A review of light interception in plant stands from leaf to canopy in different plant functional types and in species with varying shade tolerance

Changes in the efficiency of light interception and in the costs for light harvesting along the light gradients from the top of the plant canopy to the bottom are the major means by which efficient light harvesting is achieved in ecosystems. In the current review analysis, leaf, shoot and canopy level determinants of plant light harvesting, the light-driven plasticity in key traits altering light harvesting, and variations among different plant functional types and between species of different shade tolerance are analyzed. In addition, plant age- and size-dependent alterations in light harvesting efficiency are also examined. At the leaf level, the variations in light harvesting are driven by alterations in leaf chlorophyll content modifies the fraction of incident light harvested by given leaf area, and in leaf dry mass per unit area (M _A) that determines the amount of leaf area formed with certain fraction of plant biomass in the leaves. In needle-leaved species with complex foliage cross-section, the degree of foliage surface exposure also depends on the leaf total-to-projected surface area ratio. At the shoot scale, foliage inclination angle distribution and foliage spatial aggregation are the major determinants of light harvesting, while at the canopy scale, branching frequency, foliage distribution and biomass allocation to leaves (F _L) modify light harvesting significantly. F _L decreases with increasing plant size from herbs to shrubs to trees due to progressively larger support costs in plant functional types with greater stature. Among trees, F _L and stand leaf area index scale positively with foliage longevity. Plant traits altering light harvesting have a large potential to adjust to light availability. Chlorophyll per mass increases, while M _A, foliage inclination from the horizontal and degree of spatial aggregation decrease with decreasing light availability. In addition, branching frequency decreases and canopies become flatter in lower light. All these plastic modifications greatly enhance light harvesting in low light. Species with greater shade tolerance typically form a more extensive canopy by having lower M _A in deciduous species and enhanced leaf longevity in evergreens. In addition, young plants of shade tolerators commonly have less strongly aggregated foliage and flatter canopies, while in adult plants partly exposed to high light, higher shade tolerance of foliage allows the shade tolerators to maintain more leaf layers, resulting in extended crowns. Within a given plant functional type, increases in plant age and size result in increases in M _A, reductions in F _L and increases in foliage aggregation, thereby reducing plant leaf area index and the efficiency of light harvesting. Such dynamic modifications in plant light harvesting play a key role in stand development and productivity. Overall, the current review analysis demonstrates that a suite of chemical and architectural traits at various scales and their plasticity drive plant light harvesting efficiency. Enhanced light harvesting can be achieved by various combinations of traits, and these suites of traits vary during plant ontogeny.     

Plant growth-form alters the relationship between foliar morphology and species shade-tolerance ranking in temperate woody taxa

Variation in leaf size (area per leaf) and leaf dry weight per area (LWA) in relation to species shade- and drought-tolerance, characterised by Ellenberg's light (ELD) and water demand (EWD) values, respectively, were examined in 60 temperate woody taxa at constant relative irradiance. LWA was independent of plant size, but leaf size increased with total plant height at constant ELD. Canopy position also affected leaf morphology: leaves from the upper crown third had higher LWA and were larger than leaves from the lower third. Leaf size and LWA were negatively correlated, and leaf size decreased and LWA increased with decreasing species shade-tolerance. Mean LWA was similar for trees and shrubs, but trees had larger leaves than shrubs. Furthermore, all relationships were altered by plant growth-form: none of the qualitative tendencies was significant for trees. This implies the considerably lower plasticity of foliar parameters in trees than those in shrubs. Accordingly, shade-tolerance of trees, having relatively constant leaf structure, may be most affected by the variability in biomass partitioning and crown geometry which influence foliage distribution and spacing and finally determine canopy light absorptance. Alteration of leaf form and investment pattern for construction of unit foliar surface area which change the efficiency of light interception per unit biomass investment in leaves, is a competitive strategy inherent to shrubs. EWD as well as wood anatomy did not control LWA and leaf size, though there was a trend of ring-porous tree species to be more shade-tolerant than diffuse-porous trees. Since ring-porous species are more vulnerable to cavitation than diffuse-porous species, they may be constrained to environments where irradiances and consequently evaporative demand is lower.     

