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Optimality theory states that whole-tree carbon gain is maximized when leaf N and photosynthetic capacity profiles are distributed along vertical light gradients such that the marginal gain of nitrogen investment is identical among leaves. However, observed photosynthetic N gradients in trees do not follow this prediction, and the causes for this apparent discrepancy remain uncertain. Our objective was to evaluate how hydraulic limitations potentially modify crown-level optimization in Sequoiadendron giganteum (giant sequoia) trees up to 90 m tall. Leaf water potential (Ψ l ) and branch sap flow closely followed diurnal patterns of solar radiation throughout each tree crown. Minimum leaf water potential correlated negatively with height above ground, while leaf mass per area (LMA), shoot mass per area (SMA), leaf nitrogen content (%N), and bulk leaf stable carbon isotope ratios (δ(13)C) correlated positively with height. We found no significant vertical trends in maximum leaf photosynthesis (A), stomatal conductance (g s), and intrinsic water-use efficiency (A/g s), nor in branch-averaged transpiration (E L), stomatal conductance (G S), and hydraulic conductance (K L). Adjustments in hydraulic architecture appear to partially compensate for increasing hydraulic limitations with height in giant sequoia, allowing them to sustain global maximum summer water use rates exceeding 2000 kg day(-1). However, we found that leaf N and photosynthetic capacity do not follow the vertical light gradient, supporting the hypothesis that increasing limitations on water transport capacity with height modify photosynthetic optimization in tall trees.

Structural and physiological changes that occur as trees grow taller are associated with increased hydraulic constraints on leaf gas exchange, yet it is unclear if leaf-level constraints influence whole-tree growth as trees approach their maximum size. We examined variation in leaf physiology, leaf area to sapwood area ratio (L/S), and annual aboveground growth across a range of tree heights in Eucalyptus regnans. Leaf photosynthetic capacity did not differ among upper crown leaves of individuals 61.1-92.4 m tall. Maximum daily and integrated diurnal stomatal conductance (g s) averaged 36 and 34% higher, respectively, in upper crown leaves of ~60-m-tall, 80-year-old trees than in ~90-m-tall, 300-year-old trees, with larger differences observed on days with a high vapor pressure deficit (VPD). Greater stomatal regulation in taller trees resulted in similar minimum daily leaf water potentials (Ψ L) in shorter and taller trees over a broad range of VPDs. The long-term stomatal limitation on photosynthesis, as inferred from leaf δ (13)C composition, was also greater in taller trees. The δ (13)C of wood indicated that the bulk of photosynthesis used to fuel wood production in the main trunk and branches occurred in the upper crown. L/S increased with tree height, especially after accounting for size-independent variation in crown structure across 27 trees up to 99.8 m tall. Despite greater stomatal limitation of leaf photosynthesis in taller trees, total L explained 95% of the variation in annual aboveground biomass growth among 15 trees measured for annual biomass growth increment in 2006. Our results support a theoretical model proposing that, in the face of increasing hydraulic constraints with height, whole-tree growth is maximized by a resource trade-off that increases L to maximize light capture rather than by reducing L/S to sustain g s.

We studied changes in morphological and physiological characteristics of leaves and shoots along a height gradient in Sequoia sempervirens, the tallest tree species on Earth, to investigate whether morphological and physiological acclimation to the vertical light gradient was constrained by hydrostatic limitation in the upper crown. Bulk leaf water potential (Psi) decreased linearly and light availability increased exponentially with increasing height in the crown. During the wet season, Psi was lower in the outer than inner crown. C isotope composition of leaves (delta(13)C) increased with increasing height indicating greater photosynthetic water use efficiency in the upper crown. Leaf and shoot morphology changed continuously with height. In contrast, their relationships with light availability were discontinuous: morphological characteristics did not correspond to increasing light availability above 55-85 m. Mass-based chlorophyll concentration (chl) decreased with increasing height and increasing light availability. In contrast, area-based chl remained constant or increased with increasing height. Mass-based maximum rate of net photosynthesis (P (max)) decreased with increasing height, whereas area-based P (max) reached maximum at 78.4 m and decreased with increasing height thereafter. Mass-based P (max) increased with increasing shoot mass per area (SMA), whereas area-based P (max) was not correlated with SMA in the upper crown. Our results suggest that hydrostatic limitation of morphological development constrains exploitation of light in the upper crown and contributes to reduced photosynthetic rates and, ultimately, reduced height growth at the tops of tall S. sempervirens trees.

