Leaf plasticity in response to light of three evergreen species of the Mediterranean maquis

Morphological, anatomical, biochemical and physiological traits of sun and shade leaves of adult Quercus ilex, Phillyrea latifolia and Pistacia lentiscus shrub species co-occurring in the Mediterranean maquis at Castelporziano (Latium) were studied. Fully expanded sun leaves had 47% (mean of the three species) greater leaf mass area (LMA) and 31% lower specific leaf area (SLA) than shade leaves. Palisade parenchyma thickness contributed on an average 42% to the total leaf thickness, spongy layer 43%, upper epidermal cells 5%, and upper cuticle thickness 3%. Stomatal size was greater in sun (25.5 μm) than in shade leaves (23.6 μm). Total chlorophyll content per fresh mass was 71% greater in shade than in sun leaves, and nitrogen content was the highest in sun (13.7 mg g⁻¹) than in shade leaves (11.8 mg g⁻¹). Difference of net photosynthetic rates (P _N) between sun and shade leaves was 97% (mean of the three species). The plasticity index (sensu Valladares et al., New Phytol 148:79–91, 2000a) was the highest for physiological leaf traits (0.86) than for morphological, anatomical and biochemical ones. Q. ilex had the highest plasticity index of morphological, anatomical and physiological leaf traits (0.37, 0.28 and 0.71, respectively) that might explain its wider ecological distribution. The higher leaf plasticity of Q. ilex might be advantageous in response to varying environmental conditions, including global change.     

Structural and functional plasticity of<Emphasis Type="Italic"> Quercus ilex</Emphasis> seedlings of different provenances in Italy

Functional and structural leaf traits of Quercus ilex seedlings originated from parent plant acorns from three different localities in Italy were studied. Acorns from three different localities along a gradient from the north to the south of Italy: Nago (site A) in the Garda Lake region at the northernmost limit of holm oak distribution area in Italy, Castelporziano near Rome (site B), at the centre of the distribution area, and Frassanito near Otranto (site C), in a drier area in the south of Italy. Morphological and anatomical leaf traits differed between the provenances with a higher leaf mass area, total leaf thickness and the ratio of palisade to mesophyll thickness in the driest provenance (C seedlings). These traits gave C seedlings a higher water use efficiency, relative water content at predawn and photosynthetic rates than the other provenances in high air temperature conditions. The smaller leaf area of A seedlings seemed to have a higher photosynthetic capacity in low air temperature conditions than B and C seedlings. Growth analysis underlined a higher shoot relative growth rate in B seedlings explaining the highest shoot length and leaf number per shoot. The plasticity index [sensu Valladares et al. (2000) Ecology 81:1925–1936] for physiological traits of the seedlings was higher than morphological and anatomical traits, but the largest differences in plasticity among ecotypes were found for morphological and anatomical traits. The ecotypes of Q. ilex studied here seemed to integrate, at leaf level, functions of growth activity, morphology and physiology related to the climate of the original provenance.     

Differences in sapling architecture betweenFagus crenata andFagus japonica

The ecological significance of architectural patterns for saplings ofFagus crenata andFagus japonica co-occurring in a secondary oak forest were evaluated by comparing the size and shape of leaves, trunks and crowns.Fagus japonica saplings were different fromF. crenata saplings in some architectural properties: (i) the leaf area and specific leaf area were larger; (ii) the ratio of sapling height to trunk length was lower, indicating greater leaning of the trunk; and (iii) the projection area of the crown was larger and the leaf area index lower indicating less mutual shading of leaves. These architectural features indicated thatF. japonica saplings were more shade tolerant thanF. crenata andF. crenata saplings were superior toF. japonica for growth in height and could, therefore, utilize sunlight in the upper layer. An erect trunk inF. crenata and a leaning trunk inF. japonica may be important characteristics associated with the regenerations patterns of each species; regeneration from seedlings under canopy gaps in the former and vegetative regeneration by sprouting in the latter.     

