Photosynthetic and morphological acclimation of seedlings of tropical lianas to changes in the light environment.

UNLABELLED: • PREMISE OF THE STUDY: Few studies have analyzed the physiological performance of different life stages and the expression of ontogenetic niche shifts in lianas. Here, we analyzed the photosynthetic and morphological acclimation of seedlings of Stigmaphyllon lindenianum, Combretum fruticosum, and Bonamia trichantha to distinctive light conditions in a tropical dry forest and compared their response with the acclimation response of adult canopy lianas of the same species. We expected acclimation to occur faster through changes in leaf photochemistry relative to adaptation in morphology, consistent with the life history strategies of these lianas.• METHODS: Seedlings were assigned to the following light treatments: high light (HH), low light (LL), sun to shade (HL), and shade to sun (LH) in a common garden. After 40 d, HL and LH seedlings were exposed to opposite light treatments. Light response curves, the maximum photosynthetic rate in the field (Amax), and biomass allocation were monitored for another 40 d on leaves expanded before transfer.• KEY RESULTS: Photosynthetic responses, Amax, and biomass of Stigmaphyllon and Combretum varied with light availability. Physiological characters were affected by current light environment. The previous light environment (carryover effects) only influenced Amax. Morphological characters showed significant carryover effects. Stigmaphyllon showed high morphological and physiological plasticity. Sun-exposed seedlings of this liana increased stem biomass and switched from self-supporting to climbing forms.• CONCLUSIONS: Acclimation in seedlings of these lianas is consistent with the response of adult lianas in the canopy in direction, but not in magnitude. There was no evidence for ontogenetic niche shifts in the acclimation response.

Ecological and evolutionary consequences of desiccation tolerance in tropical fern gametophytes.

Ferns have radiated into the same diverse environments as spermatophytes, and have done so with an independent gametophyte that is not protected by the parent plant. The degree and extent of desiccation tolerance (DT) in the gametophytes of tropical fern species was assessed to understand mechanisms that have allowed ferns to radiate into a diversity of habitats. Species from several functional groups were subjected to a series of desiccation events, including varying degrees of intensity and multiple desiccation cycles. Measurements of chlorophyll fluorescence were used to assess recovery ability and compared with species ecology and gametophyte morphology. It is shown that vegetative DT (rare in vascular plants) is widely exhibited in fern gametophytes and the degree of tolerance is linked to species habitat preference. It is proposed that gametophyte morphology influences water-holding capacity, a novel mechanism that may help to explain how ferns have radiated into drought-prone habitats. Fern gametophytes have often been portrayed as extreme mesophytes with little tolerance for desiccation. The discovery of DT in gametophytes holds potential for improving our understanding of both the controls on fern species distribution and their evolution. It also advances a new system with which to study the evolution of DT in vascular plants.

Gametophyte ecology and demography of epiphytic and terrestrial tropical ferns.

Factors that influence the distribution of ferns are poorly understood and likely reflect the ecology of both the sporophyte and the gametophyte generation. Little study has been done on the ecology of the gametophyte generation, especially in regard to tropical species. The goal of this study was to examine demography and the influence of light and disturbance on the distribution of the gametophytes of several tropical epiphytic, hemiepiphytic, and terrestrial fern species. Through a series of observational and experimental studies, we found that increased terrestrial gametophyte density and richness were related to both increased light and disturbance. By contrast, increased light had no influence, and increased disturbance negatively affected epiphytic density. Over a 25-mo demographic study, epiphytic and hemiepiphytic species had significantly greater longevities and lower recruitment rates than terrestrial species. Such unique strategies may have evolved in response to different disturbance regimens between the two habitats. Terrestrial species encounter and are adapted to more frequent disturbance and have invested in rapid gametophyte growth and recruitment. Epiphytic species may be more influenced by bryophyte competition, and in habitats of relatively low disturbance, they have invested in greater size and longevities. In such systems, gametophytes are able to survive for years waiting for favorable recruitment conditions.

