Decline of photosynthetic capacity with leaf age and position in two tropical pioneer tree species

The effect of leaf age on photosynthetic capacity, a critical parameter in the theory of optimal leaf longevity, was studied for two tropical pioneer tree species, Cecropia longipes and Urera caracasana, in a seasonally dry forest in Panama. These species continuously produce short-lived leaves (74 and 93 d, respectively) during the rainy season (May-December) on orthotropic branches. However, they differ in leaf production rate, maximum number of leaves per branch, light environment experienced by the leaves, leaf mass per unit area, and nitrogen content. Light-saturated photosynthetic rates for marked leaves of known ages (±1 wk) were measured with two contrasting schemes (repeated measurements vs. chronosequence within branch), which overall produced similar results. In both species, photosynthetic rates and nitrogen use efficiency were negatively correlated with leaf age and positively correlated with light availability. Photosynthetic rates declined faster with leaf age in Cecropia than in Urera as predicted by the theory. The rate of decline was faster for leaves on branches with faster leaf turnover rates. Nitrogen per unit leaf area decreased with leaf age only for Urera. Leaf mass per unit area increased with leaf age, either partly (in Cecropia) or entirely (in Urera) due to ash accumulation.     

Linking leaf and root trait syndromes among 39 grassland and savannah species

Here, we tested hypothesized relationships among leaf and fine root traits of grass, forb, legume, and woody plant species of a savannah community. CO2 exchange rates, structural traits, chemistry, and longevity were measured in tissues of 39 species grown in long-term monocultures. Across species, respiration rates of leaves and fine roots exhibited a common regression relationship with tissue nitrogen (N) concentration, although legumes had lower rates at comparable N concentrations. Respiration rates and N concentration declined with increasing longevity of leaves and roots. Species rankings of leaf and fine-root N and longevity were correlated, but not specific leaf area and specific root length. The C3 and C4 grasses had lower N concentrations than forbs and legumes, but higher photosynthesis rates across a similar range of leaf N. Despite contrasting photosynthetic pathways and N2-fixing ability among these species, concordance in above- and below-ground traits was evident in comparable rankings in leaf and root longevity, N and respiration rates, which is evidence of a common leaf and root trait syndrome linking traits to effects on plant and ecosystem processes.     

Comparison of leaf life span, photosynthesis and defensive traits across seven species of deciduous broad-leaf tree seedlings

BACKGROUND AND AIMS: Leaf life span, photosynthetic parameters and defensive traits were compared across seven species of deciduous broad-leaved tree seedlings native to northern Japan to test the "cost-benefit hypothesis" that more productive leaves are more susceptible to herbivore attack than less productive leaves. METHODS: Studies were made on three early successional species, Alnus hirsuta, Betula maximowicziana and Betula platyphylla "japonica"; one mid-successional species, Ostrya japonica, and three late-successional species, Carpinus cordata, Quercus mongolica 'grosseserrata' and Acer mono. Photosynthetic parameters and defensive traits (total phenolics, condensed tannin and toughness) of leaves were measured for each species, and a bioassay test with Eri silkmoth larvae (Samia cynthia ricini) was undertaken to evaluate differences between species in susceptibility to herbivore attack. KEY RESULTS: Early successional species have a shorter leaf life span (62-88 d) than late successional species (155-187 d). Leaf nitrogen content and light-saturated photosynthetic rate per unit leaf area (P(sat)-area) and per unit leaf mass (P(sat)-mass) were negatively correlated with leaf life span. The nitrogen content of early successional species was about 30 mg g(-1) and that of late successional species was about 16 mg g(-1). Leaf toughness and the C/N ratio were positively correlated with leaf life span, although condensed tannin was not correlated with leaf life span. The bioassay test showed that the number of days the larvae survived was negatively correlated with leaf life span. Average survival of larvae feeding on leaves of A. hirsuta, which has the shortest leaf life span, was 14.4 d and that of Q. mongolica, which has the longest leaf life span, was 6.6 d. The number of days of larval survival was positively correlated with leaf nitrogen content. There was no correlation between days of larval survival and defensive traits. CONCLUSIONS: These results indicate that species with a shorter leaf life span have higher photosynthetic productivity and are more susceptible to herbivore attack than species with a longer leaf life span. This supports the "cost-benefit hypothesis".     

