Seedling growth strategies in Bauhinia species: comparing lianas and trees

BACKGROUND AND AIMS: Lianas are expected to differ from trees in their growth strategies. As a result these two groups of woody species will have different spatial distributions: lianas are more common in high light environments. This study determines the differences in growth patterns, biomass allocation and leaf traits in five closely related liana and tree species of the genus Bauhinia. METHODS: Seedlings of two light-demanding lianas (Bauhinia tenuiflora and B. claviflora), one shade-tolerant liana (B. aurea), and two light-demanding trees (B. purpurea and B. monandra) were grown in a shadehouse at 25% of full sunlight. A range of physiological, morphological and biomass parameters at the leaf and whole plant level were compared among these five species. KEY RESULTS: The two light-demanding liana species had higher relative growth rate (RGR), allocated more biomass to leaf production [higher leaf mass fraction (LMF) and higher leaf area ratio (LAR)] and stem mass fraction (SMF), and less biomass to the roots [root mass fraction (RMF)] than the two tree species. The shade-tolerant liana had the lowest RGR of all five species, and had a higher RMF, lower SMF and similar LMF than the two light-demanding liana species. The two light-demanding lianas had lower photosynthetic rates per unit area (A(area)) and similar photosynthetic rates per unit mass (A(mass)) than the trees. Across species, RGR was positively related to SLA, but not to LAR and A(area). CONCLUSIONS: It is concluded that the faster growth of light-demanding lianas compared with light-demanding trees is based on morphological parameters (SLA, LMF and LAR), and cannot be attributed to higher photosynthetic rates at the leaf level. The shade-tolerant liana exhibited a slow-growth strategy, compared with the light-demanding species.     

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.     

Light environment,sapling architecture,and leaf display in six rain forest tree species

Architecture and leaf display were compared in saplings of six rain forest tree species differing in shade tolerance. Saplings were selected along the whole light range encountered in a forest environment. Species differed largely in realized height and crown expansion per unit support biomass, but this could not be related to differences in shade tolerance. The results demonstrate that there exist various solutions to an effective expansion of plant height and crown area. It is argued that choice of the study species and the ontogenetic trajectory regarded determine to a large extent the outcome of interspecific comparisons. No evidence was found that pioneers were characterized by a multilayered and shade tolerants by a monolayered leaf distribution. Yet, sun plants had a similar crown area, a deeper crown, and a higher leaf area index compared to shade plants and their leaves were more evenly distributed along the stem. This suggests that differences in leaf layering are found between plants growing in different light environments, rather than between species differing in shade tolerance.     

Light capture efficiency decreases with increasing tree age and size in the southern hemisphere gymnosperm Agathis australis

We investigated leaf and shoot architecture in relation to growth irradiance (Q_int) in young and mature trees of a New Zealand native gymnosperm Agathis australis (D. Don) Lindl. to determine tree size-dependent and age-dependent controls on light interception efficiency. A binomial 3-D turbid medium model was constructed to distinguish between differences in shoot light interception efficiency due to variations in leaf area density, angular distribution and leaf aggregation. Because of the positive effect of light on leaf dry mass per area (M_A), nitrogen content per area (N_A) increased with increasing irradiance in both young and mature trees. At a common irradiance, N_A, M_A and the components of M_A, density and thickness, were larger in mature trees, indicating a greater accumulation of photosynthetic biomass per unit area, but also a larger fraction of support biomass in older trees. In both young and mature trees, shoot inclination angle relative to horizontal, and leaf number per unit stem length decreased, and silhouette to total leaf area ratio (S_S) increased with decreasing irradiance, demonstrating more efficient light harvesting in low light. The shoots of young trees were more horizontal and less densely leafed with a larger S_S than those of mature trees, signifying greater light interception efficiency in young plants. Superior light harvesting in young trees resulted from more planar leaf arrangement and less clumped foliage. These results suggest that the age-dependent and/or size-dependent decreases in stand productivity may partly result from reduced light interception efficiency in larger mature relative to smaller and younger plants.     