不同种源鹅掌楸苗木叶解剖性状的遗传多样性

以5个鹅掌楸种源的1年生苗木成熟叶片为材料,对叶表皮、叶片横切面及主脉横切面的16个解剖性状进行观察和变异性分析。结果表明:鹅掌楸叶解剖性状在种源间及种源内存在极丰富的变异,除下表皮密度在种源间差异不显著外,其余性状在种源间及种源内(包括下表皮密度)均存在极显著差异。种源间的平均表型分化系数为27.5%,说明鹅掌楸叶片结构的主要性状存在于种源内的变异(72.5%)远大于种源间的变异。鹅掌楸叶的解剖性状与地理生态因子的相关分析表明,其种源间的变异呈现梯度规律性,表皮各性状与经度、年均温呈负相关关系,叶片和主脉横切面各性状与经度、纬度大部分呈正相关关系,而与年降水量、年均温大都呈负相关关系。用种源间欧式距离进行UPGMA聚类,可将5个种源的鹅掌楸划分为3个类群。     

Didymella fagi sp. nov. and its anamorphAscochyta fagi, causing the yellow leaf spot disease ofFagus crenata andQuercus mongolica var.grosseserrata in Japan

AnAscochyta fungus was obtained during a survey of leaf spot diseases ofFagus crenata in the mountains of Aomori prefecture in 1995. The pathogenicity of the fungus toF. crenata was confirmed by inoculation. Its teleomorph was first found on artificially infected leaves after the leaves were placed in an incubator at 5°C with a 12-h photoperiod (approximately 500lx, daylight strip lamps) for 4 mo. The fungus was found to be the causal agent of yellow leaf spots ofF. crenata andQuercus mongolica var.grosseserrata in the field. Comparison with similar fungi so far described from Fagaceae indicated that the anamorph isAscochyta fagi and the teleomorph is an undescribed species ofDidymella. The nameDidymella fagi is introduced for the teleomorph. Correlation between the two morphs has been proved by cultural, morphological and pathological studies.     

Morpho-anatomical responses ofTrigonella foenum graecum Linn. to induced cadmium and lead stress

Effect of different concentrations of cadmium (0, 5,15, 30, 50 μg/g of soil) and lead (0,25, 50,100,200 μg/g of soil) on morphological and anatomical features ofTrigonella foenum graecum Linn, was studied at pre-flowering, flowering and post flowering stages. Morphological attributes, like number of leaves per plant, total leaf area of the plant and single leaf area, dry mass of stem, root and leaf, length of shoot, root and plant, size of stomata and stomatal pore, and the density of stomata on both epidermises were significantly reduced under metal stress at all the developmental stages. Trichome length on both epidermises increased while their density decreased under metal stress. Under cadmium stress, proportion of pith and vasculature decreased but cortex increased in the stem. Under lead stress, proportion of pith and vasculature increased but cortex decreased in the stem. In the root, proportion of vasculature and pith increased and cortex decreased in response to both cadmium and lead stresses. Dimensions of vessel element and xylem fibre in the wood of stem and root decreased under the cadmium and lead stresses. Decrease in density of vessel element in the stem and root with advancement of age was more pronounced in plants grown under cadmium and lead stresses.     

Linkage between seasonal gas exchange and hydraulic acclimation in the top canopy leaves of Fagus trees in a mesic forest in Japan

We investigated how leaf gas exchange and hydraulic properties acclimate to increasing evaporative demand in mature beech trees, Fagus crenata Blume and Fagus japonica Maxim., growing in their natural habitat. The measurements in the top canopy leaves were conducted using a 16-m-high scaffolding tower over two growing seasons. The daily maxima of net photosynthetic rate for the early growing season were close to the annual maximum value (11.9 mol m^–2 s^–1 in F. crenata and 7.7 mol m^–2 s^–1 in F. japonica). The daily maxima of water vapor stomatal conductance were highest in the summer, approximately 0.3 mol m^–2 s^–1 in F. crenata and 0.15 mol m^–2 s^–1 in F. japonica. From the early growing season to the summer season, the leaf-to-air vapor pressure deficit increased and the daily minima of leaf water potentials decreased. However, there was no loss of leaf turgor in the summer as a result of effective osmotic adjustment. Both the soil-to-leaf hydraulic conductance per unit leaf area and the twig hydraulic conductivity simultaneously increased in the summer, probably as a result of production of new vessels in the xylem. These results suggest that both osmotic adjustment and increased hydraulic conductance resulted in the largest diurnal maximum of stomatal conductance in the summer, resulting in the lowest relative stomatal limitation on net photosynthetic rate, although the leaf-to-air vapor pressure deficit was highest. These results indicate that even in a mesic forest, in which excessive hydraulic stress does not occur, the seasonal acclimation of hydraulic properties at both the single leaf and whole plant levels are important for plant carbon gain.     

