Role of flavanols in yellowing beech trees of the Black Forest

Beech leaves were sampled at the end of a prolonged hot dry period at a tree decline site in the Black Forest, Germany to investigate the potential role of flavanols in defense mechanisms against environmental stress. Green and yellowing leaves were harvested from the uppermost canopy of trees that were more than 200 years old and 30 m high. Yellowing leaves had a 7.4-fold higher concentration of total flavanols than green leaves. Green leaves contained flavanol inclusions, but during yellowing the inclusions disintegrated and the cells became filled with flavanols. Abscisic acid (ABA) stimulated the release of flavanols from intravacuolar inclusions of leaf petioles and flower pedicels. In addition, ABA caused flavanols to leach from the trichomes of beech galls. The antioxidative potential of leaf extracts, as estimated by indoleacetic acid (IAA) oxidation, was significantly higher in yellowing leaves than in green leaves. In vitro experiments revealed that (+)-catechin promoted growth of beech tissue.     

Intra-plant variation in cyanogenesis and the continuum of foliar plant defense traits in the rainforest tree Ryparosa kurrangii (Achariaceae)

At the intra-plant level, temporal and spatial variations in plant defense traits can be influenced by resource requirements, defensive priorities and storage opportunities. Across a leaf age gradient, cyanogenic glycoside concentrations in the rainforest understory tree Ryparosa kurrangii B.L. Webber were higher in young expanding leaves than in mature leaves (2.58 and 1.38 mg g(-1), respectively). Moreover, cyanogens, as an effective chemical defense against generalist herbivores, contributed to a defense continuum protecting foliar tissue during leaf development. Chemical (cyanogens and phenolic compounds) and phenological (delayed greening) defense traits protected young leaves, whereas mature leaves were largely protected by physical defense mechanisms (lamina toughness; explained primarily by leaf mass per area). Cyanogen concentration was considerably higher in floral tissue than in foliar tissue and decreased in floral tissue during development. Across contrasting tropical seasons, foliar cyanogenic concentration varied significantly, being highest in the late wet season and lowest during the pre-wet season, the latter coinciding with fruiting and leaf flushing. Cyanogens in R. kurrangii appear to be differentially allocated in a way that maximizes plant fitness but may also act as a store of reduced nitrogen that is remobilized during flowering and leaf flushing.     

Contributions of foliage distribution and leaf functions to light interception, transpiration and photosynthetic capacities in two apple cultivars at branch and tree scales

Both the spatial distribution of leaves and leaf functions affect the light interception, transpiration and photosynthetic capacities of trees, but their relative contributions have rarely been investigated. We assessed these contributions at the branch and tree scales in two apple cultivars (Malus x domestica Borkh. 'Fuji' and 'Braeburn') with contrasting architectures, by estimating their branch and tree capacities and comparing them with outputs from a radiation absorption, transpiration and photosynthesis (RATP) functional-structural plant model (FSPM). The structures of three 8-year-old trees of each cultivar were digitized to obtain 3-D representations of foliage geometry. Within-tree foliage distribution was compared with shoot demography, number of leaves per shoot and mean individual leaf area. We estimated branch and tree light interception from silhouette to total leaf area ratios (STAR), transpiration from sap flux measurements and net photosynthetic rates by the branch bag method. Based on a set of parameters we previously established for both cultivars, the outputs of the RATP model were tested against STAR values, sap fluxes and photosynthetic measurements. The RATP model was then used to virtually switch foliage distribution or leaf functions (stomatal and photosynthetic properties), or both, between cultivars and to evaluate the effects on branch and tree light interception, transpiration and photosynthetic capacities in each cultivar. 'Fuji' trees had a higher proportion of leaf area borne on long shoots, fewer leaves per unit shoot length and a larger individual leaf area than 'Braeburn' trees. This resulted in a lower leaf area density and, consequently, a higher STAR in 'Fuji' than in 'Braeburn' at both branch and tree scales. Transpiration and photosynthetic rates were significantly higher in 'Fuji' than in 'Braeburn'. Branch heterogeneity was greater in 'Braeburn' than in 'Fuji'. An analysis of the virtual switches of foliage distribution or leaf function showed that differences in leaf spatial distribution and functions had additive effects that accounted for the lower transpiration and photosynthetic rates of branches and trees of 'Braeburn' compared with 'Fuji'. Leaf distribution had a more important role at the branch scale than at the tree scale, but the leaf function effect exceeded the leaf distribution effect at both scales. Our study demonstrated the potential of FSPM to disentangle physiological differences between cultivars through in silico scenarios.     

