Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon

Relationships between tree height and crown dimensions and trunk diameter were determined for shade-tolerant species of old-growth forests of western Oregon. The study included both understory and overstory species, deciduous and evergreen angiosperms and evergreen conifers. A comparison of adult understory species with sapling overstory species of similar height showed greater crown width and trunk diameter in the former, whether the comparison is made among conifers or deciduous trees. Conifer saplings had wider crowns than deciduous saplings, but the crown widths of the two groups converged with increase in tree height. Conifer saplings had thicker trunks than deciduous saplings of similar crown width, possibly because of selection for resistance to stem bending under snow loads. The results suggest that understory species have morphologies that increase light interception and persistence in the understory, whereas overstory species allocate their biomass for efficient height growth, thereby attaining the high-light environment of the canopy. The greater crown widths and the additional strength requirements imposed by snow loads on conifer saplings result in less height growth per biomass increment in conifer saplings than in deciduous saplings. However, the convergence in crown width of the two groups at heights greater than 20 m, and the proportionately smaller effect of snow loads on large trees, may result in older conifers equalling or surpassing deciduous trees in biomass allocation to height growth.     

Evaluation of leaf display of evergreen broadleaved tree species and deciduous tree species in warm temperate conifer plantations

We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.     

Age-related trends in red spruce foliar plasticity in relation to declining productivity

Phenotypic plasticity in needle morphology with increasing tree size and age was investigated by comparing four age classes of red spruce (Picea rubens Sarg.) ranging from juvenile (3-12 years old) to mature (over 100 years old). With increase in tree age there were significant increases in leaf mass per unit area (LMA), mesophyll and vascular bundle area as a percentage of total needle cross-sectional area, and stomatal density. Within the vascular bundle, both xylem cross-sectional area and tracheid lumen area increased significantly, whereas air space as a percentage of total cross-sectional area decreased. These morphological changes were associated with a significant decrease in photosynthetic capacity and stomatal conductance, and an increase in (13)C enrichment. Although both photosynthetic capacity and whole-tree conductance decreased significantly between age classes 3 and 12 years, they did not differ between age classes 53 and 127 years, even though needle (13)C enrichment was significantly greater in the 127-year age class. Thus there appear to be compensatory mechanisms that maintain photosynthetic capacity as trees increase in size and vascular complexity, which in red spruce and other species, may affect leaf hydraulic conductance. Although increased LMA may contribute to reduced photosynthetic capacity in red spruce, similar relationships are not seen in other conifers.     

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.     

Gas exchange, leaf structure and nitrogen in contrasting successional tree species growing in open and understory sites during a drought

Seasonal ecophysiology, leaf structure and nitrogen were measured in saplings of early (Populus grandidentata Michx. and Prunus serotina J.F. Ehrh.), middle (Fraxinus americana L. and Carya tomentosa Nutt.) and late (Acer rubrum L. and Cornus florida L.) successional tree species during severe drought on adjacent open and understory sites in central Pennsylvania, USA. Area-based net photosynthesis (A) and leaf conductance to water vapor diffusion (g(wv)) varied by site and species and were highest in open growing plants and early successional species at both the open and understory sites. In response to the period of maximum drought, both sunfleck and sun leaves of the early successional species exhibited smaller decreases in A than leaves of the other species. Shaded understory leaves of all species were more susceptible to drought than sun leaves and had negative midday A values during the middle and later growing season. Shaded understory leaves also displayed a reduced photosynthetic light response during the peak drought period. Sun leaves were thicker and had a greater mass per area (LMA) and nitrogen (N) content than shaded leaves, and early and middle successional species had higher N contents and concentrations than late successional species. In both sunfleck and sun leaves, seasonal A was positively related to predawn leaf Psi, g(wv), LMA and N, and was negatively related to vapor pressure deficit, midday leaf Psi and internal CO(2). Although a significant amount of plasticity occurred in all species for most gas exchange and leaf structural parameters, middle successional species exhibited the largest degree of phenotypic plasticity between open and understory plants.     

