem II and leaf and root starch concentration were compared in healthy (asymptomatic) and declining trees. Low annual cork increment in declining trees indicated tree decline for several years. All trees showed similar water status in spring. In summer, declining trees showed lower predawn leaf water potential (?2.0 vs. ?0.8 MPa), but unexpectedly higher midday leaf water potential than healthy trees (?2.8 vs. ?3.3 MPa). The higher midday water potential was linked to by means of strongly reduced stomatal conductance and, consequently, transpiration. This study is pioneer showing that declining trees had high midday water potential. A tendency for lower sap flow driving force (the difference between predawn and midday water potential) in declining trees was also associated with reduced photosynthesis, suggesting that chronic dieback may be associated with low carbon uptake. However, starch in roots and leaves was very low and not correlated to the health status of trees. Declining trees showed lower water-use efficiency and non-photochemical quenching in summer, indicating less resistance to drought. Contrarily to chronic decline, one tree that underwent sudden death presented predawn leaf water potential below the cavitation threshold.     

Sap flow of three co-occurring Mediterranean woody species under varying atmospheric and soil water conditions

We studied the seasonal patterns of water use in three woody species co-occurring in a holm oak forest in northeastern Spain. The three species studied, Quercus ilex L., Phillyrea latifolia L. and Arbutus unedo L., constitute more than 99% of the total basal area of the forest. The study period included the dry seasons of 1999 and 2000. Water use was estimated with Granier-type sap flux sensors. Standard meteorological variables, soil water content and leaf water potentials were also monitored. All monitored individuals reduced leaf-related sap flow (Q(l)) during the summer, concurrent with an increase in soil moisture deficit (SMD). Despite similar maximum Q(l) between species, the decline in Q(l) with increasing SMD was species-dependent. The average reduction in Q(l) between early summer and the peak of the drought was 74% for A. unedo (n = 3), 58% for P. latifolia (n = 3) and 87% for Q. ilex (n = 1). The relationship between canopy stomatal conductance (G(s)) and vapor pressure deficit (D) changed during the course of the drought, with progressively lower G(s) for any given D. Summertime reductions of Q(l) and G(s) were associated with between-species differences in vulnerability to xylem embolism, and with the corresponding degree of native embolism (lowest in P. latifolia and highest in Q. ilex). Our results, combined with previous studies in the same area, outlined differences among the species studied in manner of responding to water shortage, with P. latifolia able to maintain water transport at much lower water potentials than the other two species. In an accompanying experiment, A. unedo responded to an experimental reduction in water availability by reducing Q(l) during the summer. This species also modified its water use between years according to the different seasonal patterns of precipitation. These results are discussed in relation to the possible impacts that climate change will have on Q. ilex-dominated forests.     

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.     

Water stress responses of seedlings of four Mediterranean oak species

Effects of water stress on phenology, growth, stomatal activity and water status were assessed from April to November 1996 in 2-year-old seedlings of Quercus frainetto Ten. (Quercus conferta Kit.), Quercus pubescens Willd., Quercus macrolepis Kotschy (Quercus aegilops auct.) and Quercus ilex L. growing in containers in northern Greece. All four species developed more than 50% of their total leaf area before the beginning of June--an adaptation to arid climates. Well-irrigated plants tended to develop greater individual leaf area, number of leaves per plant, total plant leaf area, height and root:shoot ratios than water-stressed plants, but the difference between treatments was not significant for any parameter in any species. Quercus macrolepis appeared to be the most drought-tolerant of the four species. It maintained the highest number of leaves of the smallest size and increased the proportion of fine roots during drought. In all species, drought caused significant decreases in stomatal conductance and predawn and midday water potentials from mid-July until the end of August, when the lowest soil water content and highest mean daily air temperatures and midday leaf temperatures occurred; however, the responses were species-specific. Among the four species, Quercus macrolepis sustained the highest stomatal conductance despite very low water potentials, thus overcoming drought by means of desiccation tolerance. Quercus ilex decreased stomatal conductance even before severe water stress occurred, thereby avoiding desication during drought. Quercus pubescens had the highest water potential despite a high stomatal conductance, indicating that its leaf water status was independent of stomatal activity. Quercus frainetto was the least drought-resistant of the four species. During drought it developed very low water potentials despite markedly reduced stomatal aperture.     