Correlation but no causation between leaf nitrogen and maximum assimilation: the role of drought and reproduction in gas exchange in an understory tropical plant Miconia ciliata (Melastomataceae).

Alternative hypotheses were tested to explain a previously reported anomaly in the response of leaf photosynthetic capacity at light saturation (A(max)) in Miconia ciliata to dry-season irrigation. The anomaly is characterized by an abrupt increase in leaf A(max) for nonirrigated plants at the onset of the rainy season to values that significantly exceeded corresponding measurements for plants that were irrigated during the previous dry season. Hypothesis 1 posits that a pulse in leaf nitrogen increases CO(2) assimilation in nonirrigated plants at the onset of the wet season and is dampened for irrigated plants; this hypothesis was rejected because, although a wet-season nitrogen pulse did occur, it was identical for both irrigated and nonirrigated plants and was preceded by the increase in assimilation by nonirrigated plants. Hypothesis 2 posits that a reproduction-related, compensatory photosynthetic response occurs in nonirrigated plants following the onset of the wet season and is dampened in irrigated plants; consistent with hypothesis 2, high maximum assimilation rates for control plants in the wet season were significantly correlated with fruiting and flowering, whereas irrigation caused flowering and fruiting in the dry season, spreading M. ciliata reproductive activity in irrigated plants across the entire year.

Variation in crown light utilization characteristics among tropical canopy trees.

BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.

Photochemical efficiency of adult and young leaves of the neotropical understory shrub Psychotria limonensis (Rubiaceae)in response to changes in the light environment

We explored the short-term adjustment in photochemical efficiency (Fv /Fm )in adult and young leaves of the understory neotropical shrub Psychotria limonensis Krause (Rubiaceae)in response to rapid changes in the light environment.Leaves were collected from 20 individual plants growing under sun and shade conditions on Gigante Peninsula,Barro Colorado Natural Monument (Republic of Panama),during the wet season of 1996. Leaves were distributed in four sequences of light treatments (AB leaves were expanded under sun and were transferred to shade,BA leaves experienced the opposite transfer,and the controls AA and BB leaves that were expanded and maintained under sun or shade conditions).Adult and young leaves did not differ in overall photochemical efficiency.Instead,differences were found among light environments,for which leaves transferred from shade to sun showed the lowest F v /F m ratios.There was no relationship between photochemical efficiency and leaf temperature.In P.limonensis,understory plants are susceptible of photoinhibition independently of the leaf ontogenetic stage.The approach utilized in this experiment allowed the rapid exploration of this capacity, and could be applied to poorly studied understory species

Se exploró el ajuste a corto plazo en la eficiencia fotosintética (Fv /Fm )en hojas jovenes y adultas del arbusto del sotobosque neotropical Psychotria limonensis Krause (Rubiaceae)en respuesta a cambios rápidos de luz ambiental. Las hojas fueron recolectadas de 20 plantas individuales bajo condiciones de sol y sombra en Peninsula Gigante, Monumento Natural Barro Colorado (Panamá),durante la estación lluviosa de 1996.Las hojas fueron distribuidas en una secuencia cuatro tratamientos de luz (AB las hojas fueron expandidas bajo el sol y fueron transferidas a la sombra,BA las hojas experimentaron la transferencia contraria,y las hojas controles AA y BB que fueron expandidas y mantenidas bajo condiciones de sol o de sombra).Las hojas adultas o jóvenes no difieren en la eficiencia fotoquímica general.Por el contrario,se encontró diferencias entre los ambientes de luz (iluminados), para los cuales las hojas transferidas de la sombra al sol mostraron las menores tasas Fv /Fm .No hubo relación entre la eficiencia fotoquímica y la temperatura de las hojas. En P.limonensis ,las plantas son suceptibles a la foto -inhibición independientemente del estado ontogenético de la hoja. El enfoque utilizado en este experimento permitió la rápida exploración de esta capacidad y demostró que puede ser utilizado en otras especies poco estudiadas del sotobosque

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 (Nmass) and light- and CO2-saturated photosynthetic rates per unit mass (Pmass) 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 Pmass, Nmass, and SLA at wetter sites. Shorter leaf life spans and more enriched foliar delta15N 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 Pmass 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.