Deciduous trees affect small-scale floristic diversity and tree regeneration in conifer forests

Abstract

There is a growing interest in the effects of deciduous trees on biodiversity, soil processes and long-term productivity in boreal, conifer-dominated forests. This study investigated whether individual birch trees allowed to grow to maturity in the coniferous forest can have a local effect on floristic richness and regeneration of tree saplings. The ground vegetation was compared in 2?m radius plots around the stem under the canopies of matched conifer–deciduous trees in a mature, conifer-dominated forest, and included in the analysis variables that could potentially mediate the tree effect (soil pH, cover of lichens, bryophytes, leaf and needle litter). The field layer vegetation was more species rich under birch (Betula pendula and B. pubescens) than under conifers (Picea abies and Pinus sylvestris), and several vascular plant species (including saplings of tree species) occurred more often under birch than under conifers. However, when the effect of the number of subordinate trees was taken into account the difference between birch and pine was not significant. The number of tree regenerations (saplings) was lowest under pines, but did not differ between spruce and birch. There were no effects of the canopy species on soil pH or on cover of lichens and bryophytes. The difference in diversity may be caused by the different effects of leaf and needle litter, and it is also likely that canopy structure has an influence via interception and throughfall and by affecting the light and microclimate.     

Internal leaf anatomy and photosynthetic resource-use efficiency: interspecific and intraspecific comparisons

Leaf mass per unit area (LMA) and internal leaf anatomy often affect net gas exchange because of their effects on internal CO2 conductance to the site of carboxylation, internal shading, competition for CO2 among carboxylation sites, nitrogen concentration and its partitioning. To evaluate effects of LMA and leaf anatomy on CO2 assimilation, water-use efficiency (WUE) and nitrogen-use efficiency (NUE), we measured LMA, leaf thickness, the thickness of mesophyll components, and gas exchange rates at ambient CO2 concentration in leaves of six woody deciduous and evergreen species with different leaf life spans. In two species, CO2 assimilation was also estimated at saturating CO2 concentrations. There were interspecific differences in all morphological variables studied. Long-lived leaves had higher LMA and were thicker than short-lived leaves. Species with high LMA had low assimilation rates and NUE, both in ambient and saturating CO2 concentrations. Thus, in species with high LMA, assimilation was reduced by non-stomatal limitations, possibly because of a lower allocation of N to the photosynthetic machinery than in species with low LMA. Within a species, thicker leaves tended to have a lower tissue density. In intraspecific comparisons under field conditions, increasing internal air volume had positive effects on WUE, probably because of enhanced internal CO2 conductance to the site of carboxylation. We conclude that, in interspecific comparisons, different patterns of N partitioning strongly influence NUE, whereas in intraspecific comparisons, internal leaf anatomy is a key factor regulating resource-use efficiency.     

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.     

Determining the shade tolerance of American chestnut using morphological and physiological leaf parameters

American chestnut (Castanea dentata (Marsh.) Borkh.) was once a principal component of the eastern deciduous forest until it became functionally extinct as a result of the invasive fungus Cryphonectria parasitica (Murr.) Barr. Restoration efforts are underway by means of a blight-resistant American-Chinese chestnut hybrid, and detailed silvicultural information, such as the shade tolerance of American chestnut and optimum site selection for restoration, is critical for planting success. In the present study, the physiological and morphological leaf characteristics of pure American chestnut seedlings, saplings, and mature trees were investigated in one of the few remaining stands of oak-chestnut vegetation (West Salem, WI) to determine shade tolerance. Seedlings, saplings, and mature trees had high maximum rates of photosynthesis, similar to shade intolerant species, and low light compensation points (LCPs), quantum efficiency, leaf mass per area (LMA), and percent nitrogen content, reminiscent of shade tolerant species. Dark respiration rates of seedlings and saplings were low, but increased in mature trees. LMA was found to significantly increase with height in the canopy, indicating a high level of light-induced plasticity. The results of this study suggest that American chestnut should be classified as intermediate in shade tolerance.     