Differences in leaf gas exchange and water relations among species and tree sizes in an Arizona pine-oak forest

We compared leaf gas exchange and water potential among the dominant tree species and major size classes of trees in an upland, pine-oak forest in northern Arizona. The study included old-growth Gambel oak (Quercus gambelii Nutt.), and sapling, pole, and old-growth ponderosa pines (Pinus ponderosa var. scopulorum Dougl. ex Laws.). Old-growth oak had higher predawn leaf water potential (Psi(leaf)) than old-growth pine, indicating greater avoidance of soil water stress by oak. Old-growth oak had higher stomatal conductance (G(w)), net photosynthetic rate (P(n)), and leaf nitrogen concentration, and lower daytime Psi(leaf) than old-growth pine. Stomatal closure started at a daytime Psi(leaf) of about -1.9 MPa for pine, whereas old-growth oak showed no obvious reduction in G(w) at Psi(leaf) values greater than -2.5 MPa. In ponderosa pine, P(n) and G(w) were highly sensitive to seasonal and diurnal variations in vapor pressure deficit (VPD), with similar sensitivity for sapling, pole, and old-growth trees. In contrast, P(n) and G(w) were less sensitive to VPD in Gambel oak than in ponderosa pine, suggesting greater tolerance of oak to atmospheric water stress. Compared with sapling pine, old-growth pine had lower morning and afternoon P(n) and G(w), predawn Psi(leaf), daytime Psi(leaf), and soil-to-leaf hydraulic conductance (K(l)), and higher foliar nitrogen concentration. Pole pine values were intermediate between sapling and old-growth pine values for morning G(w) and daytime Psi(leaf), similar to sapling pine for predawn Psi(leaf), and similar to old-growth pine for morning and afternoon P(n), afternoon G(w), K(l), and foliar nitrogen concentration. For the pines, low predawn Psi(leaf), daytime Psi(leaf), and K(l) were associated with low P(n) and G(w). Our data suggest that hydraulic limitations are important in reducing P(n) in old-growth ponderosa pine in northern Arizona, and indicate greater avoidance of soil water stress and greater tolerance of atmospheric water stress by old-growth Gambel oak than by old-growth ponderosa pine.     

Stomatal limitation to CO2 assimilation and down-regulation of photosynthesis in Quercus ilex resprouts in response to slowly imposed drought

Holm oak (Quercus ilex L.) is native to hot, dry Mediterranean forests where limited water availability often reduces photosynthesis in many species, and forest fires are frequent. Holm oaks resprout after a disturbance, with improved photosynthetic activity and water relations compared with unburned plants. To better understand the role of water availability in this improvement, watering was withheld from container-grown plants, either intact (controls) or resprouts after excision of the shoot, to gradually obtain a wide range of soil water availabilities. At high water availability, gas exchange rates did not differ between controls and resprouts. At moderate soil dryness, net photosynthesis of control plants decreased as a result of increased stomatal limitation, whereas gas exchange rates of resprouts, which had higher midday and predawn leaf water potentials, were unchanged. Under severe drought, resprouts showed a less marked decline in gas exchange than controls and maintained photosystem II integrity, as indicated by chlorophyll fluorescence measurements. Photosynthesis was down-regulated in both plant types in response to reduced CO2 availability caused by high stomatal limitation. Lower non-stomatal limitations in resprouts than in control plants, as evidenced by higher carboxylation velocity and the capacity for ribulose-1,5-bisphosphate regeneration, conferred greater drought resistance under external constraints similar to summer conditions at midday.     

Response of Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) to soil and atmospheric water deficits under Mediterranean mountain climate

The physiological responses to water deficits of Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) were studied under Mediterranean mountain climate. Minimum leaf water potentials were ?3.2 MPa for oak and ?2.1 MPa for pine, with higher predawn values for pubescent oak. Relative sap flow declined in both species when vapour pressure deficit (D) went above ca. 1.2 kPa, but stomatal control was stronger for pine during the 2003 summer drought. P. sylvestris plant hydraulic conductance on a half-total leaf area basis (k _L,s?1) was 1.2–2.6 times higher than the values shown by Q. pubescens, and it showed a considerably steeper decrease during summer. Leaf-level gas exchange was positively related to k _L,s?1 in both species. Scots pine was more vulnerable to xylem embolism and closed stomata to prevent substantial conductivity losses. The results of this study confirm that pubescent oak is more resistant to extreme drought events.     