Photochemical efficiency of adult and young leaves of the neotropical understory shrub Psychotria limonensis (Rubiaceae) in response to changes in the light environment.

We explored the short-term adjustment in photochemical efficiency (Fv/Fm) in adult and young leaves of the understory neotropical shrub Psychotria limonensis Krause (Rubiaceae) in response to rapid changes in the light environment. Leaves were collected from 20 individual plants growing under sun and shade conditions on Gigante Peninsula, Barro Colorado Natural Monument (Republic of Panama), during the wet season of 1996. Leaves were distributed in four sequences of light treatments (AB leaves were expanded under sun and were transferred to shade, BA leaves experienced the opposite transfer, and the controls AA and BB leaves that were expanded and maintained under sun or shade conditions). Adult and young leaves did not differ in overall photochemical efficiency. Instead, differences were found among light environments, for which leaves transferred from shade to sun showed the lowest Fv/Fm ratios. There was no relationship between photochemical efficiency and leaf temperature. In P. limonensis, understory plants are susceptible of photoinhibition independently of the leaf ontogenetic stage. The approach utilized in this experiment allowed the rapid exploration of this capacity, and could be applied to poorly studied understory species.

Cloud cover limits net CO2 uptake and growth of a rainforest tree during tropical rainy seasons.

Recent global-scale analyses indicate that climate variability affects net carbon storage but regard temperature and precipitation to be the main contributors. Seasonal and interannual variation in light availability may also limit CO(2) uptake. As an experimental test of light limitation by cloud cover during tropical rainy seasons and by the unusually heavy cloud cover associated with La Niña, we installed high-intensity lamps above the forest canopy to augment light for Luehea seemannii, a tropical canopy tree species, during cloudy periods of 1999-2000. Light augmentation only partially compensated for the reduction in photosynthetic photon flux density caused by clouds. Nonetheless, leaves acclimated to the augmented irradiance, and photosynthesis, vegetative growth, and reproduction increased significantly. Light, rather than water, temperature, or leaf nitrogen, was the primary factor limiting CO(2) uptake during the rainy season.

Drought constraints on leaf gas exchange by Miconia ciliata (Melastomataceae) in the understory of an eastern Amazonian regrowth forest stand.

Analyses of the effects of drought stress on Amazonian regrowth stands are lacking. We measured leaf gas exchange and leaf water potential of Miconia ciliata (Melastomataceae) in a dry-season irrigation experiment in 14-yr-old regrowth. In the dry season, irrigated plants maintained significantly higher leaf water potentials, photosynthetic capacity at light saturation (A(max)), stomatal conductance (g(s)), internal CO(2) concentration (C(i)), and lower A(max)/g(s) than control plants. The degree of dry-season down-regulation of control plant A(max), along with its fast recovery following rain, reveals the importance of occasional dry-season rains to the carbon budget of M. ciliata. During the wet season, we observed higher A(max) for control plants than for plants that had been irrigated during the dry season. We hypothesize that reduced drought constraints on photosynthesis of irrigated plants advanced the flowering and fruiting phenology of irrigated plants into the dry season. Flowers and fruits of control plants developed later, during the wet season, potentially stimulating a compensatory reproductive photosynthesis response in nearby leaves. The relative drought intolerance of M. ciliata may be a deciding factor in its ability to survive through the dynamic successional development of the regrowth stand studied.

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.

Seasonal patterns of carbohydrate storage in four tropical tree species.