The seasonal occurrence of endophytic fungus, <Emphasis Type="Italic">Mycosphaerella buna</Emphasis>, in Japanese beech, <Emphasis Type="Italic">Fagus crenata</Emphasis>

The seasonal occurrence of Mycosphaerella buna in leaves and contiguous organs of Fagus crenata was studied in a Japanese beech forest, Ibaraki, Japan, in 1998 and 1999. Mycosphaerella buna was not isolated from newly developed leaves in May, but it was isolated from asymptomatic leaves after June. The frequency of its occurrence gradually increased until leaffall. In contrast, M. buna was not isolated from overwintered buds, leaf petioles, or contiguous current-year twigs. The spermogonia and pseudothecia were observed in dead leaves after leaffall. The mature pseudothecia were found on dead leaves from May to July. The ascospores produced in the pseudothecia were suggested to infect newly developed leaves.Contribution no. 173, Laboratory of Plant Parasitic Mycology, Institute of Agriculture and Forestry, University of Tsukuba, Japan     

Temporal and spatial variations in leaf herbivory within a canopy of<Emphasis Type="Italic"> Fagus crenata</Emphasis>

This study investigated spatio-temporal variation in the leaf area consumed by insect herbivores within a canopy of Fagus crenata, with reference to the light conditions of leaf clusters. There was no clear relationship between photosynthetic photon flux density (PPFD) and consumed leaf area (CLA) in May, immediately after leaf flush, but CLA decreased with an increase in PPFD after June. Leaf mass per area, carbon concentration, C/N ratio, concentration of total phenolics, and condensed tannin concentration were higher in leaves under high light intensity than those of leaves under low light. On the other hand, the nitrogen concentration of leaves decreased as light availability increased. Consequently, within-tree variation in light availability affects the consumption of leaves by insect herbivores through temporal changes in leaf characteristics.     

Leaf traits variation during leaf expansion in <Emphasis Type="Italic">Quercus ilex</Emphasis> L.

The morphological, anatomical and physiological variations of leaf traits were analysed during Quercus ilex L. leaf expansion. The leaf water content (LWC), leaf area relative growth rate (RGR_l) and leaf dry mass relative growth rate (RGR_m) were the highest (76±2 %, 0.413 cm² cm⁻² d⁻¹, 0.709 mg mg⁻¹ d⁻¹, respectively) at the beginning of the leaf expansion process (7 days after bud break). Leaf expansion lasted 84±2 days when air temperature ranged from 13.3±0.8 to 27.6±0.9 °C. The net photosynthetic rate (P _N), stomatal conductance (g _s), and chlorophyll content per fresh mass (Chl) increased during leaf expansion, having the highest values [12.62±1.64 μmol (CO₂) m⁻² s⁻¹, 0.090 mol (H₂O) m⁻² s⁻¹, and 1.03±0.08 mg g⁻¹, respectively] 56 days after bud break. Chl was directly correlated with leaf dry mass (DM) and P _N. The thickness of palisade parenchyma contributed to the total leaf thickness (263.1±1.5 μm) by 47 %, spongy layer thickness 38 %, adaxial epidermis and cuticle thickness 9 %, and abaxial epidermis and cuticle thickness 6 %. Variation in leaf size during leaf expansion might be attributed to a combination of cells density and length, and it is confirmed by the significant (p<0.001) correlations among these traits. Q. ilex leaves reached 90 % of their definitive structure before the most severe drought period (beginning of June — end of August). The high leaf mass area (LMA, 15.1±0.6 mg cm⁻²) at full leaf expansion was indicative of compact leaves (2028±100 cells mm⁻²). Air temperature increasing might shorten the favourable period for leaf expansion, thus changing the final amount of biomass per unit leaf area of Q. ilex.     

Growth and photosynthetic response of <Emphasis Type="Italic">Fagus crenata</Emphasis> seedlings to ozone and/or elevated carbon dioxide

We investigated the effects of ozone (O₃) and/or elevated CO₂ concentration ([CO₂]) on growth and photosynthetic traits of Fagus crenata seedlings. Two-year-old seedlings were grown in four experimental treatments comprising two O₃ treatments (charcoal-filtered air and 100 nmol mol⁻¹ O₃; 6 h/day, 3 days/week) in combination with two CO₂ treatments (350 and 700 μmol mol⁻¹) for 18 weeks in environmental control growth chambers. The four treatments were designated as control, elevated O₃, elevated CO₂, and elevated CO₂ + O₃. Dry matter growth of the seedlings was greater in elevated CO₂ + O₃ than in elevated CO₂. In elevated CO₂ + O₃, a marked increase of second-flush leaves, considered a compensative response to O₃, was observed. The net photosynthetic rate of first-flush leaves in elevated CO₂ + O₃ increased earlier and was maintained for a longer period of time than that in elevated CO₂. Because emergence of second-flush leaves of F. crenata is greatly affected by the amount of assimilation products of first-flush leaves in current year, we consider that an early increase in the net photosynthetic rate of first-flush leaves contributed to the marked increase in second-flush leaf emergence under elevated CO₂ + O₃. These results imply that we must account for changes in compensative capacity with respect to not only morphological traits but also phenological traits and physiological functions such as photosynthesis when evaluating effects of O₃ on F. crenata under elevated [CO₂].     