Leaf structural and photosynthetic characteristics,and biomass allocation to foliage in relation to foliar nitrogen content and tree size in three Betula species

Young trees 0.03-1.7 m high of three coexisting Betula species were investigated in four sites of varying soil fertility, but all in full daylight, to separate nutrient and plant size controls on leaf dry mass per unit area (MA), light-saturated foliar photosynthetic electron transport rate (J) and the fraction of plant biomass in foliage (F(L)). Because the site effect was generally non-significant in the analyses of variance with foliar nitrogen content per unit dry mass (N(M)) as a covariate, N(M) was used as an explaining variable of leaf structural and physiological characteristics. Average leaf area (S) and dry mass per leaf scaled positively with N(M) and total tree height (H) in all species. Leaf dry mass per unit area also increased with increasing H, but decreased with increasing N(M), whereas the effects were species-specific. Increases in plant size led to a lower and increases in N(M) to a greater FL and total plant foliar area per unit plant biomass (LAR). Thus, the self-shading probably increased with increasing N(M) and decreased with increasing H. Nevertheless, the whole-plant average M(A), as well as M(A) values of topmost fully exposed leaves, correlated with N(M) and H in a similar manner, indicating that scaling of MA with N(M) and H did not necessarily result from the modified degree of within-plant shading. The rate of photosynthetic electron transport per unit dry mass (J(M)) scaled positively with N(M), but decreased with increasing H and M(A). Thus, increases in M(A) with tree height and decreasing nitrogen content not only resulted in a lower plant foliar area (LAR = F(L)/M(A)), but also led to lower physiological activity of unit foliar biomass. The leaf parameters (J(M), N(M) and M(A)) varied threefold, but the whole-plant characteristic FL varied 20-fold and LAR 30-fold, indicating that the biomass allocation was more plastically adjusted to different plant internal nitrogen contents and to tree height than the foliar variables. Our results demonstrate that: (1) tree height and N(M) may independently control foliar structure and physiology, and have an even greater impact on biomass allocation; and (2) the modified within-plant light availabilities alone do not explain the observed patterns. Although there were interspecific differences with respect to the statistical significance of the relationships, all species generally fit common regressions. However, these differences were consistent, and suggested that more competitive species with inherently larger growth rates also more plastically respond to N and H.     

LEAF DISPLAY,CANOPY STRUCTURE,AND LIGHT INTERCEPTION OF TWO UNDERSTORY PALM SPECIES

The implications of leaf size, leaf display, and crown size for whole-plant light interception were investigated in Geonoma cuneata and Asterogyne martiana, two understory palm species native to Central American rain forests. Adults of A. martiana had longer leaves, more leaves per plant, and greater total leaf area than G. cuneata adults. Geometric measurements within whole crowns were used to calculate light interception efficiency, a leaf-based measure of the proportion of total incident light that is intercepted by a crown. Light interception efficiency was higher in adult G. cuneata than in adult A. martiana; seedlings of the two species did not differ significantly in light interception efficiency. Decreased efficiency of adult A. martiana crowns was largely due to an increased proportion of pendent leaves. Compared to G. cuneata, adults of A. martiana had greater light interception capacity (the product of light interception efficiency and total leaf area), but they also had higher biomass costs of light interception. Lower biomass costs of light interception in adult G. cuneata enable this species to exploit successfully the most deeply shaded microsites in the rain forest understory.     

Simple equations to estimate light interception by isolated trees from canopy structure features: assessment with three-dimensional digitized apple trees

* Simple models of light interception are useful to identify the key structural parameters involved in light capture. We developed such models for isolated trees and tested them with virtual experiments. Light interception was decomposed into the projection of the crown envelope and the crown porosity. The latter was related to tree structure parameters. * Virtual experiments were conducted with three-dimensional (3-D) digitized apple trees grown in Lebanon and Switzerland, with different cultivars and training. The digitized trees allowed actual values of canopy structure (total leaf area, crown volume, foliage inclination angle, variance of leaf area density) and light interception properties (projected leaf area, silhouette to total area ratio, porosity, dispersion parameters) to be computed, and relationships between structure and interception variables to be derived. * The projected envelope area was related to crown volume with a power function of exponent 2/3. Crown porosity was a negative exponential function of mean optical density, that is, the ratio between total leaf area and the projected envelope area. The leaf dispersion parameter was a negative linear function of the relative variance of leaf area density in the crown volume. * The resulting models were expressed as two single equations. After calibration, model outputs were very close to values computed from the 3-D digitized databases.     