We examined the seasonal variation in total non-structural carbohydrate (TNC) concentrations in branch, trunk, and root tissues of Anacardium excelsum, Luehea seemannii, Cecropia longipes, and Urera caracasana growing in a seasonally dry forest in Panama. Our main goals were: (1) to determine the main sites of carbohydrate storage, and (2) to determine if seasonal patterns of carbohydrate storage are related to seasonal asynchronies in carbon supply and demand. We expected asynchronies to be related to seasonal variation in water and light availability and to foliar and reproductive phenology. Cecropia and Urera are fully drought-deciduous and so we expected them to exhibit the most dramatic seasonal variation in TNC concentrations. We predicted that maximum carbon supply would occur when canopies were at their fullest and that maximum carbon demand would occur when leaves, flowers, and fruits were produced. The concentration of total non-structural carbohydrates was assessed monthly in wood tissue of roots and in wood and bark tissue of terminal branches. Trunk tissue was sampled bimonthly. All tissues sampled served as storage sites for carbohydrates. As predicted, TNC concentrations varied most dramatically in branches of Cecropia and Urera: a 4-fold difference was observed between dry season maxima and wet season minima in branch wood tissue. Peak concentrations exceeded 25% in Urera and 30% in Cecropia. Less dramatic but significant seasonal variation was observed in Anacardium and Luehea. In all species, minimum branch TNC concentrations were measured during canopy rebuilding. In Anacardium, maximum branch TNC concentrations occurred when canopies were at their fullest. In Cecropia, Urera, and Luehea, TNC concentrations continued to increase even as canopies thinned in the early dry season. The greater photosynthetic capacity of leaves produced at the beginning of the dry season and the potential for the export of carbohydrates from senescing leaves may explain this pattern. In all species, the phenology of carbon gain was more important than the phenology of reproduction in influencing seasonal carbohydrate patterns. The combination of high TNC concentrations and the large biomass of branches, trunks, and roots indicates these species are storing and moving large quantities of carbohydrates.

Photosynthetic acclimation of the liana Stigmaphyllon lindenianum to light changes in a tropical dry forest canopy.

Tropical plant canopies show abrupt changes in light conditions across small differences in spatial and temporal scales. Given the canopy light heterogeneity, plants in this stratum should express a high degree of plasticity, both in space (allocation to plant modules as a function of opportunity for resource access) and time (photosynthetic adjustment to temporal changes in the local environment). Using a construction crane for canopy access, we studied light acclimation of the liana Stigmaphyllon lindenianum to sun and shade environments in a tropical dry forest in Panama during the wet season. Measured branches were randomly distributed in one of four light sequences: high- to low-light branches started the experiment under sun and were transferred to shade during the second part of the experiment; low- to high-light branches (LH) were exposed to the opposite sequence of light treatments; and high-light and low-light controls , which were exposed only to sun and shade environments, respectively, throughout the experiment. Shade branches were set inside enclosures wrapped in 63% greenhouse shade cloth. After 2 months, we transferred experimental branches to opposite light conditions by relocating the enclosures. Leaf mortality was considerably higher under shade, both before and after the transfer. LH branches reversed the pattern of mortality by increasing new leaf production after the transfer. Rates of photosynthesis at light saturation, light compensation points, and dark respiration rates of transferred branches matched those of controls for the new light treatment, indicating rapid photochemical acclimation. The post-expansion acclimation of sun and shade foliage occurred with little modification of leaf structure. High photosynthetic plasticity was reflected in an almost immediate ability to respond to significant changes in light. This response did not depend on the initial light environment, but was determined by exposure to new light conditions. Stigmaphyllon responded rapidly to light changes through the functional adjustment of already expanded foliage and an increase in leaf production in places with high opportunity for carbon gain.

Seasonal leaf phenotypes in the canopy of a tropical dry forest: photosynthetic characteristics and associated traits.

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 (O2max) and as light-saturated CO2 assimilation rates of intact leaves under ambient CO2 (Amax). In four species, pre-dry season leaves had significantly higher leaf mass per unit area. In these four species, O2max and Amax per unit area and maximum stomatal conductances were significantly greater in pre-dry season leaves than in early wet season leaves. In two species, Amax 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.