Differences in response to simulated herbivory between <Emphasis Type="Italic">Quercus crispula</Emphasis> and <Emphasis Type="Italic">Quercus dentata</Emphasis>

To evaluate the responses of Quercus crispula and Quercus dentata to herbivory, their leaves were subjected to simulated herbivory in early spring and examined for the subsequent changes in leaf traits and attacks by chewing herbivores in mid summer. In Quercus crispula, nitrogen content per area was higher in artificially damaged leaves than in control leaves. This species is assumed to increase the photosynthetic rate per area by increasing nitrogen content per area to compensate leaf area loss. In Quercus dentata, nitrogen content per area did not differ between artificially damaged and control leaves, while nitrogen content per mass was slightly lower in artificially damaged leaves. The difference in their responses can be attributable to the difference in the architecture of their leaves and/or the severeness of herbivory. The development of leaf area from early spring to mid summer was larger in artificially damaged leaves than in control leaves in both species, suggesting the compensatory response to leaf area loss. Leaf dry mass per unit area was also larger in artificially damaged leaves in both species, but the adaptive significance of this change is not clear. In spite of such changes in leaf traits, no difference was detected in the degree of damage by chewing herbivores between artificially damaged and controlled leaves in both species.     

Variation in mesophyll anatomy and photosynthetic capacity during leaf development in a deciduous mesophyte fruit tree (<Emphasis Type="Italic">Prunus persica</Emphasis>) and an evergreen sclerophyllous Mediterranean shrub (<Emphasis Type="Italic">Olea europaea</Emphasis>)

The relative importance that biomechanical and biochemical leaf traits have on photosynthetic capacity would depend on a complex interaction of internal architecture and physiological differences. Changes in photosynthetic capacity on a leaf area basis and anatomical properties during leaf development were studied in a deciduous tree, Prunus persica, and an evergreen shrub, Olea europaea. Photosynthetic capacity increased as leaves approached full expansion. Internal CO₂ transfer conductance (g _i) correlated with photosynthetic capacity, although, differences between species were only partially explained through structural and anatomical traits of leaves. Expanding leaves preserved a close functional balance in the allocation of resources of photosynthetic component processes. Stomata developed more rapidly in olive than in peach. Mesophyll thickness doubled from initial through final stages of development when it was twice as thick in olive as in peach. The surface area of mesophyll cells exposed to intercellular air spaces per unit leaf area tended to decrease with increasing leaf expansion, whereas, the fraction of mesophyll volume occupied by the intercellular air spaces increased strongly. In the sclerophyllous olive, structural protection of mesophyll cells had priority over efficiency of photochemical mechanisms with respect to the broad-leaved peach. The photosynthetic capacity of these woody plants during leaf development relied greatly on mesophyll properties, more than on leaf mass per area ratio (LMA) or nitrogen (N) allocation. Age-dependent changes in diffusion conductance and photosynthetic capacity affected photosynthetic relationships of peach versus olive foliage, evergreen leaves maturing functionally and structurally a bit earlier than deciduous leaves in the course of adaptation for xeromorphy.     