Mineral nutrition and growth of tropical maize as affected by soil acidity

Soil constraints linked to low pH reduce grain yield in about 10% of the maize growing area in tropical developing countries. The aim of this research was to elucidate the reasons for this maize yield reduction on an oxisol of Guadeloupe. The field experiment had two treatments: the native non-limed soil (NLI, pH 4.5, 2.1 cmol Al kg^–1, corresponding to 20% Al saturation), and the same soil limed 6 years prior to the experiment (LI, pH 5.3, 0 cmol Al kg^–1). The soils were fertilized with P and N. The above-ground biomass, root biomass at flowering, grain yield and yield components, leaf area index (LAI), light interception, radiation-use-efficiency (RUE), P and N uptake, soil water storage, and soil mineral N were measured during the maize cycle. The allometric relationships between shoot N concentration, LAI and above-ground biomass in LI were similar to those reported for maize cropped in temperate regions, indicating that these relationships are also useful to describe maize growth on tropical soils without Al toxicity. In NLI, soil acidity severely affected leaf appearance, leaf size and consequently the LAI, which was reduced by 60% at flowering, although the RUE was not affected. Therefore, the reduction in the above-ground biomass (30% at flowering) and grain yield (47%) were due to the lower LAI and light interception. At flowering, the root/shoot ratio was 0.25 in NLI and 0.17 in LI, and the root biomass in NLI was reduced by 64% compared to LI. Nitrogen uptake was also reduced in NLI in spite of high soil N availability. Nevertheless, shoot N concentration vs aboveground biomass showed a typical decline in both treatments. In NLI, the shoot P concentration vs above-ground biomass relationship showed an increase in the early stages, indicating that P uptake and root-shoot competition for the absorbed P in the early plant stages controlled the establishment and the development of the leaf area.     

Contrasting leaf chemical traits in tropical lianas and trees: implications for future forest composition

Lianas are an important growthform in tropical forests, and liana abundance and biomass may be increasing in some regions. Explanations for liana proliferation hinge upon physiological responses to changing resource conditions that would favour them over trees. Testing a chemical basis for such responses, we assessed 22 foliar traits in 778 lianas and 6496 trees at 48 tropical forest sites. Growthform differences in chemical allocation occurred on a leaf mass and area basis. Light capture-growth and maintenance-metabolism chemicals averaged 14.5 and 16.7% higher mass-based concentration in lianas than in trees globally, whereas structure and defence chemicals averaged 9.0% lower in lianas. Relative differences in chemical allocation by lianas and trees were mediated by climate with peak differences at about 2500 mm year(-1) and 25 °C. Differences in chemical traits suggest that liana expansion could be greatest in forests undergoing increased canopy-level irradiance via disturbance and climate change.     

Ontogeny, understorey light interception and simulated carbon gain of juvenile rainforest evergreens differing in shade tolerance

Background and Aims

A long-running debate centres on whether shade tolerance of tree seedlings is mainly a function of traits maximizing net carbon gain in low light, or of traits minimizing carbon loss. To test these alternatives, leaf display, light-interception efficiency, and simulated net daily carbon gain of juvenile temperate evergreens of differing shade tolerance were measured, and how these variables are influenced by ontogeny was queried.

Methods

The biomass distribution of juveniles (17–740 mm tall) of seven temperate rainforest evergreens growing in low (approx. 4 %) light in the understorey of a second-growth stand was quantified. Daytime and night-time gas exchange rates of leaves were also determined, and crown architecture was recorded digitally. YPLANT was used to model light interception and carbon gain.

Results

An index of species shade tolerance correlated closely with photosynthetic capacities and respiration rates per unit mass of leaves, but only weakly with respiration per unit area. Accumulation of many leaf cohorts by shade-tolerant species meant that their ratios of foliage area to biomass (LAR) decreased more gradually with ontogeny than those of light-demanders, but also increased self-shading; this depressed the foliage silhouette-to-area ratio (STAR), which was used as an index of light-interception efficiency. As a result, displayed leaf area ratio (LAR_d = LAR × STAR) of large seedlings was not related to species shade tolerance. Self-shading also caused simulated net daily carbon assimilation rates of shade-tolerant species to decrease with ontogeny, leading to a negative correlation of shade tolerance with net daily carbon gain of large (500 mm tall) seedlings in the understorey.

Conclusions

The results suggest that efficiency of energy capture is not an important correlate of shade tolerance in temperate rainforest evergreens. Ontogenetic increases in self-shading largely nullify the potential carbon gain advantages expected to result from low respiration rates and long leaf lifespans in shade-tolerant evergreens. The main advantage of their long-lived leaves is probably in reducing the costs of crown maintenance.     

