Fabian G. Scholz

Mechanisms contributing to seasonal homeostasis of minimum leaf water potential and predawn disequilibrium between soil and plant water potential in Neotropical savanna trees

Seasonal regulation of leaf water potential (ΨL) was studied in eight dominant woody savanna species growing in Brazilian savanna (Cerrado) sites that experience a 5-month dry season. Despite marked seasonal variation in precipitation and air saturation deficit (D), seasonal differences in midday minimum ΨL were small in all of the study species. Water use and water status were regulated by a combination of plant physiological and architectural traits. Despite a nearly 3-fold increase in mean D between the wet and dry season, a sharp decline in stomatal conductance with increasing D constrained seasonal variation in minimum ΨL by limiting transpiration per unit leaf area (E). The leaf surface area per unit of sapwood area (LA/SA), a plant architectural index of potential constraints on water supply in relation to transpirational demand, was about 1.5–8 times greater in the wet season compared to the dry season for most of the species. The changes in LA/SA from the wet to the dry season resulted from a reduction in total leaf surface area per plant, which maintained or increased total leaf-specific hydraulic conductance (Gt) during the dry season. The isohydric behavior of Cerrado tree species with respect to minimum ΨL throughout the year thus was the result of strong stomatal control of evaporative losses, a decrease in total leaf surface area per tree during the dry season, an increase in total leaf-specific hydraulic conductance, and a tight coordination between gas and liquid phase conductance. In contrast with the seasonal isohydric behavior of minimum ΨL, predawn ΨL in all species was substantially lower during the dry season compared to the wet season. During the dry season, predawn ΨL was more negative than bulk soil Ψ estimated by extrapolating plots of E versus ΨL to E=0. Predawn disequilibrium between plant and soil Ψ was attributable largely to nocturnal transpiration, which ranged from 15 to 22% of the daily total. High nocturnal water loss may also have prevented internal water storage compartments from being completely refilled at night before the onset of transpiration early in the day.

Stem and leaf hydraulics of congeneric tree species from adjacent tropical savanna and forest ecosystems.

Leaf and stem functional traits related to plant water relations were studied for six congeneric species pairs, each composed of one tree species typical of savanna habitats and another typical of adjacent forest habitats, to determine whether there were intrinsic differences in plant hydraulics between these two functional types. Only individuals growing in savanna habitats were studied. Most stem traits, including wood density, the xylem water potential at 50% loss of hydraulic conductivity, sapwood area specific conductivity, and leaf area specific conductivity did not differ significantly between savanna and forest species. However, maximum leaf hydraulic conductance (Kleaf) and leaf capacitance tended to be higher in savanna species. Predawn leaf water potential and leaf mass per area were also higher in savanna species in all congeneric pairs. Hydraulic vulnerability curves of stems and leaves indicated that leaves were more vulnerable to drought-induced cavitation than terminal branches regardless of genus. The midday Kleaf values estimated from leaf vulnerability curves were very low implying that daily embolism repair may occur in leaves. An electric circuit analog model predicted that, compared to forest species, savanna species took longer for their leaf water potentials to drop from predawn values to values corresponding to 50% loss of Kleaf or to the turgor loss points, suggesting that savanna species were more buffered from changes in leaf water potential. The results of this study suggest that the relative success of savanna over forest species in savanna is related in part to their ability to cope with drought, which is determined more by leaf than by stem hydraulic traits. Variation among genera accounted for a large proportion of the total variance in most traits, which indicates that, despite different selective pressures in savanna and forest habitats, phylogeny has a stronger effect than habitat in determining most hydraulic traits.

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r2=0.71) than to the normalized difference vegetation index (NDVI, r2=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.

Hydraulic differences along the water transport system of South American Nothofagus species: do leaves protect the stem functionality?

Hydraulic traits were studied for six Nothofagus species from South America (Argentina and Chile), and for three of these species two populations were studied. The main goal was to determine if properties of the water conductive pathway in stems and leaves are functionally coordinated and to assess if leaves are more vulnerable to cavitation than stems, consistent with the theory of hydraulic segmentation along the vascular system of trees in ecosystems subject to seasonal drought. Vulnerability to cavitation, hydraulic conductivity of stems and leaves, leaf water potential, wood density and leaf water relations were examined. Large variations in vulnerability to cavitation of stems and leaves were observed across populations and species, but leaves were consistently more vulnerable than stems. Water potential at 50% loss of maximum hydraulic efficiency (P(50)) ranged from -0.94 to -2.44 MPa in leaves and from -2.6 to -5.3 MPa in stems across species and populations. Populations in the driest sites had sapwood and leaves more vulnerable to cavitation than those grown in the wettest sites. Stronger diurnal down-regulation in leaf hydraulic conductance compared with stem hydraulic conductivity apparently has the function to slow down potential water loss in stems and protect stem hydraulics from cavitation. Species-specific differences in wood density and leaf hydraulic conductance (K(Leaf)) were observed. Both traits were functionally related: species with higher wood density had lower K(Leaf). Other stem and leaf hydraulic traits were functionally coordinated, resulting in Nothofagus species with an efficient delivery of water to the leaves. The integrity of the more expensive woody portion of the water transport pathway can thus be maintained at the expense of the replaceable portion (leaves) of the stem-leaf continuum under prolonged drought. Compensatory adjustments between hydraulic traits may help to decrease the rate of embolism formation in the trees more vulnerable to cavitation.