Distribution patterns of foliar carbon and nitrogen as affected by tree dimensions and relative light conditions in the canopy of Picea abies

 Variations in the partitioning of foliar carbon and nitrogen in combination with changes in needle and shoot structure were studied in trees of Picea abies along a vertical gradient of relative irradiance (RI). RI was the major determinant of needle morphology, causing all needle linear parameters – width, thickness and length – to increase. Due to the different responsiveness of needle thickness and width in respect of RI, the ratio of total to projected needle area increased with RI. Furthermore, shoot structure was also influenced by RI, and the ratio of shoot silhouette area to total needle area, which characterises the packing of needles and needle area within the shoot, was greater at lower values of irradiance. Needle dry weight per total needle area (LWA_t) was also increased by RI. Similarly, irrespective of the measure for surface area, needle nitrogen content per area, as the product of needle dry weight per area and nitrogen content per needle dry weight (N_m), scaled quasi-linearly with needle weight per area. Thus, the changes in needle and shoot morphology made it possible to invest more photosynthesising weight per unit light-intercepting surface there, where the pay-back due to elevated irradiances was the highest. However, N_m behaved in an entirely different manner, decreasing hyperbolically with LWA_t. Since non-structural (carbon in non-structural carbohydrates), and structural (total minus non-structural) needle carbon per dry weight also increased with LWA_t, N_m was inversely correlated with both non-structural and structural carbon. Total tree height, increasing significantly LWA_t, also influenced needle structure. It appeared that total height did not affect needle thickness or width, but larger trees had greater needle density (dry weight per volume). Because needle density was positively correlated with needle carbon content per dry weight, it was assumed that the greater values of needle carbon content can be attributed to increased lignification and thickening of needle cell walls. Thus, it appeared that the proportion of supporting structures was greater in needles of larger trees. Inasmuch as an increased fraction of supporting structures dilutes other leaf substances, including also leaf compounds responsible for CO₂-assimilation, enhanced requirement for supporting structures may be responsible for lower rates of carbon assimilation per foliage dry weight observed in large trees. Increasing water limitation with increasing tree size is discussed as a possible cause for increased needle supporting costs in large trees. Received: 2 April 1995 / Accepted: 16 February 1996     

A model separating leaf structural and physiological effects on carbon gain along light gradients for the shade-tolerant species Acer saccharum

A process-based leaf gas exchange model for C₃ plants was developed which specifically describes the effects observed along light gradients of shifting nitrogen investment in carboxylation and bioenergetics and modified leaf thickness due to altered stacking of photosynthetic units. The model was parametrized for the late-successional, shade-tolerant deciduous species Acer saccharum Marsh. The specific activity of ribulose-1,5-bisphosphate carboxylase (Rubisco) and the maximum photosynthetic electron transport rate per unit cytochrome f (cyt f) were used as indices that vary proportionally with nitrogen investment in the capacities for carboxylation and electron transport. Rubisco and cyt f per unit leaf area are related in the model to leaf dry mass per area (M_A), leaf nitrogen content per unit leaf dry mass (N_m), and partitioning coefficients for leaf nitrogen in Rubisco (P_R) and in bioenergetics (P_B). These partitioning coefficients are estimated from characteristic response curves of photosynthesis along with information on lear structure and composition. While P_R and P_B determine the light-saturated value of photosynthesis, the fraction of leaf nitrogen in thylakoid light-harvesting components (P_L) and the ratio of leaf chlorophyll to leaf nitrogen invested in light harvesting (C_B), which is dependent on thylakoid stoichiometry, determine the initial photosynthetic light utilization efficiency in the model. Carbon loss due to mitochondrial respiration, which also changes along light gradients, was considered to vary in proportion with carboxylation capacity. Key model parameters - N_m, P_R, P_B, P_LC_B and stomatal sensitivity with respect to changes in net photosynthesis (G_r) – were examined as a function of M_A, which is linearly related to irradiance during growth of the leaves. The results of the analysis applied to A. saccharum indicate that P_B and P_R increase, and G_f, P_L and C_B decrease with increasing M_A. As a result of these effects of irradiaiice on nitrogen partitioning, the slope of the light-saturated net photosynthesis rate per unit leaf dry mass (A^m_max) versus N_m relationship increased with increasing growth irradiance in mid-season. Furthermore, the nitrogen partitioning coefficients as well as the slopes of A^m_max versus N_m were independent of season, except during development of the leaf photosynthetic apparatus. Simulations revealed that the acclimation to high light increased A^m_max by 40% with respect to the low light regime. However, light-saturated photosynthesis per leaf area (A^a_max) varied 3-fold between these habitats, suggesting that the acclimation to high light was dominated by adjustments in leaf anatomy (A^a_max=A^m_maxM_A) rather than in foliar biochemistry. This differed from adaptation to low light, where the alterations in foliar biochemistry were predicted to be at least as important as anatomical modifications. Due to the light-related accumulation of photosynthetic mass per unit area, A^a_max depended on M_A and leaf nitrogen per unit area (N_a). However, N_a conceals the variation in both M_A and N_m (N_a=N_mM_A), and prevents clear separation of anatomical adjustments in foliage structure and biochemical modifications in foliar composition. Given the large seasonal and site nutrient availability-related variation in N_m, and the influences of growth irradiance on nitrogen partitioning, the relationship between A^a_max and N_a is universal neither in time nor in space and in natural canopies at mid-season is mostly driven by variability in M_A. Thus, we conclude that analyses of the effects of nitrogen investments on potential carbon acquisition should use mass-based rather than area-based expressions.     