Leaf-age effects on seasonal variability in photosynthetic parameters and its relationships with leaf mass per area and leaf nitrogen concentration within a Pinus densiflora crown

In the temperate zone of Japan, Pinus densiflora Sieb. et Zucc. bears needles of up to three age classes in the upper crown and up to five age classes in the lower crown. To elucidate the effects of leaf age on photosynthetic parameters and its relationships with leaf mass per unit area (LMA) and leaf nitrogen (N(l)) concentration on an area (N(a)) and mass (N(m)) basis, we measured seasonal variations in LMA, N(l), light-saturated photosynthetic rate (A(max)), stomatal conductance (g(s)), maximum rate of carboxylation (V(cmax)) and maximum rate of electron transport (J(max)) in leaves of all age classes in the upper and lower crown. Leaf mass per unit area increased by 27% with increasing leaf age in the lower crown, but LMA did not depend on age in the upper crown. Leaf age had a significant effect on N(m) but not on N(a) in both crown positions, indicating that decreases in N(m) resulted from dilution. Photosynthetic parameters decreased significantly with leaf age in the lower crown (39% for A(max) and 43% for V(cmax)), but the effect of leaf age was not as great in the upper crown, although these parameters exhibited seasonal variation in both crown positions. Regression analysis indicated a close relationship between LMA and N(a), regardless of age class or when each age class was pooled (r(2) = 0.57-0.86). Relationships between LMA and N(a) and among A(max), V(cmax) and J(max) were weak or not significant when all age classes were examined by regression analysis. However, compared with older leaves, relationships among LMA, N(a) and A(max) were stronger in younger leaves. These results indicate that changes in LMA and N(l) mainly reflect light acclimation during leaf development, but they are only slightly affected by irradiance in mature leaves. In conclusion, LMA and N(l) are useful parameters for estimating photosynthetic capacity, but age-related effects need to be taken into account, especially in evergreen conifers.     

Patterns of stomatal conductance among blue oak (Quercus douglasii) size classes and populations: implications for seedling establishment

Quercus douglasii Hook. & Arn. (blue oak) is a deciduous white oak that is currently failing to regenerate throughout much of its range in California, USA. Patterns of water use were observed in adult trees, saplings and seedlings to determine if ontogenetic changes in water use occur, which might be important in the establishment of this long-lived perennial species in a Mediterranean-type system. Seasonal and diurnal stomatal conductance (g(s)), late-season predawn xylem water potentials (Psi(pre)), carbon isotopic ratio (delta(13)C) and soil water status were compared among the three size classes at three sites differing in mean precipitation and soil water characteristics. Comparisons were also made between microsites with and without regeneration (defined by the presence or absence of saplings). Overall patterns of water use were consistent among the three sites, except that, at the site with the highest rainfall, Q. douglasii plants had higher g(s) and more positive Psi(pre) values. Although no differences in water use patterns were found between regeneration and non-regeneration microsites, the observed ontogenetic differences in water use may have important implications for Q. douglasii establishment. Compared with adult trees and saplings, seedlings had higher gas exchange rates during periods of high soil water content (early in the season and in the morning). Seedling g(s) was correlated with percent extractable soil water (ESW) throughout the season; adult tree and sapling g(s) was correlated with ESW between June and September. Despite experiencing greater water stress (indicated by more negative Psi(pre) values) than older trees, seedlings had more negative delta(13)C values, implying lower water-use efficiencies.     

Acclimation of tropical tree species to hurricane disturbance: ontogenetic differences