Effects of leaf emergence on leaf lifespan are independent of life form and successional status

Abstract The longevity of a leaf is related to the benefit that the plant is able to derive from it. This benefit varies among seasons and as more leaves emerge, such that leaf lifespan can be limited by canopy position rather than physiological age. Using interval‐censored failure time analysis, we investigate leaf lifespan for 34 Mediterranean species in a previously published dataset involving species with different life forms and functional strategies. Failure time regression models were used to determine leaf lifespan, and to investigate how these effects varied among species. Median lifespan estimated for each species with two methods differed by less than 10% on average, but varied from 0.02–19.5% depending on the shape of the underlying failure time distribution. Within shoots, later‐emerging leaves had shorter lifespans for species with longer periods of leaf emergence, and the reverse was true for species with short emergence. Having accounted for the within‐shoot effect, leaves emerging in spring had shorter lifespans, particularly herbaceous species, whereas the reverse was true woody species. These effects were consistent among life forms and successional stages, and consistent with theories of within‐shoot translocation of resources following self‐shading.     

Relationships between photosynthetic activity and silica accumulation with ages of leaf in Sasa veitchii (Poaceae, Bambusoideae)

BACKGROUND AND AIMS: Bamboos have long-lived, evergreen leaves that continue to accumulate silica throughout their life. Silica accumulation has been suggested to suppress their photosynthetic activity. However, nitrogen content per unit leaf area (N(area)), an important determinant of maximum photosynthetic capacity per unit leaf area (P(max)), decreases as leaves age and senescence. In many species, P(max) decreases in parallel with the leaf nitrogen content. It is hypothesized that if silica accumulation affects photosynthesis, then P(max) would decrease faster than N(area), leading to a decrease in photosynthetic rate per unit leaf nitrogen (photosynthetic nitrogen use efficiency, PNUE) with increasing silica content in leaves. METHODS: The hypothesis was tested in leaves of Sasa veitchii, which have a life span of 2 years and accumulate silica up to 41 % of dry mass. Seasonal changes in P(max), stomatal conductance, N(area) and silica content were measured for leaves of different ages. KEY RESULTS: Although P(max) and PNUE were negatively related with silica content across leaves of different ages, the relationship between PNUE and silica differed depending on leaf age. In second-year leaves, PNUE was almost constant although there was a large increase in silica content, suggesting that leaf nitrogen was a primary factor determining the variation in P(max) and that silica accumulation did not affect photosynthesis. PNUE was strongly and negatively correlated with silica content in third-year leaves, suggesting that silica accumulation affected photosynthesis of older leaves. CONCLUSIONS: Silica accumulation in long-lived leaves of bamboo did not affect photosynthesis when the silica concentration of a leaf was less than 25 % of dry mass. Silica may be actively transported to epidermal cells rather than chlorenchyma cells, avoiding inhibition of CO2 diffusion from the intercellular space to chloroplasts. However, in older leaves with a larger silica content, silica was also deposited in chlorenchyma cells, which may relate to the decrease in PNUE.     