Structural and physiological responses of two invasive weeds, <Emphasis Type="Italic">Mikania micrantha</Emphasis> and <Emphasis Type="Italic">Chromolaena odorata</Emphasis>, to contrasting light and soil water conditions

To better understand the requirement of light and soil water conditions in the invasion sites of two invasive weeds, Mikania micrantha and Chromolaena odorata, we investigated their structural and physiological traits in response to nine combined treatments of light [full, medium and low irradiance (LI)] and soil water (full, medium and low field water content) conditions in three glasshouses. Under the same light conditions, most variables for both species did not vary significantly among different water treatments. Irrespective of water treatment, both species showed significant decreases in maximum light saturated photosynthetic rate (P _max), photosynthetic nitrogen-use efficiency, and relative growth rate under LI relative to full irradiance; specific leaf area, however, increased significantly from full to LI though leaf area decreased significantly, indicating that limited light availability under extreme shade was the critical factor restricting the growth of both species. Our results also indicated that M. micrantha performed best under a high light and full soil water combination, while C. odorata was more efficient in growth under a high light and medium soil water combination.     

The effect of growth irradiance on leaf anatomy and photosynthesis in Acer species differing in light demand

Variation in light demand is a major factor in determining the growth and survival of trees in a forest. There is strong relation between the light‐demand and the effect of growth irradiance on leaf morphology and photosynthesis in three Acer species: A. rufinerve (light‐demanding), A. mono (intermediate) and A. palmatum (shade‐tolerant). The increase in mesophyll thickness and surface area of chloroplasts facing the intercellular airspaces (S_c) with growth irradiance was highest in A. rufinerve. Although the increase in light‐saturated photosynthesis (A_max) was similar among the species, the increase in water use efficiency (WUE) was much higher in A. rufinerve than that in the other species, indicating that the response to water limitation plays an important role in leaf photosynthetic acclimation to high light in A. rufinerve. The low CO₂ partial pressure at the carboxylation site (C_c) in A. rufinerve (130 µmol mol^?1) at high irradiance was caused by low stomatal and internal conductance to CO₂ diffusion, which minimized the increase in A_max in A. rufinerve despite its high Rubisco content. Under shade conditions, interspecific differences in leaf features were relatively small. Thus, difference in light demand related to leaf acclimation to high light rather than that to low light in the Acer species.     