黄土高原先锋种猪毛蒿叶片形态解剖与生理特征对立地的适应性

为了揭示黄土丘陵沟壑区撂荒地植被演替前期优势种猪毛蒿(Artemisia scoparia)对该区立地环境的适应性,探讨猪毛蒿演替生态位的变化,研究了陕北黄土丘陵沟壑区3种立地环境下(阳峁坡、峁顶、阴峁坡)猪毛蒿叶片形态解剖和生理特征的变化,以及这些变化与生态因子之间的相互关系。结果表明:(1)猪毛蒿叶片具有适应该区半干旱环境的形态及解剖结构:叶片针形化、具表皮毛、环栅型叶肉组织、海绵组织特化为贮水组织、维管束退化、具裂生分泌腔,C3植物呈现类似CAM植物的叶片特性;(2)在土壤、空气湿度相对干燥和强光生境的阳峁坡与峁顶,猪毛蒿具有较小的叶面积、发达的栅栏组织、致密的表皮毛和紧密的细胞间隙,而在生境条件较好的阴峁坡则呈相反趋势;(3)阳峁坡猪毛蒿叶片相对含水量和叶绿素含量较小,超氧阴离子自由基增加,但植物体内超氧化物歧化酶和抗坏血酸含量增加以清除植物体内产生的活性氧;(4)冗余及相关性分析表明,猪毛蒿叶片形态、解剖和生理指标的可塑性对立地光照强度、土壤水分和有机质含量较为敏感,同时其形态解剖与生理可塑性可共同调节来适应生境。综合分析,猪毛蒿对陕北黄土丘陵沟壑区撂荒初期光照强度大、土壤贫瘠但土层干化现象尚未出现的立地环境有较好的适应性,使其成为黄土高原植被自然演替过程中的先锋物种。     

不同种源樟树叶片形态特征及生长差异分析

为了解不同种源樟树叶片形态特征和生长差异,该文以30个种源樟树为研究对象,对其叶长、叶宽、叶柄长、周长、叶面积、长宽比、形态因子、株高、地径等指标进行测定和差异性分析.结果表明:(1)30个种源间叶片性状的变异系数为3.88％～16.14％,显示不同种源樟树叶片形态特征存在显著差异;叶长、叶宽、叶柄长、周长、面积、叶厚...     

不同种源桃金娘表型性状多样性研究

为解析桃金娘表型性状多样性及其种源间关系，该文以20个不同来源的桃金娘为研究对象，在同质园栽培条件下，对其营养器官和花器官表型性状进行观测，采用方差分析、变异分析、Shannon-Wiener多样性指数分析和聚类分析等方法，探讨不同种源桃金娘表型性状多样性。结果表明：(1)不同种源桃金娘表型性状存在显著差异(P<0.05),Shannon-Wiener多样性指数均值在1.35以上，表型性状多样性丰富。(2)种源内表型性状变异系数均值在10.81%～63.75%之间，种源间的变异系数均值在13.08%～74.04%之间，种源间变异(23.33%)高于种源内变异(19.79%),营养器官变异(29.52%)高于花器官变异(14.06%)。(3)部分性状存在极显著或显著相关性，株高与分枝数呈极显著负相关，而与叶长、叶宽和叶面积等却呈显著正相关。(4)在欧式距离10处，20个种源桃金娘可分为A、B、C三类，A类包含8个种源，该类种源表现为植株高大、分枝少、叶大和花大等特征；B类包含11个种源，该类种源表现为株高中等、叶较大和花中等等特征；C类仅包含1个种源，表现为植株低矮、分枝多、叶小和...     

<Emphasis Type="SmallCaps">P</Emphasis>lastic responses to temporal variation in moisture availability: consequences for water use efficiency and plant performance

The ability to appropriately modify physiological and morphological traits in response to temporal variation should increase fitness. We used recombinant hybrid plants generated by crossing taxa in the Piriqueta caroliniana complex to assess the effects of individual leaf traits and trait plasticities on growth in a temporally variable environment. Recombinant hybrids were used to provide a wide range of trait expression and to allow an assessment of the independent effects of individual traits across a range of genetic backgrounds. Hybrid genotypes were replicated through vegetative propagation and planted in common gardens at Archbold Biological Station in Venus, Florida, where they were monitored for growth, leaf morphological characters, and integrated water use efficiency (WUE) (C isotope ratio; δ¹³C) for two successive seasons. Under wet conditions only leaf area had significant effects on plant growth, but as conditions became drier, growth rates were greatest in plants with narrow leaves and higher trichome densities. Plants with higher WUE exhibited increased growth during the dry season but not during the wet season. WUE during the dry season was increased for plants with smaller, narrower leaves that had higher trichome densities and increased reflectance. Examination of alternative path models revealed that during the dry season leaf traits had significant effects on plant growth only through their direct effects on WUE, as estimated from δ¹³C. Over the entire growing season, plants with a greater ability to produce smaller and narrower leaves with higher trichome densities in response to reduced water availability had the greatest growth rate. These findings suggest that plants making appropriate changes to leaf morphology as conditions became dry had increased WUE, and that the ability to adjust leaf phenotypes in response to environmental variation is a mechanism by which plants increase fitness.     