We investigated acclimation responses of seedlings and saplings of the pioneer species Cecropia schreberiana Miq. and three non-pioneer species, Dacryodes excelsa Vahl, Prestoea acuminata (Willdenow) H.E. Moore var. montana (Graham) Henderson and Galeano, and Sloanea berteriana Choisy ex DC, following a hurricane disturbance in a lower montane wet forest in Puerto Rico. Measurements were made, shortly after passage of the hurricane, on leaves expanded before the hurricane (pre-hurricane leaves) and, at a later time, on recently matured leaves that developed after the hurricane (post-hurricane leaves) from both seedlings and saplings at sites that were severely damaged by the hurricane (disturbed sites) and at sites with little disturbance (undisturbed sites). Pre-hurricane leaves of the non-pioneer species had relatively low light-saturated photosynthetic rates (A(max)) and stomatal conductance (g(s)); neither A(max) nor g(s) responded greatly to the increase in irradiance that resulted from the disturbance, and there were few significant differences between seedlings and saplings. Pre-hurricane leaves of plants at undisturbed sites had low dark respiration rates per unit area (R(d)) and light compensation points (LCP), whereas pre-hurricane leaves of plants at disturbed sites had significantly higher R(d) and LCP. Post-hurricane leaves of plants at disturbed sites had significantly higher A(max) and R(d) than plants at undisturbed sites. Compared with seedlings, saplings had higher A(max) and R(d) and showed greater acclimation to the increase in irradiance that followed the disturbance. Post-hurricane leaves of the non-pioneer species had significantly lower A(max) and were less responsive to changes in irradiance than the pioneer species C. schreberiana. Variation in A(max) across light environments and stages was strongly related to differences in leaf mass per unit area (LMA), especially in the non-pioneer species. As indicated by V(cmax) or J(max) per unit nitrogen, light acclimation of A(max) was determined by leaf morphology (LMA) for the non-pioneer species and by both leaf morphology and leaf biochemistry for C. schreberiana. Ontogenetic changes in A(max) were attributable to changes in leaf morphology. The ontogenetic component of variation in A(max) across light environments and stages differed among species, ranging from 36 to 59% for the non-pioneer species (D. excelsa, 59.3%; P. acuminata var. montana, 44.7%; and S. berteriana, 36.3%) compared with only 17% in the pioneer species C. schreberiana.     

Physiological responses of three deciduous conifers (Metasequoia glyptostroboides, Taxodium distichum and Larix laricina) to continuous light: adaptive implications for the early Tertiary polar summer

Polar regions were covered with extensive forests during the Cretaceous and early Tertiary, and supported trees comparable in size and productivity to those of present-day temperate forests. With a winter of total or near darkness and a summer of continuous, low-angle illumination, these temperate, high-latitude forests were characterized by a light regime without a contemporary counterpart. Although maximum irradiances were much lower than at mid-latitudes, the 24-h photoperiod provided similar integrated light flux. Taxodium, Larix and Metasequoia, three genera of deciduous conifers that occurred in paleoarctic wet forests, have extant, closely related descendents. However, the contemporary relative abundance of these genera differs greatly from that in the paleoarctic. To provide insight into attributes that favor competitive success in a continuous-light environment, we subjected saplings of these genera to a natural photoperiod or a 24-h photoperiod and measured gas exchange, chlorophyll fluorescence, non-structural carbohydrate concentrations, biomass production and carbon allocation. Exposure to continuous light significantly decreased photosynthetic capacity and quantum efficiency of photosystem II in Taxodium and Larix, but had minimal influence in Metasequoia. In midsummer, foliar starch concentration substantially increased in both Taxodium and Larix saplings grown in continuous light, which may have contributed to end-product down-regulation of photosynthetic capacity. In contrast, Metasequoia allocated photosynthate to continuous production of new foliar biomass. This difference in carbon allocation may have provided Metasequoia with a two fold advantage in the paleoarctic by minimizing depression of photosynthetic capacity and increasing photosynthetic surface.     

Rate of stomatal opening, shoot hydraulic conductance and photosynthetic characteristics in relation to leaf abscisic acid concentration in six temperate deciduous trees