Size-dependent variability of leaf and shoot hydraulic conductance in silver birch

Variation in leaf and shoot hydraulic conductance was examined on detached shoots of silver birch (Betula pendula Roth), cut from the lower third (shade leaves) and upper third of the crown (sun leaves) of large trees growing in a natural temperate forest stand. Hydraulic conductances of whole shoots (K _S), leaf blades (K _lb), petioles (K _P) and branches (i.e. leafless stem; K _B) were determined by water perfusion using a high-pressure flow meter in quasi-steady state mode. The shoots were exposed to irradiance of photosynthetic photon flux density of 200–250 μmol m⁻² s⁻¹, using different light sources. K _lb depended significantly (P < 0.001) on light quality, canopy position and leaf blade area (A _L). K _lb increased from crown base to tree top, in parallel with vertical patterns of A _L. However, the analysis of data on shade and sun leaves separately revealed an opposite trend: the bigger the A _L the higher K _lb. Leaf anatomical study of birch saplings revealed that this trend is attributable to enhanced vascular development with increasing leaf area. Hydraulic traits (K _S, K _B, K _lb) of sun shoots were well co-ordinated and more strongly correlated with characteristics of shoot size than those of shade shoots, reflecting their greater evaporative load and need for stricter adjustment of hydraulic capacity with shoot size. K _S increased with increasing xylem cross-sectional area to leaf area ratio (Huber value; P < 0.01), suggesting a preferential investment in water-conducting tissue (sapwood) relative to transpiring tissue (leaves), and most likely contributing to the functional stability of the hydraulic system, essential for fast-growing pioneer species.     

Changes in foliage distribution with relative irradiance and tree size: Differences between the saplings ofAcer platanoides andQuercus robur

Dependencies of foliage arrangement and structure on relative irradiance and total height (TH) were studied in saplings ofAcer platanoides andQuercus robur. The distribution of relative foliar area and dry weight (leaf area and weight in a crown layer per total tree leaf area and weight, respectively) were examined with respect to relative height (RH, height in the crown per TH) and characterized by the Weibull function. The distributions of relative area and weight were nearly identical, and the differences between them were attributable to a systematic decline in leaf dry weight per area with increasing crown depth. Foliage distribution was similarly altered by tree size in both species; RH at foliage maximum was lower and relative canopy size (RCS, length of live crown per TH) greater in taller trees. However, the distribution was more uniform inA. platanoides than inQ. robur. Apart from the size effects, relative irradiance also influenced canopy structure; RCS increased inQ. platanoides and decreased inQ. robur with increasing irradiance. As crown architecture was modified by irradiance, foliage distribution was shifted upwards with decreasing irradiance inA. platanoides, but it was independent of irradiance inQ. robur. Higher foliage maximum at lower irradiance in more shade-tolerantA. platanoides is likely to contribute towards more efficient foliar display for light interception and increase the competitive ability of this species in light-limited environments. Consequently, these differences in crown architecture and foliage distribution may partly explain the superior behavior ofA. platanoides in understory.     

A tropical epiphytic orchid uses a low‐light interception strategy in a spatially heterogeneous light environment

Light is considered a non‐limiting factor for vascular epiphytes. Nevertheless, an epiphyte's access to light may be limited by phorophyte shading and the spatio‐temporal environmental patchiness characteristic of epiphytic habitats. We assessed the extent to which potential light interception in Rodriguezia granadensis, an epiphytic orchid, is determined by individual factors (plant size traits and leaf traits), or environmental heterogeneity (light patchiness) within the crown of the phorophyte, or both. We studied 104 adult plants growing on Psidium guajava trees in two habitats with contrasting canopy cover: a dry tropical forest edge, and isolated trees in a pasture. We recorded the number of leaves and the leaf area, the leaf position angles, and the potential exposure of the leaf surface to direct irradiance (silhouette area of the leaf blade), and the potential irradiance incident on each plant. We found the epiphytes experience a highly heterogeneous light environment in the crowns of P. guajava. Nonetheless, R. granadensis plants displayed a common light interception strategy typical of low‐light environments, resembling terrestrial, forest understory plants. Potential exposure of the total leaf surface to direct irradiance correlated positively with plant size and within‐plant variation in leaf orientation. In many‐leaved individuals, within‐plant variation in leaf angles produced complementary leaf positions that enhanced potential light interception. This light interception strategy suggests that, in contrast to current wisdom, enhancing light capture is important for vascular epiphytes in canopies with high spatio‐temporal heterogeneity in light environments.     

Diaheliotropic leaf movement enhances leaf photosynthetic capacity and photosynthetic light and nitrogen use efficiency via optimising nitrogen partitioning among photosynthetic components in cotton (Gossypium hirsutum L.)