Specific leaf area and leaf nitrogen concentration in annual and perennial grass species growing in Mediterranean old-fields

 We evaluated the hypothesis that photosynthetic traits differ between leaves produced at the beginning (May) and the end (November–December) of the rainy season in the canopy of a seasonally dry forest in Panama. Leaves produced at the end of the wet season were predicted to have higher photosynthetic capacities and higher water-use efficiencies than leaves produced during the early rainy season. Such seasonal phenotypic differentiation may be adaptive, since leaves produced immediately preceding the dry season are likely to experience greater light availability during their lifetime due to reduced cloud cover during the dry season. We used a construction crane for access to the upper canopy and sampled 1- to 2-month-old leaves marked in monthly censuses for six common tree species with various ecological habits and leaf phenologies. Photosynthetic capacity was quantified as light- and CO₂-saturated oxygen evolution rates with a leaf-disk oxygen electrode in the laboratory (O_2max) and as light-saturated CO₂ assimilation rates of intact leaves under ambient CO₂ (A_max). In four species, pre-dry season leaves had significantly higher leaf mass per unit area. In these four species, O_2max and A_max per unit area and maximum stomatal conductances were significantly greater in pre-dry season leaves than in early wet season leaves. In two species, A_max for a given stomatal conductance was greater in pre-dry season leaves than in early wet season leaves, suggesting a higher photosynthetic water-use efficiency in the former. Photosynthetic capacity per unit mass was not significantly different between seasons of leaf production in any species. In both early wet season and pre-dry season leaves, mean photosynthetic capacity per unit mass was positively correlated with nitrogen content per unit mass both within and among species. Seasonal phenotypic differentiation observed in canopy tree species is achieved through changes in leaf mass per unit area and increased maximum stomatal conductance rather than by changes in nitrogen allocation patterns. Received: 7 March 1996 / Accepted: 1 August 1996     

Leaf dynamics,self-shading and carbon gain in seedlings of a tropical pioneer tree

We examined leaf dynamics and leaf age gradients of photosynthetic capacity and nitrogen concentration in seedlings of the tropical pioneer tree, Heliocarpus appendiculatus, grown in a factorial design under controlled conditions with two levels each of nutrients, ambient light (light levels incident above the canopy), and self-shading (the gradient of light levels from upper to lower leaves on the shoot). Correlations among these parameters were examined in order to determine the influence of self-shading, and the regulation of standing leaf numbers, on leaf longevity and its association with leaf photosynthetic capacity. Leaf longevity and the number of leaves on the main shoot were both reduced in high light, while in the low light environment, they were reduced in the steeper self-shading gradient. In high nutrients, leaf longevity was reduced whereas leaf number increased. Leaf initiation rates were higher in the high nutrient treatment but were not influenced by either light treatment. Maximum-light saturated photosynthetic rate, on an area basis, was greater in the high light and nutrient treatments, while the decline in photosynthetic capacity in realtion to leaf position on the shoot was more rapid in high light and in low nutrients. Leaf longevity was negatively correlated among treatments with initial photosynthetic capacity. The leaf position at which photosynthetic capacity was predicted to reach zero was positively correlated with the number of leaves on the shoot, supporting the hypothesis that leaf numbers are regulated by patterns of self-shading. The negative association of longevity and initial photosynthetic capacity apparently arises from different associations among gradients of photosynthetic capacity, leaf numbers and leaf initiation rates in relation to light and nutrient availability. The simultaneous consideration of age and position of leaves illuminates the role of self-shading as an important factor influencing leaf senescence and canopy structure and dynamics.     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

Annual photosynthetic activities of temperate evergreen and deciduous broadleaf tree species with simultaneous and successive leaf emergence in response to altitudinal air temperature

To explain why the composition of evergreen and deciduous forests changes along air temperature gradients, we measured several traits of single leaves from temperate deciduous and evergreen broadleaf trees with simultaneous and successive leaf emergence growing at different altitudes in the field. The parameters included seasonal net photosynthetic rate, longevity, mass per area, nitrogen content, and photosynthetic nitrogen-use efficiency. With decreasing altitude, the leaf longevity of deciduous broadleaf trees increased, whereas the maximum net photosynthetic rate decreased. In contrast, leaf longevity of evergreen broadleaf trees decreased, whereas the minimum net photosynthetic rate in winter increased. Along the air temperature gradient, the annual production of deciduous trees with simultaneous leaf emergence may be constant, because the integrated lifetime net photosynthetic rate (ILNPR) of a single leaf changed little. In comparison, deciduous trees with successive leaf emergence may show enhanced annual production with increasing air temperature, by increasing the total leaf number per branch and tree under an extended growing season. Temperate evergreen broadleaf tree species may also show increased annual production with increasing air temperature by sufficiently raising the number of the first-year leaves to the total leaves of branch and tree, which is accelerated by raising the integrated first-year net photosynthetic rate of the single leaf, despite little change in the ILNPR. With increasing air temperature from cool-temperate to warm-temperate zones, evergreen broadleaf tree species gain an advantage of the annual production over deciduous broadleaf tree species with simultaneous leaf emergence.     