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.     

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

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

Photosynthetic differences between saplings and adult trees: an integration of field results by meta-analysis

Ontogenetic changes in gas exchange parameters provide both insight into mechanisms underlying tree growth patterns, and data necessary to scale environmental impacts on young trees to predict responses of older trees. We present a quantitative review and meta-analysis of field measurements of gas exchange parameters in saplings and mature trees of 35 tree species (seven conifers, seven temperate deciduous trees, and 21 tropical evergreen trees). Data for saplings were obtained in both understory environments and open areas or large gaps. We also present data on ontogenetic changes in photosynthesis for Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg., species of particular interest because of their large maximal heights and long life-spans. Among tree species, there is evidence for both ontogenetic increases and ontogenetic decreases in photosynthetic capacity on a leaf area basis (A(area)). Overall, A(area) is generally higher for upper-canopy leaves of adult trees than for saplings, especially in temperate deciduous trees. However, the pattern for photosynthetic capacity on a leaf mass basis (A(mass)) is the reverse of that observed for A(area). Saplings of both conifers and broad-leaved trees, even when acclimated to low-light conditions, characteristically have a higher A(mass) than adult trees. This pattern is driven largely by an ontogenetic increase in leaf mass per unit area (LMA), as found in 100% of studies reviewed. Data for Pacific Northwest conifers, although including measurements on some of the tallest trees studied, did not differ greatly from patterns found in other tree species. We conclude that ontogenetic changes in LMA are the single most consistent difference between saplings and adult trees, and that changes in LMA and related aspects of leaf morphology may be critical to understanding both variation in gas exchange during tree growth, and stage-dependent responses of trees to environmental change.     

Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine

As forests age, their structure and productivity change, yet in some cases, annual rates of water loss remain unchanged. To identify mechanisms that might explain such observations, and to determine if widely different age classes of forests differ functionally, we examined young (Y, approximately 25 years), mature (M, approximately 90 years) and old (O, approximately 250 years) ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) stands growing in a drought-prone region of central Oregon. Although the stands differed in tree leaf area index (LAIT) (Y = 0.9, M = 2.8, O = 2.1), cumulative tree transpiration measured by sap flow did not differ substantially during the growing season (100-112 mm). Yet when water was readily available, transpiration per unit leaf area of the youngest trees was about three times that of M trees and five times that of O trees. These patterns resulted from a nearly sixfold difference in leaf specific conductance (KL) between the youngest and oldest trees. At the time of maximum transpiration in the Y stand in May-June, gross carbon uptake (gross ecosystem production, GEP) was similar for Y and O stands despite an almost twofold difference in stand leaf area index (LAIS). However, the higher rate of water use by Y trees was not sustainable in the drought-prone environment, and between spring and late summer, KL of Y trees declined fivefold compared with a nearly twofold decline for M trees and a < 30% reduction in O trees. Because the Y stand contained a significant shrub understory and more exposed soil, there was no appreciable difference in mean daily latent energy fluxes between the Y stand and the older stands as measured by the eddy-covariance technique. These patterns resulted in 60 to 85% higher seasonal GEP and 55 to 65% higher water-use efficiency at the M and O stands compared with the Y stand.     

Ontogenetic transition of leaf physiology and anatomy from seedlings to mature trees of a rain forest pioneer tree, Macaranga gigantea