• Phototropic leaf movement of plants is an effective mechanism for adapting to light conditions. Light is the major driver of plant photosynthesis. Leaf N is also an important limiting factor on leaf photosynthetic potential. Cotton (Gossypium hirsutum L.) exhibits diaheliotropic leaf movement. Here, we compared the long‐term photosynthetic acclimation of fixed leaves (restrained) and free leaves (allowed free movement) in cotton.
• The fixed leaves and free leaves were used for determination of PAR, leaf chlorophyll concentration, leaf N content and leaf gas exchange. The measurements were conducted under clear sky conditions at 0, 7, 15 and 30 days after treatment (DAT).
• The results showed that leaf N allocation and partitioning among different components of the photosynthetic apparatus were significantly affected by diaheliotropic leaf movement. Diaheliotropic leaf movement significantly increased light interception per unit leaf area, which in turn affected leaf mass per area (LMA), leaf N content (N_A) and leaf N allocation to photosynthesis (N_P). In addition, cotton leaves optimised leaf N allocation to the photosynthetic apparatus by adjusting leaf mass per area and N_A in response to optimal light interception.
• In the presence of diaheliotropic leaf movement, cotton leaves optimised their structural tissue and photosynthetic characteristics, such as LMA, N_A and leaf N allocation to photosynthesis, so that leaf photosynthetic capacity was maximised to improve the photosynthetic use efficiency of light and N under high light conditions.

     

Within-canopy variation in the rate of development of photosynthetic capacity is proportional to integrated quantum flux density in temperate deciduous trees

The present study was undertaken to test for the hypothesis that the rate of development in the capacity for photosynthetic electron transport per unit area (J_max;A), and maximum carboxylase activity of Rubisco (V_cmax;A) is proportional to average integrated daily quantum flux density (Q_int) in a mixed deciduous forest dominated by the shade‐intolerant species Populus tremula L., and the shade‐tolerant species Tilia cordata Mill. We distinguished between the age‐dependent changes in net assimilation rates due to modifications in leaf dry mass per unit area (M_A), foliar nitrogen content per unit dry mass (N_M), and fractional partitioning of foliar nitrogen in the proteins of photosynthetic electron transport (F_B), Rubisco (F_R) and in light‐harvesting chlorophyll‐protein complexes (V_cmax;A ∝ M_AN_MF_R; J_max;A ∝ M_AN_MF_B). In both species, increases in J_max;A and V_cmax;A during leaf development were primarily determined by nitrogen allocation to growing leaves, increases in leaf nitrogen partitioning in photosynthetic machinery, and increases in M_A. Canopy differences in the rate of development of leaf photosynthetic capacity were mainly controlled by the rate of change in M_A. There was only small within‐canopy variation in the initial rate of biomass accumulation per unit Q_int (slope of M_A versus leaf age relationship per unit Q_int), suggesting that canopy differences in the rate of development of J_max;A and V_cmax;A are directly proportional to Q_int. Nevertheless, M_A, nitrogen, J_max;A and V_cmax;A of mature leaves were not proportional to Q_int because of a finite M_A in leaves immediately after bud‐burst (light‐independent component of M_A). M_A, leaf chlorophyll contents and chlorophyll : N ratio of mature leaves were best correlated with the integrated average quantum flux density during leaf development, suggesting that foliar photosynthetic apparatus, once developed, is not affected by day‐to‐day fluctuations in Q_int. However, for the upper canopy leaves of P. tremula and for the entire canopy of T. cordata, there was a continuous decline in N contents per unit dry mass in mature non‐senescent leaves on the order of 15–20% for a change of leaf age from 40 to 120 d, possibly manifesting nitrogen reallocation to bud formation. The decline in N contents led to similar decreases in leaf photosynthetic capacity and foliar chlorophyll contents. These data demonstrate that light‐dependent variation in the rate of developmental changes in M_A determines canopy differences in photosynthetic capacity, whereas foliar photosynthetic apparatus is essentially constant in fully developed leaves.     