Summer field performance of <Emphasis Type="Italic">Quercus petraea</Emphasis> (Matt.) Liebl and <Emphasis Type="Italic">Quercus pyrenaica</Emphasis> Willd seedlings,planted in three sites with contrasting canopy cover

In 2000, one-year-old seedlings of pyrenean oak (Quercus pyrenaica Willd.) and sessile oak (Quercus petraea [Matt.] Liebl) were planted in a thinned and an unthinned plot in a pinewood (Pinus sylvestris), and in a nearby clearing. In summer 2002 and 2003, water relations and gas exchange parameters were measured to address the impact of drought on the seedlings. Chlorophyll a fluorescence was also measured to explore leaf photochemistry and a possible non-stomatal limitation to photosynthesis (A). Reduction in stomatal conductance (g) in response to the decrease of predawn water potential (Ψ_pd) resulted the main cause affecting net carbon uptake. Water potential at midday (Ψ_md) was similar in both species but Quercus petraea was more sensitive to soil water deployment occurred along summer, showing slightly lower Ψ_pd because worse recover of water potential during night. Rate of photosynthesis was higher in Q.␣pyrenaica probably in relation to its greater leaf mass per area (LMA) and nitrogen content per leaf area (N_a). Mortality was highest in the clearing and lowest in the thinned pinewood. Throughout the summer, soil moisture was higher in the thinned area, possibly because of the reduction in tree transpiring surface and interception of rainfall. Accordingly, Ψ_pd of both species was higher in the thinned site.     

Quercus species differ in water and nutrient characteristics in a resource-limited fall-line sandhill habitat

We compared co-occurring mature Quercus laevis Walt. (turkey oak), Q. margaretta Ashe (sand post oak) and Q. incana Bartr. (bluejack oak) trees growing in resource-limited sandhill habitats of the southeastern United States for water and nutrient characteristics. The Quercus spp. differed in their distribution along soil water and nutrient gradients, and in their access to and use of water, even though the study year was wetter than average with no mid-season drought. Quercus laevis had the greatest access to soil water (least negative pre-dawn water potential, psi(pd)) and the most conservative water-use strategy based on its relatively low stomatal conductance (g(s)), high instantaneous water-use efficiency (WUE), least negative midday water potential (psy(md)) and high leaf specific hydraulic conductance (K(L)). Quercus margaretta had the least conservative water-use characteristics, exhibiting relatively high g(s), low instantaneous WUE, most negative psi(md), and low K(L). Quercus margaretta also had a low photosynthetic nitrogen-use efficiency (PNUE), but a high leaf phosphorus concentration. Quercus incana had the poorest access to soil water, but intermediate water-use characteristics and leaf nutrient characteristics more similar to those of Q. laevis. There were no species differences for photosynthesis (A), leaf nitrogen on an area basis, or seasonally integrated WUE (delta13C). Both A and g(s) were positively correlated for each species, but A and g(s) were generally not correlated with psi(pd), psi(md) or delta psi(pd-md). Although we found differences in resource use and resource status among these sandhill Quercus spp., the results are consistent with the interpretation that they are generally drought avoiders. Quercus laevis may have an advantage on xeric ridges because of its greater ability to access soil water and use it more conservatively compared with the other Quercus spp.     

Atmospheric and soil drought reduce nocturnal conductance in live oaks

Nocturnal and daytime whole-canopy transpiration rate (E) and conductance (g = E/VPD, where VPD is leaf to air vapor pressure difference) were assessed gravimetrically in drought-treated and well-watered 3-year-old saplings of live oak species (Quercus series Virentes Nixon) from the southeastern USA (Quercus virginiana Mill.) and Central America (Q. oleoides Cham. and Schlecter). Our objectives were to: (1) quantify nocturnal and daytime E and g in a controlled environment; (2) determine the impact of severe drought on nocturnal E and g; and (3) examine whether unavoidable water loss through the epidermis could account for nocturnal water loss. We calculated daytime E during peak daylight hours (between 0930 and 1330 h) and nocturnal E during complete darkness (between 2200 and 0500 h). In addition to reducing E and g during the daytime, drought-treated plants reduced nocturnal E and g on a whole-canopy basis by 62-64% and 59-61%, respectively, and on a leaf-level basis by 27-28% and 19-26%, respectively. In well-watered plants, nocturnal g declined with increasing VPD, providing evidence for stomatal regulation of nocturnal transpiration. In drought-treated plants, g was low and there was no relationship between nocturnal g and VPD, indicating that water loss could not be reduced further through stomatal regulation. Both daytime and nocturnal g declined curvilinearly with predawn water potential for all plants, but nocturnal g was unrelated to predawn water potentials below -1 MPa. The reductions in daytime and nocturnal E and g during drought were associated with decreases in whole-plant and leaf hydraulic conductances. Observed nocturnal g was within the same range as epidermal conductance for oak species determined in previous studies under a range of conditions. Nocturnal E rose from 6-8% of daytime E for well watered plants to 19-20% of daytime E for drought-treated plants. These results indicate that, during drought, saplings of live oak species reduce g to a minimum through stomatal closure, and experience unavoidable water loss through the epidermis.     