Correlations between leaf abscisic acid concentration ([ABA]), stomatal conductance (gs), rate of stomatal opening in response to an increase in leaf water potential (si), shoot hydraulic conductance (L) and photosynthetic characteristics were examined in saplings of six temperate deciduous tree species: Acer platanoides L., Padus avium Mill., Populus tremula L., Quercus robur L., Salix caprea L. and Tilia cordata Mill. Species-specific values of foliar [ABA] were negatively related to the mean values of gs, si, L and light- and CO2- saturated net photosynthesis (P(max)), thus providing strong correlative evidence of a scaling of foliar gas exchange and hydraulic characteristics with leaf endogenous [ABA]. In addition, we suggest that mean gs, si, L and Pmax for mature leaves may partly be determined by the species-specific [ABA] during leaf growth. The most drought-intolerant species had the lowest [ABA] and the highest gs, suggesting that interspecific differences in [ABA] may be linked to differences in species-specific water-use efficiency. Application of high concentrations of exogenous ABA led to large decreases in gs, si and P(max), further underscoring the direct role of ABA in regulating stomatal opening and photosynthetic rate. Exogenous ABA also decreased L, but the decreases were considerably smaller than the decreases in gs, si and Pmax. Thus, exogenous ABA predominantly affected the stomata directly, but modification of L by ABA may also be an important mechanism of ABA action. We conclude that interspecific variability in endogenous [ABA] during foliage growth and in mature leaves provides an important factor explaining observed differences in L, gs, si and Pmax among temperate deciduous tree species.     

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.     

Enhancement of understory productivity by asynchronous phenology with overstory competitors in a temperate deciduous forest

Some saplings and shrubs growing in the understory of temperate deciduous forests extend their periods of leaf display beyond that of the overstory, resulting in periods when understory radiation, and hence productivity, are not limited by the overstory canopy. To assess the importance of the duration of leaf display on the productivity of understory and overstory trees of deciduous forests in the north eastern United States, we applied the simulation model, BIOME-BGC with climate data for Hubbard Brook Experimental Forest, New Hampshire, USA and mean ecophysiological data for species of deciduous, temperate forests. Extension of the overstory leaf display period increased overstory leaf area index (LAI) by only 3 to 4% and productivity by only 2 to 4%. In contrast, extending the growing season of the understory relative to the overstory by one week in both spring and fall, increased understory LAI by 35% and productivity by 32%. A 2-week extension of the growing period in both spring and fall increased understory LAI by 53% and productivity by 55%.     

Hydraulic conductances of angiosperms versus conifers: similar transport sufficiency at the whole-plant level

Bond's "slow seedling" hypothesis proposes that, because of slow growth rates caused by an inefficient transport system and low leaf photosynthetic capacity, gymnosperm seedlings are weak competitors with angiosperms in productive habitats. We measured component (shoot, leaf, and root) and whole-plant hydraulic conductances of sapling-sized tropical plants growing on nitrogen-poor white sand in Borneo. After accounting for size effects, there were no significant differences in conductances between evergreen angiosperms (nine species) and conifers (three species). Plant successional status or transpiration rate seemed more important than soil fertility in determining hydraulic conductance-colonizers had significantly higher whole-plant conductance than late-succession species. Contrary to prediction, leaf hydraulic conductance (normalized by projected leaf area) was unrelated to complexity of venation in conifers and angiosperms, but was highly correlated with whole-plant conductance. Analyses of published data showed that leafless branches of temperate deciduous angiosperms had higher leaf-area normalized hydraulic conductivity than conifers, but there was no significant difference in adult, whole-plant conductance between these taxa. Thus, at the branch level, conifers with narrow tracheids have less efficient transport than angiosperms with wider vessels, but variations in other resistance components and hydraulic architecture (e.g., sapwood/leaf area ratio) ultimately equalize the sufficiency of water transport to leaves of conifers and angiosperms. Although failing to support one of the proposed mechanisms, our findings did not refute the "slow seedling" hypothesis per se.     

Diversity of deciduousness and phenological traits of key Indian dry tropical forest trees

? In seasonally dry tropical forests deciduousness (leaflessness) is an important strategy of trees to survive in water stress period during summer. Deciduousness is a reflection of interacted effect of seasonal drought, tree characteristics and soil moisture conditions.

? The present study aims to document the diversity in leaf pheno-phases in terms of duration of deciduousness (which is reciprocal to growing season length), wood density, leaf mass per area (LMA) and leaf strategy index in 24 important tree species growing in the Vindhyan dry tropical forest in India.