Significance of leaf longevity in plants

The cost–benefit model of leaf life span is extended by incorporating the cost of construction and maintenance of supporting structures, such as shoots and roots. Published data on leaf longevities were reviewed, and they support qualitative predictions of optimal leaf life span based on the present model. In relation to life form, leaf life span increases in the following sequence, due to increasing costs of support structures: floating leaves of aquatic plants < annual herbs < perennial herbs < deciduous trees. Within species, leaf life span increases with plant size as support costs per unit of leaf increase. This prediction is supported by comparisons of seedlings and adults of temperate trees. These results suggest that the construction and maintenance costs of supporting structures significantly influence life span of individual leaves. Leaf longevity, in turn, connects the ecophysiology of the individual leaf to growth of the individual plant and energy and matter cycling at the ecosystem level.     

Contrasting patterns of photosynthetic acclimation and photoinhibition in two evergreen herbs from a winter deciduous forest

The relationship between the microclimate within an Oak-Hickory forest and photosynthetic characters of two resident evergreen herbs with contrasting leaf phenologies was investigated on a monthly basis for 1 full year. Heuchera americana has leaf flushes in the spring and fall, with average leaf life spans of 6–7 months. Hexastylis arifolia produces a single cohort of leaves each spring with a leaf life span of 12–13 months. We predicted that among evergreen plants inhabiting a seasonal habitat, a species for which the frequency of leaf turnover is greater than the frequency of seasonal extremes would have a greater annual range in photosynthetic capacity than a species that only produced a single flush of leaves during the year. Photosynthetic parameters, including apparent quantum yield, maximum photosynthetic capacity (P_max), temperature of maximum photosynthesis, photochemical efficiency of PSII and leaf nitrogen (N) and chlorophyll concentrations, were periodically measured under laboratory conditions in leaves sampled from natural populations of both species. Mature leaves of both species acclimated to changing understory conditions with the mean seasonal differences being significantly greater for Heuchera than for Hexastylis. Area based maximum photosynthetic rates at 25°C were approximately 250% and 100% greater in winter leaves than summer leaves for Heuchera and Hexastylis respectively. Nitrogen concentrations were highest in winter leaves. Chlorophyll concentrations were highest in summer leaves. Low P_max/N values for these species suggest preferential allocation of leaf nitrogen into non-photosynthetic pools and/or light-harvesting function at the expense of photosynthetic enzymes and electron transport components. Despite the increase in photosynthetic capacity, there was evidence of chronic winter photoinhibition in Hexastylis, but not in Heuchera. Among these ecologically similar species, there appears to be a trade-off between the frequency of leaf production and the balance of photosynthetic acclimation and photoinhibition.     

Nitrogen resorption and photosynthetic activity over leaf life span in an evergreen shrub, Rhododendron ferrugineum, in a subalpine environment