Size-dependent mortality in a Neotropical savanna tree: the role of height-related adjustments in hydraulic architecture and carbon allocation

Size-related changes in hydraulic architecture, carbon allocation and gas exchange of Sclerolobium paniculatum (Leguminosae), a dominant tree species in Neotropical savannas of central Brazil (Cerrado), were investigated to assess their potential role in the dieback of tall individuals. Trees greater than approximately 6-m-tall exhibited more branch damage, larger numbers of dead individuals, higher wood density, greater leaf mass per area, lower leaf area to sapwood area ratio (LA/SA), lower stomatal conductance and lower net CO(2) assimilation than small trees. Stem-specific hydraulic conductivity decreased, while leaf-specific hydraulic conductivity remained nearly constant, with increasing tree size because of lower LA/SA in larger trees. Leaves were substantially more vulnerable to embolism than stems. Large trees had lower maximum leaf hydraulic conductance (K(leaf)) than small trees and all tree sizes exhibited lower K(leaf) at midday than at dawn. These size-related adjustments in hydraulic architecture and carbon allocation apparently incurred a large physiological cost: large trees received a lower return in carbon gain from their investment in stem and leaf biomass compared with small trees. Additionally, large trees may experience more severe water deficits in dry years due to lower capacity for buffering the effects of hydraulic path-length and soil water deficits.

The stem xylem of Patagonian shrubs operates far from the point of catastrophic dysfunction and is additionally protected from drought-induced embolism by leaves and roots.

Hydraulic architecture was studied in shrub species differing in rooting depth in a cold desert in Southern Argentina. All species exhibited strong hydraulic segmentation between leaves, stems and roots with leaves being the most vulnerable part of the hydraulic pathway. Two types of safety margins describing the degree of conservation of the hydraulic integrity were used: the difference between minimum stem or leaf water potential (Ψ) and the Ψ at which stem or leaf hydraulic function was reduced by 50% (Ψ - Ψ50), and the difference between leaf and stem Ψ50. Leaf Ψ50 - stem Ψ50 increased with decreasing rooting depth. Large diurnal decreases in root-specific hydraulic conductivity suggested high root vulnerability to embolism across all species. Although stem Ψ50 became more negative with decreasing species-specific Ψsoil and minimum stem Ψ, leaf Ψ50 was independent of Ψ and minimum leaf Ψ. Species with embolism-resistant stems also had higher maximum stem hydraulic conductivity. Safety margins for stems were >2.1 MPa, whereas those for leaves were negative or only slightly positive. Leaves acted as safety valves to protect the integrity of the upstream hydraulic pathway, whereas embolism in lateral roots may help to decouple portions of the plant from the impact of drier soil layers.

Water relations and hydraulic architecture in Cerrado trees: adjustments to seasonal changes in water availability and evaporative demand

O objetivo deste estudo foi determinar os ajustamentos na morfologia e fisiologia que permitem arvores das savanas neotropicais do Brasil Central (Cerrado) de evitar deficits hidricos e de manter um balanco hidrico interno praticamente constante apesar das variacoes sazonais da precipitacao e no deficit de saturacao do ar (D). A precipitacao na area de estudo e fortemente sazonal, com cerca de cinco meses praticamente sem chuva durante os quais D e duas vezes maior aos valores medidos na epoca chuvosa. Como consequencia da flutuacao sazonal das chuvas e de D, o potencial hidrico do solo muda substancialmente, nos primeiros 100 cm do solo, mas permanece quase constante abaixo de 2 m de profundidade. A arquitetura hidraulica e os parâmetros relacionados a relacoes hidricas das arvores do Cerrado se ajustaram durante a estacao seca para evitar o deficit hidrico crescente e assegurar a homeostase nos valores minimos de potencial hidrico foliar ψL e na perda total diaria de agua pela planta (iso-hidria). O comportamento iso-hidrico das arvores do Cerrado foi o resultado de uma diminuicao da superficie foliar total por arvore, um forte controle estomatico das perdas por evaporacao, um aumento na condutividade hidraulica especifica da folha e na condutância hidraulica foliar e um aumento da quantidade de aguas retirada dos reservatorios internos do caule, durante a estacao seca. A eficiencia no transporte de agua aumentou, nas mesmas proporcoes, nas folhas e nos ramos terminais durante a estacao seca. Todos estes ajustamentos sazonais foram importantes para a manutencao de ψL acima de limiares criticos, com isto contribuindo para uma reducao na formacao de embolismos nos ramos e ajudando a evitar a perda de turgor em tecidos foliares durante a epoca seca. Esses ajustes permitem que os ramos das especies lenhosas do Cerrado operem bem distanciados do ponto de disfuncao catastrofica para a cavitacao, enquanto as folhas operam proximas e sofrem embolismos em uma base diaria, especialmente durante a estacao seca.