In a field study, we compared anatomy and diurnal gas exchange and chlorophyll fluorescence in sunlit mature leaves of Macaranga gigantea (Reichb. f. and Zoll.) Muell. seedlings, saplings, an adult tree and suckers originating from stumps. We tested the hypothesis that the pattern of resource use shifts across various life stages with ontogenetic changes in leaf anatomy and physiology. Among leaves of different developmental stages, seedling leaves were the smallest and thinnest, whereas adult tree leaves were the largest and thickest, and the air space within the lamina was largest in seedling leaves and smallest in adult tree leaves. Photosynthetic nitrogen-use efficiency (PNUE) was higher in seedling and sapling leaves than in adult tree leaves. Mean PNUE in seedling leaves was 1.6 times that in adult tree leaves. Nevertheless, among the developmental stages, net photosynthetic rate (Pn) per unit leaf area was lowest in seedling leaves because they have the lowest nitrogen (N) content per unit leaf area. In situ water vapor stomatal conductance (g(s) at a given leaf-to-air vapor pressure deficit was highest in sapling leaves, suggesting that they have a high hydraulic efficiency per unit leaf area. Among developmental stages, intrinsic water-use efficiency (Pn/g(s)) and photochemical capacity of photosystem II were lowest in seedling leaves. Sapling leaves had the highest N concentration and Pn per unit dry mass and the highest g(s), indicating that the gradual transition from the seedling stage to the sapling stage is accompanied by an accumulation of N in plant bodies and the development of hydraulic systems to counteract unfavorable environmental stresses. The properties of adult tree leaves (low PNUE, high carbon:N ratio, small and dense cells and thick lamina) indicate that, during the transition from the sapling stage to the adult tree stage, the priority of resource use in leaves gradually shifts from enhancement of photosynthetic performance to defense against herbivory and mechanical damage. Leaf morphology and physiology were coordinated with the differences in resource use at each life stage.     

Differences in leaf phenology between juvenile and adult trees in a temperate deciduous forest

In a deciduous forest, differences in leaf phenology between juvenile and adult trees could result in juvenile trees avoiding canopy shade for part of the growing season. By expanding leaves earlier or initiating senescence later than canopy trees, juvenile trees would have some period in high light and therefore greater potential carbon gain. We observed leaf phenology of 376 individuals of 13 canopy tree species weekly over 3 years in a deciduous forest in east central Illinois, USA. Our objectives were: (1) to quantify for each species the extent of differences in leaf phenology between juvenile and conspecific adult trees; and (2) to determine the extent of phenological differences between juvenile Aesculus glabra Willd. and Acer saccharum Marsh. trees in understory and gap microhabitats. All species displayed phenological differences between life stages. For 10 species, bud break was significantly earlier, by an average of 8 days, for subcanopy individuals than for canopy individuals. In 11 species, completion of leaf expansion was earlier, by an average of 6 days, for subcanopy individuals than for canopy individuals. In contrast, there were no significant differences between life stages for start of senescence in 10 species and completion of leaf drop in nine species. For eight species, leaf longevity was significantly greater for subcanopy individuals than for canopy individuals by an average of 7 days (range = 4-10 days). Leaf phenology of subcanopy individuals of both Aesculus glabra and Acer saccharum responded to gap conditions. Leaf longevity was 11 days less in the understory than in gaps for Aesculus glabra, but 14 days more in the understory than in gaps for Acer saccharum. Therefore, leaf phenology differed broadly both between life stages and within the juvenile life stage in this community. A vertical gradient in temperature sums is the proposed mechanism explaining the patterns. Temperature sums accumulated more rapidly in the sheltered understory than in an open elevated area, similar to the canopy. Early leaf expansion by juvenile trees may result in a period of disproportionately higher carbon gain, similar to gains made during summer months from use of sun flecks.     

Dynamics of nonstructural carbohydrates and biomass yield in a fodder legume tree at different harvest intensities

Tropical tree fodder is harvested by frequent prunings, and resprouting depends on nonstructural carbohydrate reserves in the remaining tree parts. We studied the effects of three pruning intensities (removal of all leaves and branches leaving 1 m of stem once a year (T-12), or every 6 months (T-6), and about 50% pruning every 2 months (P-2)) on regrowth and the dynamics of soluble sugars and starch in the legume tree Gliricidia sepium (Jacq.) Walp. growing under humid tropical conditions in Guadeloupe, Lesser Antilles. Carbohydrates were sampled in roots, stems and branches. Among pruned trees, trees in the T-6 harvest regime had the highest leaf fodder yield (0.73 kg tree(-1) year(-1)). High litter loss reduced leaf yield of T-12 trees, but compared with the other treatments, T-12 trees produced the most branch biomass (3.43 kg tree(-1)). Among treatments, P-2 trees had an intermediate leaf fodder yield and the lowest branch production. Sucrose, glucose and fructose were the most common sugars in all biomass compartments. Mannose, pinitol and an unidentified cyclitol were relatively abundant in branches. Root sugar and starch concentrations were unaffected by harvest regime. There was a significant interactive effect of harvest intensity and regrowth time on stem sugar concentration. Stem starch concentration was highest in T-12 trees. After a year of fodder harvesting, whole-tree reserves of nonstructural carbohydrates were highest in T-12 trees; however, a larger proportion of reserves were located in roots and stems of T-6 and P-2 trees. These reserves, which were not lost in pruning and contributed to regrowth of G. sepium after pruning, may explain the relatively small effects of harvesting regime on soluble sugar and starch concentrations.     