We compared ammonium and nitrate nutrition in Eriophorum scheuchzeri and E. vaginatum, two Alaskan sedges that are native to high- and low-fertility sites, respectively. When grown in solution culture, the two species were similar in their kinetics of NH inf4 sup+ NO inf3 sup- absorption: at nitrogen concentrations below 50 μM, net NH inf4 sup+ and NO inf3 sup- were absorbed at similar rates, but at higher concentrations, net uptake of NO inf3 sup- was significantly faster than that of NH inf4 sup+ . The two species also showed similar abilities to assimilate NO inf3 sup- . Growth of E. vaginatum under NO inf3 sup- nutrition was only slightly less than that under NH inf4 sup+ . The observed similarities between these species from contrasting edaphic habitats indicate that factors other than tissue-specific rates of nitrogen acquisition and assimilation may underlie local adaptation to soil N fertility. Moreover, the capacity of these species to exploit NO inf3 sup- as a N source supports the view that NO inf3 sup- availability may be significant even in wet, acidic, arctic soils.

Two annual species of Bromus, an invader (B. hordeaceus, ex B. mollis) and a non-invader (B. intermedius), were grown for 28 days in growth chambers, at 5 and 100 μM NO 3 - in flowing nutrient solution. No differences between the two species were observed at either NO 3 - level, in terms of relative growth rate (RGR) or its components, dry matter partitioning, specific NO 3 - absorption rate, nitrogen concentration, and other characteristics of NO 3 - uptake and photosynthesis. The effects of decreasing NO 3 - concentration in the solution were mainly to decrease the NO 3 - concentration in the plants through decreased absorption rate, and to decrease the leaf area ratio through increased specific leaf mass and decreased leaf mass ratio. Organic nitrogen concentration varied little between the two treatments, which may be the reason why photosynthetic rates were not altered. Consequently, RGR was only slightly decreased in the 5-μM treatment compared to the 100-μM treatment. This is in contrast with other species, where growth is reduced at much higher NO 3 - concentrations. These discrepancies may be related to differences in RGR, since a log-linear relationship was found between RGR and the NO 3 - concentration at which growth is first reduced. In addition, a strong linear relationship was found between the RGR of these species and their maximum absorption rate for nitrate, suggesting that the growth of species with low maximum RGR may be partly regulated by nutrient uptake.

10 broadleafed trees and shrubs native to the mediterranean climactic zone in California were surveyed for their photosynthetic and stomatal responses to SO 2 . These species ranged from drought deciduous to evergreen and had diverse responses to SO 2 . These results suggest an approach for predicting SO 2 resistances of plants.We found that conductance values of plants in SO 2 -free air can be used to estimate the quantity of SO 2 which plants absorb. These estimates are based on conductance values for plants in non-limiting environmental conditions. SO 2 absorption quantities are then used to predict relative photosynthesis following the fumigation. Thus, relative photosynthesis of plants following fumigation can be predicted on the basis of conductance in SO 2 -free air. This approach to predicting SO 2 resistances of plants includes analysis of their stomatal responses to fumigation, their characteristics of SO 2 adsorption and absorption, and their change in photosynthesis resulting from SO 2 stress.

Measurements and modeling of photosynthesis, respiration and growth in flowers of Diplacus aurantiacus, a semidrought-deciduous shrub, indicate that they can provide 18%-25% of their total carbon requirements through photosynthesis of flower parts. Daily photosynthetic carbon fixation exceeds daily respiratory CO 2 loss during most non-fruiting stages of development. However, this carbon gain fails to meet the requirements for new biomass construction during bud growth and corolla expansion. During fruiting stages, insufficient carbon is fixed to fully supply either respiration or growth.The calyx performs most of the flower's photosynthesis throughout the life of the flower. However, during stages of fruit development, the contribution of the ovary to flower photosynthesis may equal that of the calyx.