Why are Sun Leaves Thicker than Shade Leaves? — Consideration based on Analyses of CO2 Diffusion in the Leaf

A ) depend not only on photosynthetic biochemistry but also on mesophyll structure. Because resistance to CO₂ diffusion from the substomatal cavity to the stroma is substantial, it is likely that mesophyll structure affects A through affecting diffusion of CO₂ in the leaf. To evaluate effects of various aspects of mesophyll structure on photosynthesis, we constructed a one-dimensional model of CO₂ diffusion in the leaf. When mesophyll thickness of the leaf is changed with the Rubisco content per unit leaf area kept constant, the maximum A occurs at an almost identical mesophyll thickness irrespective of the Rubisco contents per leaf area. On the other hand, with an increase in Rubisco content per leaf area, the mesophyll thickness that realizes a given photosynthetic gain per mesophyll thickness (or per leaf cost) increases. This probably explains the strong relationship between A and mesephyll thickness. In these simulations, an increase in mesophyll thickness simultaneously means an increase in the diffusional resistance in the intercellular spaces (R _ias), an increase in the total surface area of chloroplasts facing the intercellular spaces per unit leaf area (S _c ), and an increase in construction and maintenance cost of the leaf. Leaves can increase S _c and decrease R _ias also by decreasing cell size. Leaves with smaller cells are mechanically stronger. However, actual leaves do not have very small cells. This could be because actual leaves exhibiting considerable rates of leaf area expansion, adequate heat capacitance, high efficiency of N and/or P use, etc, are favoured. Relationships between leaf longevity and mesophyll structure are also discussed. Received 20 September 2000/ Accepted in revised form 4 January 2001     

Herbivory alters plant carbon assimilation,patterns of biomass allocation and nitrogen use efficiency

Herbivory can trigger physiological processes resulting in leaf and whole plant functional changes. The effects of chronic infestation by an insect on leaf traits related to carbon and nitrogen economy in three Prunus avium cultivars were assessed. Leaves from non-infested trees (control) and damaged leaves from infested trees were selected. The insect larvae produce skeletonization of the leaves leaving relatively intact the vein network of the eaten leaves and the abaxial epidermal tissue. At the leaf level, nitrogen content per mass (N_mass) and per area (N_area), net photosynthesis per mass (A_mass) and per area (A_area), photosynthetic nitrogen-use efficiency (PNUE), leaf mass per area (LMA) and total leaf phenols content were measured in the three cultivars. All cultivars responded to herbivory in a similar fashion. The N_mass, A_mass, and PNUE decreased, while LMA and total content of phenols increased in partially damaged leaves. Increases in herbivore pressure resulted in lower leaf size and total leaf area per plant across cultivars. Despite this, stem cumulative growth tended to increase in infected plants suggesting a change in the patterns of biomass allocation and in resources sequestration elicited by herbivory. A larger N investment in defenses instead of photosynthetic structures may explain the lower PNUE and A_mass observed in damaged leaves. Some physiological changes due to herbivory partially compensate for the cost of leaf removal buffering the carbon economy at the whole plant level.     

Leaf functional traits of Neotropical savanna trees in relation to seasonal water deficit

The seasonal savannas (cerrados) of Central Brazil are characterized by a large diversity of evergreen and deciduous trees, which do not show a clear differentiation in terms of active rooting depth. Irrespective of the depth of the root system, expansion of new foliage in deciduous species occurs at the end of the dry season. In this study, we examined a suite of leaf traits related to C assimilation, water and nutrients (N, P) in five deciduous and six evergreen trees that were among the dominant families of cerrado vegetation. Maximum CO₂ assimilation on a mass basis (A_mass) was significantly correlated with leaf N and P, and specific leaf area (SLA; leaf area per unit of leaf mass). The highest leaf concentrations of both nutrients were measured in the newly mature leaves of deciduous species at the end of the dry period. The differences in terms of leaf N and P between evergreen and deciduous species decreased during the wet season. Deciduous species also invested less in the production of non-photosynthetic leaf tissues and produced leaves with higher SLA and maintained higher water use efficiency. Thus, deciduous species compensated for their shorter leaf payback period by maintaining higher potential payback capacity (higher values of A_mass) and lower leaf construction costs (higher SLA). Their short leafless period and the capacity to flush by the end of the dry season may also contribute to offset the longer payback period of evergreen species, although it may involve the higher cost of maintaining a deep-root system or a tight control of plant water balance in the shallow-rooted ones.     