Electric resistance as a measure of tree water status during seasonal drought in a tropical dry forest in Costa Rica

Variation in electric resistance of stem tissues was used to measure differences and changes in water status among trees in a tropical dry forest in Costa Rica during the dry season. For more than 30 tree species, stem water content (SWC), measured as electric resistance between nails driven 20 mm deep into tree trunks, correlated well with wood density, saturation water content, dehydration, measured with the pressure chamber, and tree development during drought. At dry sites, SWC was lowest in hardwood trees (characterized by high wood density) and highest in stem-succulent lightwood trees (characterized by low wood density). Among hardwood trees, SWC varied with soil water availability. During the dry season, SWC declined before leaf shedding and increased during rehydration preceding bud break. The time course of seasonal changes in SWC apparently constitutes an indirect measure of variation in the relative water content of outer stem tissues, which determines development of dry-forest trees during the dry season.     

不同林龄樟子松叶片养分含量及其再吸收效率

树木叶片的养分再吸收效率能够反映树木对养分保存、利用以及对养分贫瘠环境的适应能力。以科尔沁沙地东南缘章古台地区樟子松人工林为研究对象,分析了11、20、29、45年生树木叶片的基本特征、养分含量及其再吸收效率。结果表明:叶片衰老后其质量和面积明显减少;叶片凋落前的平均养分含量没有表现出随樟子松年龄增加而出现有规律的变化;凋落叶片中的N、P、K、Mg含量表现出随年龄增加而增加的趋势,而Ca的趋势与之相反;11年生和20年生的樟子松叶片N、P、K的再吸收效率相似,都显著高于29年生和45年生樟子松(P<0.05),而樟子松叶片对Mg的再吸收效率表现出随年龄增大而显著降低,Ca随叶片的衰老而不断累积,再吸收效率表现为负值,20年生的樟子松叶片Ca再吸收效率最大,11年生和45年生最低。樟子松叶片的N、P、K、Mg养分再吸收效率随年龄增加而降低的趋势表明,随年龄增加樟子松对贫瘠养分生境的适应能力逐渐降低,反映了樟子松养分保存方面的衰退特征。     

Response of giant sequoia canopy foliage to elevated concentrations of atmospheric ozone

We examined the physiological response of foliage in the upper third of the canopy of 125-year-old giant sequoia (Sequoiadendron giganteum Buchholz.) trees to a 61-day exposure to 0.25x, 1x, 2x or 3x ambient ozone concentration. Four branch exposure chambers, one per ozone treatment, were installed on 1-m long secondary branches of each tree at a height of 34 m. No visible symptoms of foliar ozone damage were apparent throughout the 61-day exposure period and none of the ozone treatments affected branch growth. Despite the similarity in ozone concentrations in the branch chambers within a treatment, the trees exhibited different physiological responses to increasing ozone uptake. Differences in diurnal and seasonal patterns of g(s) among the trees led to a 2-fold greater ozone uptake in tree No. 2 compared with trees Nos. 1 and 3. Tree No. 3 had significantly higher CER and g(s) at 0.25x ambient ozone than trees Nos. 1 and 2, and g(s) and CER of tree No. 3 declined with increasing ozone uptake. The y-intercept of the regression for dark respiration versus ozone uptake was significantly lower for tree No. 2 than for trees Nos. 1 and 3. In the 0.25x and 1x ozone treatments, the chlorophyll concentration of current-year foliage of trees Nos. 1 and 2 was significantly higher than that of current-year foliage of tree No. 3. Chlorophyll concentration of current-year foliage on tree No. 1 did not decline with increasing ozone uptake. In all trees, total needle water potential decreased with increasing ozone uptake, but turgor was constant. Although tree No. 2 had the greatest ozone uptake, g(s) was highest and foliar chlorophyll concentration was lowest in tree No. 3 in the 0.25x and 1x ambient atmospheric ozone treatments.     