The Structural Adaptation of Aerial Parts of Invasive <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> to Water Regime

Alternanthera philoxeroides has successfully invaded diverse habitats with considerably various water availability, threatening biological diversity in many parts of the world. Because its genetic variation is very low, phenotypic plasticity is believed to be the primary strategy for adapting to the diverse habitats. In the present paper, we investigated the plastic changes of anatomical traits of the aerial parts of A. philoxeroides from flooding to wet then to drought habitat; the results are as follows: A. philoxeroides could change anatomical structures sensitively to adapt to water regime. As a whole, effects of water regime on structures in stem were greater than those in leaf. Except for principal vein diameter and stoma density on leaf surfaces, all other structural traits were significantly affected by water regime. Among which, cuticular wax layer, collenchyma cell wall, phloem fiber cell wall, and hair density on both leaf surfaces thickened significantly with decrease of water availability, whereas, pith cavity and vessel lumen in stem lessened significantly; wet habitat is vital for the spread of A. philoxeroides from flooding to drought habitat and vice versa, because in this habitat, it had the greatest structural variations; when switching from flooding to wet then to drought habitat, the variations of cuticular wax layer, collenchyma cell wall, phloem fiber cell wall, pith cavity area ratio, diameter of vessel lumen, and hair density on both leaf surfaces, played the most important role. These responsive variables contribute most to the adaptation of A. philoxeroides to diverse habitats with considerably various water availability.     

同期生长在种源地与异地同质园刨花楠叶性状分析

为辨别环境变化与遗传因素对植物叶片主要功能性状的影响,以同期生长在4种源地母树林下及异地同质园的1.5年生刨花楠苗木为研究对象,对其叶片表型及养分性状进行对比分析。结果显示：(1)刨花楠叶面积、叶厚、叶干物质含量等叶片表型性状受遗传与环境因素共同影响;叶片碳(C)含量受遗传因素调控,环境对其影响较小;叶片氮、磷(N、P)含量主要受环境因素影响;(2)不同种源刨花楠比叶面积、叶厚、叶干物质含量、叶形指数等性状变异系数较大(8.85%—37.03%),其中江西遂川种源变异系数相对较大,而湖南茶陵种源则相对较小,各种源都倾向于通过调节比叶面积、叶厚、叶片氮磷含量等性状以适应生境变化;(3)种源地与同质园刨花楠的比叶面积虽均与叶厚呈显著负相关,但同质园刨花楠比叶面积与叶形指数呈显著正相关,与叶片氮含量无明显相关,而种源地刨花楠比叶面积则与叶形指数无明显相关,与叶片氮含量则呈显著负相关;(4)不同种源苗木叶性状指标在种源地与同质园间存在不同的协调与权衡,体现了植物在不同生境下的适应策略。其中湖南茶陵种源在两种生境下都具有更保守的资源获取策略,而江西安福种源对环境变化则更为敏感,资源获取策略更为灵...     

Coordinated changes in photosynthesis,water relations and leaf nutritional traits of canopy trees along a precipitation gradient in lowland tropical forest

We investigated leaf physiological traits of dominant canopy trees in four lowland Panamanian forests with contrasting mean annual precipitation (1,800, 2,300, 3,100 and 3,500 mm). There was near complete turn-over of dominant canopy tree species among sites, resulting in greater dominance of evergreen species with long-lived leaves as precipitation increased. Mean structural and physiological traits changed along this gradient as predicted by cost–benefit theories of leaf life span. Nitrogen content per unit mass (N_mass) and light- and CO₂-saturated photosynthetic rates per unit mass (P_mass) of upper canopy leaves decreased with annual precipitation, and these changes were partially explained by increasing leaf thickness and decreasing specific leaf area (SLA). Comparison of 1,800 mm and 3,100 mm sites, where canopy access was available through the use of construction cranes, revealed an association among extended leaf longevity, greater structural defense, higher midday leaf water potential, and lower P_mass, N_mass, and SLA at wetter sites. Shorter leaf life spans and more enriched foliar ¹⁵N values in drier sites suggest greater resorption and re-metabolism of leaf N in drier forest. Greater dominance of short-lived leaves with relatively high P_mass in drier sites reflects a strategy to maximize photosynthesis when water is available and to minimize water loss and respiration costs during rainless periods. Overall, our study links coordinated change in leaf functional traits that affect productivity and nutrient cycling to seasonality in lowland tropical forests.