Optimisation of spatial allocation patterns in lianas compared to trees used for support

There are only limited possibilities to study the competition between trees and lianas in the top canopy of tropical rain forests. Furthermore, the important question how the leaf traits are related to twig support is rarely studied, especially regarding growing space partitioning between the self-supporting and the climbing growth form. Our study used the hot-air balloon within the “Operation Canopee” in the Masoala National Park, Madagascar, to test the differences in spatial allocation patterns of leaves and twigs in lianas and tree parts used for support. The sampling design emphasised to obtain a common assembly of twigs and leaves from both, trees and lianas. The results from the top canopy were compared to the data from the understorey regarding biomass and nutrients in leaves and distal twigs. In the understorey the reduction in structural investment was much stronger in lianas than in trees. The results showed that lianas reduced carbon investment per volume, but increased leaf nitrogen concentration and leaf area ratio (LAR), the latter driven by a reduction in leaf mass per area (LMA). In the top canopy, lianas contributed about one third of the leaf area density of 3 m² m⁻³. For distal twigs, no relationship was found between twig biomass per volume and leaf area density for trees, but lianas balanced both structural parameters closely. The climbers benefit from the external support provided by the trees and optimise the area of assimilation tissue at low per volume investment for mechanical stability. Several traits such as low LMA and high leaf nitrogen concentrations together with higher LAR and optimised spatial investment advantage the climbing growth form and enable a fast acquisition of growing space. The results emphasize the necessity to consider spatial and structural features of growing space acquisition when dealing with plant competition. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The expression of light-related leaf functional traits depends on the location of individual leaves within the crown of isolated Olea europaea trees

Background The spatial arrangement and expression of foliar syndromes within tree crowns can reflect the coupling between crown form and function in a given environment. Isolated trees subjected to high irradiance and concomitant stress may adjust leaf phenotypes to cope with environmental gradients that are heterogeneous in space and time within the tree crown. The distinct expression of leaf phenotypes among crown positions could lead to complementary patterns in light interception at the crown scale.Methods We quantified eight light-related leaf traits across 12 crown positions of ten isolated Olea europaea trees in the field. Specifically, we investigated whether the phenotypic expression of foliar traits differed among crown sectors and layers and five periods of the day from sunrise to sunset. We investigated the consequences in terms of the exposed area of the leaves at the tree scale during a single day.Key Results All traits differed among crown positions except the length-to-width ratio of the leaves. We found a strong complementarity in the patterns of the potential exposed area of the leaves among day periods as a result of a non-random distribution of leaf angles across the crown. Leaf exposure at the outer layer was below 60 % of the displayed surface, reaching maximum interception during morning periods. Daily interception increased towards the inner layer, achieving consecutive maximization from east to west positions within the crown, matching the sun’s trajectory.Conclusions The expression of leaf traits within isolated trees of O. europaea varies continuously through the crown in a gradient of leaf morphotypes and leaf angles depending on the exposure and location of individual leaves. The distribution of light-related traits within the crown and the complementarity in the potential exposure patterns of the leaves during the day challenges the assumption of low trait variability within individuals.     

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.     

Scaling‐up from leaf to canopy‐aggregate properties in sclerophyll shrub species

Abstract: Plant species vary widely in their average leaf lifespan (LL) and specific leaf area (SLA, leaf area per dry mass). The negative LL–SLA relationship commonly seen among species represents an important evolutionary trade‐off, with higher SLA indicating greater potential for fast growth (higher rate of return on a given investment), but longer LL indicating a longer duration of the revenue stream from that investment. We investigated how these leaf‐economic traits related to aggregate properties of the plant crown. Across 14 Australian sclerophyll shrub species, those with long LL accumulated more leaf mass and leaf area per unit ground area. Light attenuation through their canopies was more severe. Leaf accumulation and light attenuation were more weakly related to SLA than to LL. The greater accumulation of foliage in species with longer LL and lower SLA may counterbalance their generally lower photosynthetic rates and light‐capture areas per gram of leaf.     

Phenotypic correlates of the lianescent growth form: a review

Background

As proposed by Darwin, climbers have been assumed to allocate a smaller fraction of biomass to support organs in comparison with self-supporting plants. They have also been hypothesized to possess a set of traits associated with fast growth, resource uptake and high productivity.

Scope

In this review, these hypotheses are evaluated by assembling and synthesizing published and unpublished data sets from across the globe concerning resource allocation, growth rates and traits of leaves, stems and roots of climbers and self-supporting species.