Tree phenology and water availability in semi-arid agroforestry systems

Four common agroforestry trees, including both exotic and native species, were used to provide a range of leafing phenologies to test the hypothesis that temporal complementarity between trees and crops reduces competition for water in agroforestry systems during the cropping period and improves utilisation of annual rainfall. Species examined included Melia volkensii, which sheds its leaves twice a year, Senna spectabilis and Gliricidia sepium, which shed their leaves once during the long dry season, and the evergreen Croton megalocarpus. Phenological patterns were examined in relation to climatic conditions in the bimodal rainfall regions of Kenya to identify factors which dictate the intensity of competition between trees and crops.

The main differences in leaf cover patterns were between indigenous and exotic tree species. The Central American species, S. spectabilis and G. sepium, shed their foliage during the dry season before the short rains, whereas the native species, M. volkensii and C. megalocarpus, exhibited reduced leaf cover during both dry seasons. C. megalocarpus was the only species to maintain leaf cover throughout the 2-year experimental period. M. volkensii and S. spectabilis exhibited similar leafing phenology, losing almost all leaf cover during the long dry season (July–October) and flushing before the onset of the ensuing rains. S. spectabilis lost few leaves during the short dry season, whereas M. volkensii shed a greater proportion of its foliage before flushing prior to the long rains (March–July). M. volkensii lost much of its leaf cover during the 1997/1998 short rains (October–February), when soil water content was unusually high. Although essentially evergreen, leaf cover in C. megalocarpus decreased during the dry season and increased rapidly during periods of high rainfall. G. sepium exhibited a period of low leaf cover during the long dry season and did not regain full leaf cover until mid-way through the short rains. The mechanisms responsible for these phenological changes and the implications of tree phenology for resource utilisation and competition with crops are discussed.     

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.     

Effects of pruning on the concentration of secondary metabolites in Colophospermum mopane leaves

Colophospermum mopane, commonly known as mopane, produces secondary metabolites during the growing season. However, there is still insufficient knowledge on the quantity of secondary metabolites and the effect of browsers on the concentration of secondary metabolites. A pruning experiment was conducted in the Musina Nature Reserve, Limpopo province, South Africa, to simulate the effect of browsers on the concentration of secondary metabolites in mopane leaves. The twigs from 40 selected experimental mopane trees were pruned back 50 mm from their tips using a hand shear. The total amount of leaf and shoot biomass removed from branches of selected experimental trees was less than 10%. Three independent samples composed of seven mopane leaves per 40 mopane trees were randomly collected from the canopies of experimental and control trees per sampling event. The dried leaves from each three independent samples were then mixed separately and a pooled sample of 10 g per treatment per sample cycle (55 d) was used for determination of total phenols (TP), condensed tannins (CT) and protein-precipitating tannins (PPT). Results showed that <10% pruning does not have an effect on the amount of secondary metabolites in the mopane leaves. The concentration of TP, CT and PPT increased during leaf flush in October and then declined as the leaves matured and aged. It is concluded that the amount of secondary metabolites in mopane leaves is not dependent on <10% pruning, but appeared to be associated with leaf growth stages. The ability of mopane to produce secondary metabolites has implications on the seasonal diet composition and distribution of browsers in mopane woodland.     

Water-use strategies in two co-occurring Mediterranean evergreen oaks: surviving the summer drought

In the Mediterranean evergreen oak woodlands of southern Portugal, the main tree species are Quercus ilex ssp. rotundifolia Lam. (holm oak) and Quercus suber L. (cork oak). We studied a savannah-type woodland where these species coexist, with the aim of better understanding the mechanisms of tree adaptation to seasonal drought. In both species, seasonal variations in transpiration and predawn leaf water potential showed a maximum in spring followed by a decline through the rainless summer and a recovery with autumn rainfall. Although the observed decrease in predawn leaf water potential in summer indicates soil water depletion, trees maintained transpiration rates above 0.7 mm day(-1) during the summer drought. By that time, more than 70% of the transpired water was being taken from groundwater sources. The daily fluctuations in soil water content suggest that some root uptake of groundwater was mediated through the upper soil layers by hydraulic lift. During the dry season, Q. ilex maintained higher predawn leaf water potentials, canopy conductances and transpiration rates than Q. suber. The higher water status of Q. ilex was likely associated with their deeper root systems compared with Q. suber. Whole-tree hydraulic conductance and minimum midday leaf water potential were lower in Q. ilex, indicating that Q. ilex was more tolerant to drought than Q. suber. Overall, Q. ilex seemed to have more effective drought avoidance and drought tolerance mechanisms than Q. suber.     