Here, the advantages for a shrub of having long vs short-lived leaves was investigated in Rhododendron ferrugineum by following nitrogen(15N) and carbon(14C) resorption and translocation, and photosynthetic capacity over the life span. Mean leaf life span was 19 months. Nitrogen (N) resorption in attached leaves occurred mainly in the first year (23%) and reached a maximum of 31% in the second. Although, resorption was similar in attached and fallen 1-yr-old leaves, it was on average one-third higher in fallen than in attached older leaves. Final N resorption of a leaf compartment reached 41%, half occurring from healthy leaves during the first year. Photosynthetic capacity decreased slightly during the life span. Before shoot growth, plant photosynthesis was mainly supported by 1-yr-old leaves, although the contribution of the 2-yr-old leaves was nonnegligible (15% of the capacity and higher carbon transfer toward the roots). After shoot growth, the current-year leaves made the greatest contribution. Our results suggest that short-lived leaves (half of the cohort) are mainly involved in a photosynthetic function, having a high photosynthetic capacity and drawing most of their resorbed N towards current-year leaves; and long-lived leaves are also involved in a conservative function, increasing N resorption and mean residence time (MRT).     

Leaf life span plasticity in tropical seedlings grown under contrasting light regimes

BACKGROUND AND AIMS: The phenotypic plasticity of leaf life span in response to low resource conditions has a potentially large impact on the plant carbon budget, notably in evergreen species not subject to seasonal leaf shedding, but has rarely been well documented. This study evaluates the plasticity of leaf longevity, in terms of its quantitative importance to the plant carbon balance under limiting light. METHODS: Seedlings of four tropical tree species with contrasting light requirements (Alstonia scholaris, Hevea brasiliensis, Durio zibethinus and Lansium domesticum) were grown under three light regimes (full sunlight, 45 % sunlight and 12 % sunlight). Their leaf dynamics were monitored over 18 months. RESULTS: All species showed a considerable level of plasticity with regard to leaf life span: over the range of light levels explored, the ratio of the range to the mean value of life span varied from 29 %, for the least plastic species, to 84 %, for the most. The common trend was for leaf life span to increase with decreasing light intensity. The plasticity apparent in leaf life span was similar in magnitude to the plasticity observed in specific leaf area and photosynthetic rate, implying that it has a significant impact on carbon gain efficiency when plants acclimate to different light regimes. In all species, median survival time was negatively correlated with leaf photosynthetic capacity (or its proxy, the nitrogen content per unit area) and leaf emergence rate. CONCLUSIONS: Longer leaf life spans under low light are likely to be a consequence of slower ageing as a result of a slower photosynthetic metabolism.     

Coordination of leaf N,anatomy, photosynthetic capacity,and hydraulics enhances evergreen expansive potential

Key message

Reduced leaf longevity, N-fixation, and enhanced hydraulic capacity combined support greater shifts in seasonal photosynthetic capacity of an expansive understory evergreen woody species relative to co-occurring less expansive evergreen species.

Abstract

Physiological functioning typically declines with increased leaf life span. While an evergreen leaf habit is generally associated with reduced leaf N, physiological capacity, and slower growth, most expansive woody species are evergreens and/or N fixers. An evergreen leaf habit enables year-round activity and less investment in carbon and nutrients, while N-fixation enhances photosynthetic capacity. Our objective was to compare anatomy and physiology of three woody evergreens Ilex opaca Aiton (Aquifoliaceae), Kalmia latifolia L. (Ericaceae), and Myrica cerifera (Myricaceae) of varying leaf longevity, N-fixation capability, and known expansive potential in a deciduous forest understory to determine if seasonal physiological performance integrated these factors. We hypothesized that I. opaca (non-expansive) and K. latifolia (moderately expansive), which have longer leaf longevities, would have reduced physiological performance compared to M. cerifera (expansive), which has shorter leaf longevity, and symbiotically fixes atmospheric N. Stomatal conductance to water vapor, photosynthetic and hydraulic capacities, specific leaf area, and leaf %N decreased with increasing leaf life span; however, trends among species were not consistent seasonally. While hydraulic capacity remained constant throughout the year, photosynthetic capacity did not. During the growing season, M. cerifera displayed photosynthetic capacity similar to deciduous species, yet, during the winter, photosynthetic capacity was similar to the slower-growing evergreens. Reduced leaf life span, enhanced hydraulic capacity, and nitrogen fixation support the seasonal shift in photosynthetic capacity observed in M. cerifera. This “hybrid” strategy enables M. cerifera to maximize productivity during months of optimal conditions, thereby promoting rapid growth and expansion in the understory.     