? On the basis of phenological observations, the tree species were categorized into two main groups: leaf exchanging species exhibiting overlapping periods of leaf fall and leaf flush, and deciduous species whose timings of leaf flush and leaf fall differ resulting in a time lag (deciduousness) between the completion of leaf fall and initiation of leaf flush. Presence of wide range of deciduousness duration (from ca. a week to 7 months) among dry tropical trees indicates large variations in their growing season length. In the tree species studied, as the duration of deciduousness increased, leaf flushing period decreased significantly but leaf fall period showed little variation.

? Differing deciduousness in tree species exhibited substantial differences in their leafing (vegetative growth) pattern, as reflected by ratio of durations of leaf flush to leaf fall (leaf strategy index). Across different species, duration of deciduousness was significantly positively correlated with leaf strategy index, and significantly negatively correlated with both wood density and LMA.

? Wide variations in deciduousness, leaf strategy index, wood density and LMA in the 24 species investigated indicate considerable functional diversity in tree species growing in Vindhyan dry tropical region. Variation in seasonal duration of deciduousness among species is reflections of differences in tree functional traits like stem wood density, leaf strategy index and LMA.

     

Leaf phenology,photosynthesis, and the persistence of saplings and shrubs in a mature northern hardwood forest

We quantified leaf phenologies of saplings and overstory trees of sugar maple (Acer saccharum Marsh.) and American beech (Fagus grandifolia Ehrh.), and the shrub hobblebush viburnum (Viburnum alnifolium Marsh.) in a 72-year-old northern hardwood forest. Seasonal changes in irradiance in the shrub layer, and in the leaf CO(2) exchange of viburnum, and sugar maple and beech saplings were also measured. Leaf expansion occurred earlier in the spring and green leaves were retained later in the autumn in saplings and shrubs than in overstory trees. During the spring light phase (before overstory closure), large CO(2) gains by all three shrub-layer species occurred as a result of a combination of relatively large leaf area, high photosynthetic capacity, and high irradiance. Throughout the summer shade phase, photosynthetic capacity at a given irradiance remained relatively constant, but CO(2) gain was typically limited by low irradiances. Even though irradiance in the shrub layer increased during the autumn light phase as the overstory opened, CO(2) gains were modest compared to springtime values because of declining leaf area and photosynthetic capacity in all three species. The CO(2) gains during the spring light phase, and to a lesser extent during the autumn light phase, may be important to the carbon balance and long-term persistence of saplings and shrubs in the usually light-limited shrub layer of a northern hardwood forest. Therefore, for some late-successional species, leaf phenology may be an important characteristic that permits their long-term persistence in the shrub layer of mature northern hardwood forests.     

Leaf gas exchange in the frankincense tree (Boswellia papyrifera) of African dry woodlands

A conceptual model was tested for explaining environmental and physiological effects on leaf gas exchange in the deciduous dry tropical woodland tree Boswellia papyrifera (Del.) Hochst. For this species we aimed at (i) understanding diurnal patterns in leaf gas exchange, (ii) exploring cause-effect relationships among external environment, internal physiology and leaf gas exchange, and (iii) exploring site differences in leaf gas exchange in response to environmental variables. Diurnal courses in gas exchange, underlying physiological traits and environmental variables were measured for 90 trees on consecutive days at two contrasting areas, one at high and the other at low altitude. Assimilation was highest in the morning and slightly decreased during the day. In contrast, transpiration increased from early morning to midday, mainly in response to an increasing vapor pressure deficit (VPD) and gradual stomatal closure. The leaf water potential varied relatively little and did not influence gas exchange during the measurement period. Our results suggest that the same cause-effect relationships function at contrasting areas. However, leaves at the higher altitude had higher photosynthetic capacity, reflecting acclimation to higher light levels. Trees at both areas nevertheless achieved similar leaf assimilation rates since assimilation was down-regulated by stomatal closure due to the higher VPD at the higher altitude, while it became more light limited at the lower altitude. Gas exchange was thus limited by a high VPD or low light levels during the wet season, despite the ability of the species to acclimate to different conditions.