Comparative physiology and demography of three Neotropical forest shrubs: alternative shade-adaptive character syndromes.

A suite of functionally-related characters and demography of three species of Neotropical shadeadapted understory shrubs (Psychotria, Rubiaceae) were studied in the field over five years. Plants were growing in large-scale irrigated and control treatments in gaps and shade in old-growth moist forest at Barro Colorado Island, Panama. Irrigation demonstrated that dry-season drought limited stomatal conductance, light saturated photosynthesis, and leaf longevity in all three species. Drought increased mortality of P. furcata. In contrast, irrigation did not affect measures of photosynthetic capacity determined with an oxygen electrode or from photosynthesis-CO2 response curves in the field. Drought stress limited field photosynthesis and leaf and plant survivorship without affecting photosynthetic capacity during late dry season. Leaves grown in high light in naturally occurring treefall gaps had higher photosynthetic capacity, dark respiration and mass per unit area than leaves grown in the shaded understory. P. furcata had the lowest acclimation to high light for all of these characters, and plant mortality was greater in gaps than in shaded understory for this species. The higher photosynthetic capacity of gap-grown leaves was also apparent when photosynthetic capacity was calculated on a leaf mass basis. Acclimation to high light involved repackaging (higher mass per unit leaf area) as well as higher photosynthetic capacity per unit leaf mass in these species. The three species showed two distinct syndromes of functionally-related adaptations to low light. P. limonensis and P. marginata had high leaf longevity (∼3 years), high plant survivorship, low leaf nitrogen content, and high leaf mass per unit area. In contrast, P. furcata had low leaf survivorship (∼1 year), high plant mortality (77-96% in 39 months), low leaf mass per unit area, high leaf nitrogen content, and the highest leaf area to total plant mass; the lowest levels of shelf shading, dark respiration and light compensation; and the highest stem diameter growth rates. This suite of characters may permit higher whole-plant carbon gain and high leaf and population turnover in P. furcata. Growth in deep shade can be accomplished through alternative character syndromes, and leaf longevity may not be correlated with photosynthetic capacity in shade adapted plants.

Drought acclimation among tropical forest shrubs (Psychotria, Rubiaceae).

Mechanisms of dry-season drought resistance were evaluated for five evergreen shrubs (Psychotria, Rubiaceae) which occur syntopically in tropical moist forest in central Panama. Rooting depths, leaf conductance, tissue osmotic potentials and elasticity, and the timing of leaf production were evaluated. From wet to dry season, tissue osmotic potentials declined and moduli of elasticity increased in four and five species, respectively. Irrigation only affected osmotic adjustment by P. furcata. The other seasonal changes in leaf tissue properties represented ontogenetic change. Nevertheless, they made an important contribution to dry-season turgor maintenance. Small between-year differences in dry season rainfall had large effects on plant water status. In 1986, 51 mm of rain fell between 1 January and 31 March, and pre-dawn turgor potentials averaged <0.1 MPa for all five Psychotria species in March (Wright 1991). In 1989, 111 mm of rain fell in the same period, pre-dawn turgor potentials averaged from 0.75 to 1.0 MPa for three of the species in April, and only P. chagrensis lost turgor. The relation between leaf production and drought differed among species. P. limonensis was buffered against drought by the lowest dry-season conductances and the deepest roots (averaging 244% deeper than its congeners) and was the only species to produce large numbers of leaves in the dry season. P. chagrensis was most susceptible to drought, and leaf production ceased as turgor loss developed. For the other species, water stress during severe dry seasons may select against dry-season leaf production.

Comparative life history and physiology of two understory Neotropical herbs.