Leaf life spans in wild perennial herbaceous plants: a survey and attempts at a functional interpretation

Summary Leaf longevity in 29 herbaceous plant species of Central Europe was studied by inspecting tagged leaves at weekly intervals. About half of the species are elements of the lowland meadow flora, the other half comprises a representative sample of species from the highest sites where vascular plants grow in the Alps. Shaded and water-stressed sites were avoided. Overall mean leaf longevity did not differ significantly between sites and amounted to 71±5 days at low and 68±4 days at high altitude. Leaf life spans ranged (with no clear altitudinal trend) from 41 to 95 days. Low-altitude forbs and grasses produced several leaf cohorts during their growth period, while most alpine species produced only one. Correlations were found between leaf duration and percent nitrogen content and carbon-cost/carbon-gain ratios, but not with leaf dry mass per unit leaf area and photosynthetic capacity alone. As leaf life spans increase, more C tends to be invested per unit CO₂ uptake and less N is invested per unit invested C. Thus, mass relationships rather than area relationships seem to be linked to leaf life span in these species, suggesting that leaf duration is associated with properties other than the efficiency of light utilization (e.g. mechanical strength, herbivory or pathogen resistance). It seems that the explanations of leaf duration that have been developed for evergreen/deciduous plants and for plants along steep light gradients do not apply to the variable life spans in leaves of perennial herbaceous plants of open habitats.     

植物叶片最大羧化速率及其对环境因子响应的研究进展

植物叶片最大羧化速率对环境因子的响应关系是陆地生态系统生产力与碳收支研究的重要方面。论文从测定方法、影响因子与模拟模型3方面综述了植物叶片最大羧化速率及其对环境因子响应研究的最新进展,指出现有的植物叶片最大羧化速率对单个环境因子的响应研究严重制约着陆地生态系统生产力的准确评估。为弄清植物叶片最大羧化速率对环境因子的综合响应关系,迫切需要加强以下研究:(1)植物叶片最大羧化速率的生物与环境控制机制研究;(2)生物与环境因子协同作用下的植物叶片最大羧化速率定量模拟及其尺度化研究;(3)植物叶片最大羧化速率的环境因子阈值研究。     

Small trees,big problems: Comparative leaf function under extreme edaphic stress

Premise of the Study

The pygmy forest, a plant community of severely stunted conifers and ericaceous angiosperms, occurs on patches of highly acidic, nutrient‐poor soils along the coast of Northern California, USA. This system is an excellent opportunity to study the effect of severe nutrient deficiency on leaf physiology in a naturally‐occurring ecosystem. In this study, we seek to understand the physiological mechanisms stunting the plants' growth and their implications for whole plant function.

Methods

We measured 14 traits pertaining to leaf photosynthetic function or physical structure on seven species. Samples were taken from the pygmy forest community and from conspecifics growing on higher‐nutrient soils, where trees may grow over 30 m tall.

Key Results

Pygmy plants of most species maintained similar area‐based photosynthetic and stomatal conductance rates to conspecific controls, but had lower specific leaf area (leaf area divided by dry weight), lower percent nitrogen, and less leaf area relative to xylem growth. Sequoia sempervirens, a species rare in the pygmy forest, had a categorically different response from the more common plants and had remarkably low photosynthetic rates.

Conclusions

Pygmy plants were not stunted by low photosynthetic rates on a leaf‐area basis; instead, several species had restricted whole‐plant photosynthesis due to low leaf area production. Pygmy plants of all species showed signs of greater carbon investment in their leaves and higher production of nonphotosynthetic leaf tissue, further contributing to slow growth rates.     