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.     

Leaf senescence and late-season net photosynthesis of sun and shade leaves of overstory sweetgum (Liquidambar styraciflua) grown in elevated and ambient carbon dioxide concentrations

We examined the effects of elevated CO2 concentration ([CO2]) on leaf demography, late-season photosynthesis and leaf N resorption of overstory sweetgum (Liquidambar styraciflua L.) trees in the Duke Forest Free Air CO2 Enrichment (FACE) experiment. Sun and shade leaves were subdivided into early leaves (formed in the overwintering bud) and late leaves (formed during the growing season). Overall, we found that leaf-level net photosynthetic rates were enhanced by atmospheric CO2 enrichment throughout the season until early November; however, sun leaves showed a greater response to atmospheric CO2 enrichment than shade leaves. Elevated [CO2] did not affect leaf longevity, emergence date or abscission date of sun leaves or shade leaves. Leaf number and leaf area per shoot were unaffected by CO2 treatment. A simple shoot photosynthesis model indicated that elevated [CO2] stimulated photosynthesis by 60% in sun shoots, but by only 3% in shade shoots. Whole-shoot photosynthetic rate was more than 12 times greater in sun shoots than in shade shoots. In senescent leaves, elevated [CO2] did not affect residual leaf nitrogen, and nitrogen resorption was largely unaffected by atmospheric CO2 enrichment, except for a small decrease in shade leaves. Overall, elevated [CO2] had little effect on the number of leaves per shoot at any time during the season and, therefore, did not change seasonal carbon gain by extending or shortening the growing season. Stimulation of carbon gain by atmospheric CO2 enrichment in sweetgum trees growing in the Duke Forest FACE experiment was the result of a strong stimulation of photosynthesis throughout the growing season.     

Effects of leaf age and tree size on stomatal and mesophyll limitations to photosynthesis in mountain beech (Nothofagus solandrii var. cliffortiodes)

Mesophyll conductance, g(m), was estimated from measurements of stomatal conductance to carbon dioxide transfer, g(s), photosynthesis, A, and chlorophyll fluorescence for Year 0 (current-year) and Year 1 (1-year-old) fully sunlit leaves from short (2 m tall, 10-year-old) and tall (15 m tall, 120-year-old) Nothofagus solandrii var. cliffortiodes trees growing in adjacent stands. Rates of photosynthesis at saturating irradiance and ambient CO(2) partial pressure, A(satQ), were 25% lower and maximum rates of carboxylation, V(cmax), were 44% lower in Year 1 leaves compared with Year 0 leaves across both tree sizes. Although g(s) and g(m) were not significantly different between Year 0 and Year 1 leaves and g(s) was not significantly different between tree heights, g(m) was significantly (19%) lower for leaves on tall trees compared with leaves on short trees. Overall, V(cmax) was 60% higher when expressed on the basis of CO(2) partial pressure at the chloroplasts, C(c), compared with V(cmax) on the basis of intercellular CO(2) partial pressure, C(i), but this varied with leaf age and tree size. To interpret the relative stomatal and mesophyll limitations to photosynthesis, we used a model of carbon isotopic composition for whole leaves incorporating g(m) effects to generate a surface of 'operating values' of A over the growing season for all leaf classes. Our analysis showed that A was slightly higher for leaves on short compared with tall trees, but lower g(m) apparently reduced actual A substantially compared with A(satQ). Our findings showed that lower rates of photosynthesis in Year 1 leaves compared with Year 0 leaves were attributable more to increased biochemical limitation to photosynthesis in Year 1 leaves than differences in g(m). However, lower A in leaves on tall trees compared with those on short trees could be attributed in part to lower g(m) and higher stomatal, L(s), and mesophyll, L(m), limitations to photosynthesis, consistent with steeper hydraulic gradients in tall trees.