Demography and physiology of two broad-leaved understory tropical herbs (Marantaceae) were studied in gaps and shaded understory in large-scale irrigated and control treatments during the dry season at Barro Colorado Island (BCI), Panama. Because photosynthetic acclimation potential may not predict light environments where tropical species are found, we studied a suite of physiological features to determine if they uniquely reflect the distribution of each species. Calathea inocephala and Pleiostachya pruinosa grow and reproduce in gaps, persist in shade, and have equivalent rates of leaf production. Calathea leaves survived 2 to 3 times as long as leaves of Pleiostachya and plants of Pleiostachya were 6 to 8 times more likely to die as plants of Calathea during 3.5 years of study. Pleiostachya had lowest survival in shade and when not irrigated during the dry season, while Calathea survived well in both habitats and both treatments. Pleiostachya had higher photosynthetic capacity and stomatal conductance than Calathea and acclimated to gaps by producing leaves with higher photosynthetic capacity. Calathea had lower mesophyll CO2 concentrations than Pleiostachya. Both species had similar dark respiration rates and light compensation points, and water-use and nitrogen-use efficiencies were inversely related between species. Species showed no differences in leaf osmotic potentials at full turgor. Calathea roots were deeper and had tuberous swellings.Leaf-level assimilation and potential water loss are consistent with where these species are found, but photosynthetic acclimation to high light does not reflect both species' abilities to grow and reproduce in gaps. Pleiostachya's gap-dependent, rapid growth and reproduction require high rates of carbon gain in short-lived leaves, which can amortize their cost quickly. High rates of water loss are associated with reduced longevity during drought. Calathea's roots may confer greater capacitance, while its leaves are durable, long-lived and have lower water loss, permitting persistence long after gap closure.

Oxygen isotope ratio stratification in a tropical moist forest.

Oxygen isotope ratios were determined in leaf cellulose from two plant species at Barro Colorado (Republic of Panama) in 4 different plots, two of which were undergoing an irrigation treatment during the dry season. There is a gradient in δ18O values of leaf cellulose from the understory to canopy leaves, reflecting the differences in relative humidity between these two levels of the forest. This gradient is most pronounced in irrigated plots. For irrigated plots there was a highly significant correlation between δ18O and δ13C values, which was not observed in control plots. This relationship can be explained by humidity controlling stomatal conductance. Low humidity affects δ18O values of leaf water during photosynthesis, which isotopically labels cellulose during its synthesis. Low humidity also decreases stomatal conductance, which affects discrimination against carbon-13 by photosynthetic reactions, thus affecting the δ13C values of photosynthates. WUE values calculated by using plant carbon and oxygen isotope ratios were similar to those observed with gas exchange measurements in other tropical and temperate area. Thus the concurrent analysis of carbon and oxygen isotope ratios of leaf material can potentially be useful for long term estimation of assimilation and evapotranspiration regimes of plants.

Photosynthetic acclimation and water-use efficiency of three species of understory herbaceous bamboo (Gramineae) in Panama.

To assess the role of photosynthetic acclimation in the response of tropical understory herbs to treefall light gaps, photosynthetic response curves were determined for three species of herbaceous bamboo growing in treatments of sun and shade at Barro Calorado Island, Panama. Increased maximum photosynthetic capacity did not always accompany higher ramet production in the sun treatment. Pharus latifolius reproduced abundantly in both treatments, and produced more ramets and developed higher maximum photosynthetic capacity under higher irradiance. Streptochaeta spicata also produced a high percentage of reproductive ramets in both treatments and produced more ramets in the sun, did not show any significant differences in photosynthetic parameters between treatments. Streptochaeta sodiroana did not change maximum photosynthetic capacity in the sun, and had higher photosynthetic efficiency and lower mortality in the shade. Stable carbon isotope composition of leaves indicated that all three species developed higher water-use efficiency under higher irradiance. Photosynthetic flexibility may contribute to the ability of P. latifolius to reproduce in treefall gaps, whereas S. spicata and S. sodiroana may maintain the ability to fix carbon efficiently in low irradiance even when growing or persisting in gaps.