黑龙江省次生林主要组成树种光合能力与叶片含氮量研究

以黑龙江省次生林主要组成树种蒙古栎、白桦、水曲柳、山杨、胡桃楸、黄波罗为研究对象,测定自然状态下这6个树种的光合能力,并分析光合能力与叶片含氮量之间的关系.研究结果表明,树种的光合能力存在明显的季节变化,不同树种间的光合能力、光合潜力存在差异.生长季中,胡桃楸具有最高的光合能力最大值,白桦具有最高的年平均光合能力,蒙古栎具有最大的光合潜力.蒙古栎叶片含氮量与光合能力线性正相关(r=0.97),白桦、水曲柳叶片含氮量与光合能力呈二次曲线相关(r=0.61,r=0.51).     

Functional traits and trait coordination change over the life of a leaf in a tropical fern species

Premise

Plant ecological strategies are often defined by the integration of underlying traits related to resource acquisition, allocation, and growth. Correlations between key traits across diverse plants suggest that variation in plant ecological strategies is largely driven by a fast–slow continuum of plant economics. However, trait correlations may not be constant through the life of a leaf, and it is still poorly understood how trait function varies over time in long-lived leaves.

Methods

Here, we compared trait correlations related to resource acquisition and allocation across three different mature frond age cohorts in a tropical fern species, Saccoloma inaequale.

Results

Fronds exhibited high initial investments of nitrogen and carbon, but with declining return in photosynthetic capacity after the first year. In the youngest fronds, we found water-use efficiency to be significantly lower than in the oldest mature fronds due to increased transpiration rates. Our data suggest that middle-aged fronds are more efficient relative to younger, less water-use efficient fronds and that older fronds exhibit greater nitrogen investments without higher photosynthetic return. In addition, several trait correlations expected under the leaf economics spectrum (LES) do not hold within this species, and some trait correlations only appear in fronds of a specific developmental age.

Conclusions

These findings contextualize the relationship between traits and leaf developmental age with those predicted to underlie plant ecological strategy and the LES and are among the first pieces of evidence for when relative physiological trait efficiency is maximized in a tropical fern species.

     

Ontogenetic transition in leaf traits: a new cost associated with the increase in leaf longevity

Aims Recent work has identified a worldwide 'economics' spectrum of correlated leaf traits that mainly reflects the compromises between maximizing leaf longevity and short-term productivity. However, during the early stages of tree growth different species tend to exhibit a common strategy, because competition for soil water and nutrients forces the maximization of short-term productivity owing to the need for rapid growth during the most vulnerable part of the tree's life cycle. Accordingly, our aim here was to compare the variations that occur during ontogeny in the different leaf traits (morphology and leaf chemical composition) of several coexisting Mediterranean woody species differing in their leaf life spans and to test our hypothesis that tree species with a long leaf life span should exhibit larger shifts in leaf characteristics along ontogeny.Methods Six Mediterranean tree species differing in leaf life span, selected from three plots located in central-western Spain, were studied during three growth stages: seedlings, juveniles and mature trees. Leaf life span, leaf morphology (leaf area, dry weight, thickness and mass per unit area) and chemical composition (N and fibre concentrations) were measured in all six species. The magnitude of the ontogenetic changes in the different traits was estimated and related to the mean leaf longevity of the different species.Important findings Along ontogeny, strong changes were observed in all variables analysed. The early growth stages showed lower leaf thickness, leaf thickness and mass per unit area and N, cellulose and hemicellulose concentrations than mature trees, but a higher lignin content. However, these changes were especially marked in species with a longer leaf life span at maturity. Interspecific differences in leaf life span, leaf morphology and chemical composition were stronger at the mature stage than at the seedling stage. We conclude that greater plasticity and more intense strategy shifts along ontogeny are necessarily associated with long leaf life span. Our results thus provide a new aspect that should be incorporated into the analysis of the costs and benefits associated with the different strategies related to leaf persistence displayed by the different species. Accordingly, the intensity of the alterations in leaf traits among different growth stages should be added to the suite of traits that change along the leaf economics spectrum.