Hydraulic redistribution in dwarf Rhizophora mangle trees driven by interstitial soil water salinity gradients: impacts on hydraulic architecture and gas exchange

Rhizophora mangle L. trees of Biscayne National Park (Florida, USA) have two distinct growth forms: tall trees (5-10 m) growing along the coast and dwarf trees (1 m or less) growing in the adjacent inland zone. Sharp decreases in salinity and thus increases in soil water potential from surface soil to about a depth of 1 m were found at the dwarf mangrove site but not at the tall mangrove site. Consistent with our prediction, hydraulic redistribution detected by reverse sap flow in shallow prop roots was observed during nighttime, early morning and late afternoon in dwarf trees, but not in tall trees. In addition, hydraulic redistribution was observed throughout the 24-h period during a low temperature spell. Dwarf trees had significantly lower sapwood-specific hydraulic conductivity, smaller stem vessel diameter, lower leaf area to sapwood area ratio (LA/SA), smaller leaf size and higher leaf mass per area. Leaves of dwarf trees had lower CO(2) assimilation rate and lower stomatal conductance compared to tall trees. Leaf water potentials at midday were more negative in tall trees that are consistent with their substantially higher stomatal conductance and LA/SA. The substantially lower water transport efficiency and the more conservative water use of dwarf trees may be due to a combination of factors such as high salinity in the surface soil, particularly during dry periods, and substantial reverse sap flow in shallow roots that make upper soil layers with high salinity a competing sink of water to the transpiring leaves. There may also be a benefit for the dwarf trees in having hydraulic redistribution because the reverse flow and the release of water to upper soil layers should lead to dilution of the high salinity in the rhizosphere and thus relieve its potential harm to dwarf R. mangle trees.

Why do trees adjust water relations and hydraulic architecture in response to nutrient availability

Fil: Goldstein, Guillermo Hernan. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Oficina de Coordinacion Administrativa Ciudad Universitaria. Instituto de Ecologia, Genetica y Evolucion de Buenos Aires; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Ecologia, Genetica y Evolucion. Laboratorio de Ecologia Funcional; Argentina. University of Miami. Departament of Biology; Estados Unidos

Water storage dynamics in the main stem of subtropical tree species differing in wood density, growth rate and life history traits

Wood biophysical properties and the dynamics of water storage discharge and refilling were studied in the trunk of canopy tree species with diverse life history and functional traits in subtropical forests of northeast Argentina. Multiple techniques assessing capacitance and storage capacity were used simultaneously to improve our understanding of the functional significance of internal water sources in trunks of large trees. Sapwood capacitances of 10 tree species were characterized using pressure-volume relationships of sapwood samples obtained from the trunk. Frequency domain reflectometry was used to continuously monitor the volumetric water content in the main stems. Simultaneous sap flow measurements on branches and at the base of the tree trunk, as well as diurnal variations in trunk contraction and expansion, were used as additional measures of stem water storage use and refilling dynamics. All evidence indicates that tree trunk internal water storage contributes from 6 to 28% of the daily water budget of large trees depending on the species. The contribution of stored water in stems of trees to total daily transpiration was greater for deciduous species, which exhibited higher capacitance and lower sapwood density. A linear relationship across species was observed between wood density and growth rates with the higher wood density species (mostly evergreen) associated with lower growth rates and the lower wood density species (mostly deciduous) associated with higher growth rates. The large sapwood capacitance in deciduous species may help to avoid catastrophic embolism in xylem conduits. This may be a low-cost adaptation to avoid water deficits during peak water use at midday and under temporary drought periods and will contribute to higher growth rates in deciduous tree species compared with evergreen ones. Large capacitance appears to have a central role in the rapid growth patterns of deciduous species facilitating rapid canopy access as these species are less shade tolerant than evergreen species.