Conclusions

The majority of studies offer little support for the smaller allocation of biomass to stems or greater relative growth rates in climbers; however, these results are based on small sized (<1 kg) plants. Simulations based on allometric biomass equations demonstrate, however, that larger lianas allocate a greater fraction of above-ground biomass to leaves (and therefore less biomass to stems) compared with similar sized trees. A survey of leaf traits of lianas revealed their lower average leaf mass per area (LMA), higher N and P concentration and a slightly higher mass-based photosynthetic rate, as well as a lower concentration of phenolic-based compounds than in woody self-supporting species, consistent with the specialization of lianas towards the fast metabolism/rapid turnover end of the global trait spectra. Liana stems have an efficient hydraulic design and unique mechanical features, while roots appear to penetrate deeper soil levels than in trees and are often able to generate hydraulic pressure. Much remains to be learned, however, about these and other functional specializations of their axial organs and the associated trade-offs. Developmental switches between self-supporting, searcher and climbing shoots within the same individual are a promising field of comparative studies on trait association in lianas. Finally, some of the vast trait variability within lianas may be reduced when species with different climbing mechanisms are considered separately, and when phylogenetic conservatism is accounted for.     

Seasonal differences in leaf-level physiology give lianas a competitive advantage over trees in a tropical seasonal forest

Lianas are an important component of most tropical forests, where they vary in abundance from high in seasonal forests to low in aseasonal forests. We tested the hypothesis that the physiological ability of lianas to fix carbon (and thus grow) during seasonal drought may confer a distinct advantage in seasonal tropical forests, which may explain pan-tropical liana distributions. We compared a range of leaf-level physiological attributes of 18 co-occurring liana and 16 tree species during the wet and dry seasons in a tropical seasonal forest in Xishuangbanna, China. We found that, during the wet season, lianas had significantly higher CO₂ assimilation per unit mass (A _mass), nitrogen concentration (N _mass), and δ¹³C values, and lower leaf mass per unit area (LMA) than trees, indicating that lianas have higher assimilation rates per unit leaf mass and higher integrated water-use efficiency (WUE), but lower leaf structural investments. Seasonal variation in CO₂ assimilation per unit area (A _area), phosphorus concentration per unit mass (P _mass), and photosynthetic N-use efficiency (PNUE), however, was significantly lower in lianas than in trees. For instance, mean tree A _area decreased by 30.1% from wet to dry season, compared with only 12.8% for lianas. In contrast, from the wet to dry season mean liana δ¹³C increased four times more than tree δ¹³C, with no reduction in PNUE, whereas trees had a significant reduction in PNUE. Lianas had higher A _mass than trees throughout the year, regardless of season. Collectively, our findings indicate that lianas fix more carbon and use water and nitrogen more efficiently than trees, particularly during seasonal drought, which may confer a competitive advantage to lianas during the dry season, and thus may explain their high relative abundance in seasonal tropical forests.     

Crown development in a pioneer tree, Rhus trichocarpa, in relation to the structure and growth of individual branches

Based on an allometric reconstruction, the structure and biomass-allocation patterns of branches and current-year shoots were investigated in branches of various heights in the pioneer tree Rhus trichocarpa, to evaluate how crown development is achieved and limited in association with height. Path analysis was conducted to explore the effects of light availability, basal height and size of individual branches on branch structure and growth. Branch angle was affected by basal height, whereas branch mass was influenced primarily by light availability. This result suggests that branch structure is strongly constrained by basal height, and that trees mediate such constraints under different light environments. Previous-year leaf area and light availability showed positive effects on current-year stem mass. In contrast, branch basal height and mass negatively affected current-year stem mass. Moreover, the length of stems of a given diameter decreased with increasing branch height. Therefore the cost of biomass investment for a unit growth in length is greater for branches of larger size and at upper positions. Vertical growth rate in length decreased with increasing height. Height-dependent changes in stem allometry and angle influenced the reduction in vertical growth rate to a similar degree.     