Leaf traits of natural populations of <Emphasis Type="Italic">Adiantum reniforme</Emphasis> var. <Emphasis Type="Italic">sinensis</Emphasis>, endemic to the Three Gorges region in China

Leaf mass per unit area (LMA), carbon and nitrogen contents, leaf construction cost, and photosynthetic capacity (P _max) of Adiantum reniforme var. sinensis, an endangered fern endemic to the Three Gorges region in southwest China, were compared in five populations differing in habitat such as soil moisture and irradiance. The low soil moisture and high irradiance habitat population exhibited significantly higher LMA, area-based leaf construction (CC_A), and carbon content (C_A), but lower leaf nitrogen content per unit dry mass (N_M) than the other habitat populations. The high soil moisture and low irradiance habitat populations had the lowest CC_A, but their cost/benefic ratios of CCA/P _max were similar to the medium soil moisture and irradiance habitat population due to their lower leaf P _max. Hence A. reniforme var. sinensis prefers partially shaded, moist but well-drained, slope habitats. Due to human activities, however, its main habitats now are cliffs or steeply sloped bare rocks with poor and thin soil. The relatively high energy requirements and low photosynthetic capacity in these habitats could limit the capability of the species in extending population or interspecific competition and hence increase its endangerment.     

Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees

We investigated how water transport capacity, wood density and wood anatomy were related to leaf photosynthetic traits in two lowland forests in Panama. Leaf-specific hydraulic conductivity (k_L) of upper branches was positively correlated with maximum rates of net CO₂ assimilation per unit leaf area (A_area) and stomatal conductance (g_s) across 20 species of canopy trees. Maximum k_L showed stronger correlation with A_area than initial k_L suggesting that allocation to photosynthetic potential is proportional to maximum water transport capacity. Terminal branch k_L was negatively correlated with A_area/g_s and positively correlated with photosynthesis per unit N, indicating a trade-off of efficient use of water against efficient use of N in photosynthesis as water transport efficiency varied. Specific hydraulic conductivity calculated from xylem anatomical characteristics (k_theoretical) was positively related to A_area and k_L, consistent with relationships among physiological measurements. Branch wood density was negatively correlated with wood water storage at saturation, k_L, A_area, net CO₂ assimilation per unit leaf mass (A_mass), and minimum leaf water potential measured on covered leaves, suggesting that wood density constrains physiological function to specific operating ranges. Kinetic and static indices of branch water transport capacity thus exhibit considerable co-ordination with allocation to potential carbon gain. Our results indicate that understanding tree hydraulic architecture provides added insights to comparisons of leaf level measurements among species, and links photosynthetic allocation patterns with branch hydraulic processes.     

Leaf position,light levels,and nitrogen allocation in five species of rain forest pioneer trees

We used path analysis to ask whether leaf position or leaf light level was a better predictor of within-plant variation in leaf nitrogen concentration in five species of rain forest pioneer trees (Cecropia obtusifolia, Ficus insipida, Heliocarpus appendiculatus, Piper auritum, and Urera caracasana) from the Los Tuxtlas Biological Station, Veracruz, Mexico. Three hundred seventy-five leaves on 28 plants of the five species were analyzed for leaf nitrogen concentration, leaf mass per area, and leaf light interception at different positions (= nodes) along a shoot. Mean values of leaf nitrogen concentration ranged from 0.697 to 0.993 g/m² in the five species, and varied by as much as 2.24 g/m² among leaves on individual plants. Leaf position on the shoot explained significantly more of the within-plant variation in leaf nitrogen concentration than did leaf light level in four of the five species: Cecropia obtusifolia, Heliocarpus appendiculatus, Piper auritum (branch leaves only), and Urera caracasana. However, individual species differed considerably in the patterns of nitrogen allocation and leaf mass per area among leaves on a shoot. These results suggest that leaf nitrogen deployment in these plants is, in part, developmentally constrained and related to the predictability of canopy light distribution associated with plant growth form.     