Responses of deciduous forest trees to severe drought in Central Europe

In 2003, Central Europe experienced the warmest summer on record combined with unusually low precipitation. We studied plant water relations and phenology in a 100-year- old mixed deciduous forest on a slope (no ground water table) near Basel using the Swiss Canopy Crane (SCC). The drought lasted from early June to mid September. We studied five deciduous tree species; half of the individuals were exposed to elevated CO(2) concentration ([CO(2)]) (530 ppm) using a free-air, atmospheric CO(2)-enrichment system. In late July, after the first eight weeks of drought, mean predawn leaf water potential about 30 m above ground was -0.9 MPa across all trees, dropping to a mean of -1.5 MPa in mid-August when the top 1 m of the soil profile had no plant accessible moisture. Mean stomatal conductance and rates of maximum net photosynthesis decreased considerably in mid-August across all species. However, daily peak values of sap flow remained surprisingly constant over the whole period in Quercus petraea (Matt.) Liebl., and decreased to only about half of the early summer maxima in Fagus sylvatica L. and Carpinus betulus L. (stomatal down- regulation of flux). Although we detected no differences in most parameters between CO(2)-treated and control trees, predawn leaf water potential tended to be less negative in trees exposed to elevated [CO(2)]. Leaf longevity was greater in 2003 compared with the previous years, but the seasonal increase in stem basal area reached only about 75% of that in previous years. Our data suggest that the investigated tree species, particularly Q. petraea, did not experience severe water stress. However, an increased frequency of such exceptionally dry summers may have a more serious impact than a single event and would give Q. petraea a competitive advantage in the long run.     

Leaf traits and tree rings suggest different water-use and carbon assimilation strategies by two co-occurring Quercus species in a Mediterranean mixed-forest stand in Tuscany, Italy

We compared the water-use characteristics of co-occurring mature Quercus cerris L. and Quercus pubescens Willd. trees growing in resource-limited (mainly water) hilly habitats in Tuscany, Italy. The species differed in their distribution along soil water gradients and in their access to, and use of, water, even though the study year was wetter than average, though with a summer drought. Compared with Q. cerris, Q. pubescens had greater access to soil water (less negative predawn water potentials) and a more conservative water-use strategy based on its relatively low stomatal conductance, high instantaneous water-use efficiency, less negative midday water potential and high soil-to-leaf hydraulic conductance. Quercus cerris had less conservative water-use characteristics than Q. pubescens, exhibiting relatively high stomatal conductance, low instantaneous water-use efficiency, more negative midday water potentials and low soil-to-leaf hydraulic conductance; however, Q. cerris had higher photosynthetic rates than Q. pubescens. Photosynthesis and stomatal conductance were positively correlated in both species. Although a strong correlation between ring widths and precipitation patterns was not found, some dry periods influenced ring-width growth. Quercus pubescens has always grown faster than Q. cerris, probably because of more efficient water use, although stand dynamics (driven by exogenous disturbance factors, including coppicing, browsing and competition) cannot be excluded. Ring-width variability, as well as tree-ring growth in dry years, which should be unaffected by stand dynamics, were higher in Q. pubescens than in Q. cerris. Moreover, Q. pubescens recovered completely after the drought in the seventies, even showing higher tree-ring growth than in the recent past, whereas Q. cerris showed a minor growth decline followed by a recovery to values comparable with those observed before the 1970s drought. Beginning in the early eighties, tree-ring growth decreased in both species, though Q. pubescens showed consistently higher values than Q. cerris. These differences can be explained by differences in water-use efficiency. Despite differences between the species in water use and water status, the results are consistent with the interpretation that both are drought tolerant, but that Q. pubescens is at an advantage on xeric ridges because of its greater ability to access soil water and use it more conservatively compared with Q. cerris.     