Physiological basis of spacing effects on tree growth and form in Eucalyptus globulus

We examine the effects of spacing and layout on the growth and form of 3- to 4-year-old Eucalyptus globulus in a farm forestry context. Four planting layouts were chosen. These represented the range commonly in use in farm forestry: block plantings (2Ǹ m), triple rows (2Ǹ m) at 10-m intervals, single rows (2᎒ m) and isolated trees (10᎒ m). The physiological significance of key results is interpreted in terms of changes in the parameters of a simple plantation growth model. Under conditions where levels of direct light are high, for example during summer, block-planted trees intercepted only 38% of the light intercepted by isolated trees. On a stand basis, however, the combination of incident radiation and ground coverage declined with lower stand densities. While stand leaf area index declined from around 6 to 1 with increased spacing, individual tree leaf areas rose from around 50 m² in block plantings to 150 m² in isolated trees. The proportion of above-ground biomass found in stems declined with increasing spacing as the mass in foliage and branches increased. Stems accounted for 65% of above-ground biomass in block-planted trees but only 35% in isolated trees. The contributions of leaves and branches correspondingly rose from 19% to 35% and from 16% to 29%, respectively. Changes in biomass distribution were accompanied by increasing branch number, branch thickness, flatter branch angles and the longer retention of lower branches with greater spacing. These changes have implications for the merchantability of the timber. The efficiency of above-ground radiation conversion was constant at 0.67 g MJ^-1 irrespective of spacing. We estimated that foliar maintenance respiration (R_m) accounted for about 90% of above-ground R_m. On a stand basis R_m costs block plantings 23.90 t DM ha^-1 year^-1 (50% annual above-ground photosynthetic production) compared with 6.22 t DM ha^-1 year^-1 (40% annual above-ground photosynthetic production) in stands of isolated trees.     

Seedlings of temperate rainforest conifer and angiosperm trees differ in leaf area display

Background and Aims

The contemporary relegation of conifers mainly to cold or infertile sites has been ascribed to low competitive ability, as a result of the hydraulic inefficiency of tracheids and their seedlings'' initial dependence on small foliage areas. Here it is hypothesized that, in temperate rainforests, the larger leaves of angiosperms also reduce self-shading and thus enable display of larger effective foliage areas than the numerous small leaves of conifers.

Methods

This hypothesis was tested using 3-D modelling of plant architecture and structural equation modelling to compare self-shading and light interception potential of seedlings of six conifers and 12 angiosperm trees from temperate rainforests. The ratio of displayed leaf area to plant mass (LAR_d) was used to indicate plant light interception potential: LAR_d is the product of specific leaf area, leaf mass fraction, self-shading and leaf angle.

Results

Angiosperm seedlings self-shaded less than conifers, mainly because of differences in leaf number (more than leaf size), and on average their LAR_d was about twice that of conifers. Although specific leaf area was the most pervasive influence on LAR_d, differences in self-shading also significantly influenced LAR_d of large seedlings.

Conclusions

The ability to deploy foliage in relatively few, large leaves is advantageous in minimizing self-shading and enhancing seedling light interception potential per unit of plant biomass. This study adds significantly to evidence that vegetative traits may be at least as important as reproductive innovations in explaining the success of angiosperms in productive environments where vegetation is structured by light competition.     

Increase in light interception cost and metabolic mass component of leaves are coupled for efficient resource use in the high altitude vegetation

Global syntheses of leaf trait scaling relationships report an increase in light interception costs or ‘diminishing returns’ with increase in leaf area. However, variation in light interception costs across ecological gradients and plant strategies to cope up with these costs are not adequately understood. We analyzed leaf area (A) – leaf dry mass (M), leaf water mass (W) – M and W – A scaling relationships in plants occurring in a high altitude region of western Himalaya across environmental gradients to understand changes in light interception cost and metabolic mass component. M represents light interception cost, whereas, W is considered as a proxy of metabolic mass component for liquid phase being the ultimate source of metabolic activity. Trait values were measured from 9278 leaves belonging to 136 dominant species occurring at different sites, slope aspects, elevations and habitat types. Overall, light interception cost increased with increasing A (scaling exponent (α) < 1 in A–M relationship) and metabolic mass component increased disproportionately high with increasing M and A. We found significant differences in scaling exponents of leaf trait relationship between sites, elevations, slope aspects and habitat types, indicating that increase in light interception cost was more evident at higher elevations, southern slopes and open habitats. Further, with increase in light interception cost, metabolic mass component also increased (α > 1 in W–M and W–A relationships). The changes in scaling exponents of various leaf trait relationships across ecological gradients indicated that vegetation of different regions have differences in light interception cost and metabolic mass component. Moreover, increasing light interception cost (increase in mechanical and hydraulic tissues) with increasing A and increasing metabolic mass (leaf thickness) with increasing A and M are favored in high altitude vegetation. This could be a key strategy of high altitude plants for efficient resource capture and use in harsh environments.