Influence of nutrient versus water supply on hydraulic architecture and water balance in Pinus taeda

We investigated the hydraulic consequences of a major decrease in root‐to‐leaf area ratio (A_R:A_L) caused by nutrient amendments to 15‐year‐old Pinus taeda L. stands on sandy soil. In theory, such a reduction in A_R:A_L should compromise the trees’ ability to extract water from drying sand. Under equally high soil moisture, canopy stomatal conductance (G_S) of fertilized trees (F) was 50% that of irrigated/fertilized trees (IF), irrigated trees (I), and untreated control trees (C). As predicted from theory, F trees also decreased their stomatal sensitivity to vapour pressure deficit by 50%. The lower G_S in F was associated with 50% reduction in leaf‐specific hydraulic conductance (K_L) compared with other treatments. The lower K_L in F was in turn a result of a higher leaf area per sapwood area and a lower specific conductivity (conducting efficiency) of the plant and its root xylem. The root xylem of F trees was also 50% more resistant to cavitation than the other treatments. A transport model predicted that the lower A_R:A_L in IF trees resulted in a considerably restricted ability to extract water during drought. However, this deficiency was not exposed because irrigation minimized drought. In contrast, the lower A_R:A_L in F trees caused only a limited restriction in water extraction during drought owing to the more cavitation resistant root xylem in this treatment. In both fertilized treatments, approximate safety margins from predicted hydraulic failure were minimal suggesting increased vulnerability to drought‐induced dieback compared with non‐fertilized trees. However, IF trees are likely to be so affected even under a mild drought if irrigation is withheld.     

Growth and crown efficiency of height repressed lodgepole pine; are suppressed trees more efficient?

Leaf area, crown projection area and growth over the last 5 years were measured to assess growth efficiency (GE) and crown efficiency (CE) of dominant (D), codominant (CD) and suppressed (SP) trees growing in height-repressed (P sites) and normally developing (M sites) lodgepole pine stands. Leaf area index (LAI), hydraulic characteristics, and needle nutrient concentrations were also measured. Volume growth of P site trees between 1994 and 1999 was 46% that of M site trees. Volume growth was closely associated with both hydraulic supply capacity (Q*) and leaf area. Height repression was not associated with lower GE, but P site trees had CE that was 24.5% lower than M site trees. Average GE of D and CD trees was 28% lower than that of SP trees, while mean CE for the D trees was 46% greater than that of CD, and 80% greater than for SP trees. Between M and P sites, canopy LAI and Q* per unit leaf area did not differ. Needle nitrogen (N) concentrations of M site trees were 7.6% greater than for P site trees. SP tree needles had the highest concentration of N and phosphorus. The nutrient advantage enjoyed by SP trees presumably allowed them to maintain higher GE for a given Q*/A_l. The fastest growing trees were the large D and CD trees from M sites. As LAI did not differ between sites, height repression on P sites may be a result of total leaf area being distributed among too many small trees.     

Bundle sheath extensions affect leaf structural and physiological plasticity in response to irradiance

Coordination between structural and physiological traits is key to plants' responses to environmental fluctuations. In heterobaric leaves, bundle sheath extensions (BSEs) increase photosynthetic performance (light‐saturated rates of photosynthesis, A_max) and water transport capacity (leaf hydraulic conductance, K_leaf). However, it is not clear how BSEs affect these and other leaf developmental and physiological parameters in response to environmental conditions. The obscuravenosa (obv) mutation, found in many commercial tomato varieties, leads to absence of BSEs. We examined structural and physiological traits of tomato heterobaric and homobaric (obv) near‐isogenic lines grown at two different irradiance levels. K_leaf, minor vein density, and stomatal pore area index decreased with shading in heterobaric but not in homobaric leaves, which show similarly lower values in both conditions. Homobaric plants, on the other hand, showed increased A_max, leaf intercellular air spaces, and mesophyll surface area exposed to intercellular airspace (S_mes) in comparison with heterobaric plants when both were grown in the shade. BSEs further affected carbon isotope discrimination, a proxy for long‐term water‐use efficiency. BSEs confer plasticity in traits related to leaf structure and function in response to irradiance levels and might act as a hub integrating leaf structure, photosynthetic function, and water supply and demand.