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.     

Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

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.     

Response of Quercus pyrenaica (melojo oak) to soil water deficit: a case study in Spain

In the present study, carried out from 2004 to 2006, leaf and stem water potential and stem water content were measured in Quercus pyrenaica Willd. individuals in an experimental forested catchment located in Central Western Spain under Mediterranean subhumid conditions. These indicators of tree water status were compared with soil moisture contents measured in the same area from 0 to 1 m depth and from 0 to 2.5 m depth during the last year of the study. The objectives were to clarify the seasonal and year-to-year variations in tree water status, to examine applicability of stem water content as useful water stress indicator and to discuss how deep soil water and root uptake contribute to survival during the long dry summer. Seasonal variations in the tree variables measured revealed a typical pattern, with maximum values at the end of spring followed by a progressive decline during the summer drought in response to the decrease in soil water content (almost exhausted at 0–100 cm depth). The relatively high values and the non-significant variation in predawn leaf water potentials (except for 2005, which was exceptionally dry) indicate that no clear water stress situations occurred. This may be explained in terms of a progressive absorption of water from the deeper layers. The results also suggest that the stem water content is a more sensitive indicator of long-term water limitation than the other variables measured.     

Effects of root damage associated with Phytophthora cinnamomi on water relations,biomass accumulation,mineral nutrition and vulnerability to water deficit of five oak and chestnut species

The effects of root damage associated with Phytophthora cinnamomi on water relations, biomass accumulation, mineral nutrition and vulnerability to water deficit were investigated in pedunculate oak (Quercus robur), red oak (Quercus rubra) and holm oak (Quercus ilex) saplings over two years. Comparison was made with sweet chestnut (Castanea sativa), a susceptible species to infection by P. cinnamomi, and with a resistant hybrid chestnut (Castanea crenata × C. sativa). Trees were inoculated in 1998 and were subjected to water shortage in 1999. All inoculated sweet chestnuts died before the application of water shortage. Hybrid chestnut, pedunculate oak and red oak displayed low root susceptibility to P. cinnamomi. In these species, water relations, aerial growth and mineral nutrition were slightly affected by inoculation. By contrast, holm oak was the most susceptible oak species to P. cinnamomi as inoculated well‐watered trees displayed the highest root loss (67%) and a 10% mortality. Root loss was associated with a decrease in predawn leaf water potential, a 61% reduction in stomatal conductance, a 55% reduction in aerial biomass, a decrease in leaf carbon isotope discrimination and reduced leaf N and P contents in comparison with controls. In hybrid chestnut and pedunculate oak, water shortage resulted in a similar decrease of predawn leaf water potential, stomatal conductance and aerial biomass in inoculated and non‐inoculated trees. In red and holm oaks, soil volumetric water content of inoculated trees subjected to water shortage remained high. The effects observed in those trees were similar to those of inoculated well‐watered trees and were probably the result of root infection only.     

Responses to water stress in two Eucalyptus globulus clones differing in drought tolerance

We evaluated drought resistance mechanisms in a drought-tolerant clone (CN5) and a drought-sensitive clone (ST51) of Eucalyptus globulus Labill. based on the responses to drought of some physiological, biophysical and morphological characteristics of container-grown plants, with particular emphasis on root growth and hydraulic properties. Water loss in excess of that supplied to the containers led to a general decrease in growth and significant reductions in leaf area ratio, specific leaf area and leaf-to-root area ratio. Root hydraulic conductance and leaf-specific hydraulic conductance decreased as water stress became more severe. During the experiment, the drought-resistant CN5 clone maintained higher leaf water status (higher predawn and midday leaf water potentials), sustained a higher growth rate (new leaf area expansion and root growth) and displayed greater carbon allocation to the root system and lower leaf-to-root area ratio than the drought-sensitive ST51 clone. Clone CN5 possessed higher stomatal conductances at moderate stress as well as higher hydraulic conductances than Clone ST51. Differences in the response to drought in root biomass, coupled with changes in hydraulic properties, accounted for the clonal differences in drought tolerance, allowing Clone CN5 to balance transpiration and water absorption during drought treatment and thereby prolong the period of active carbon assimilation.