Heterogeneous Light Environment Research Articles

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

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

Physiological responses of seedlings of different Quercus castaneifolia C.A. Mey. provenances to heterogeneous light environments

In this study, five biochemical traits obtained for seedlings of five different provenances of Quercus castaneifolia C.A. Meyer were used to evaluate the available light environment using instantaneous electronic readings as a standard. Experimental design was executed under controlled conditions at six different irradiance levels (10, 20, 30, 50, 60, and 70% of full light). Results show that among total chlorophyll (Chlmass), chlorophyll a to b ratio (Chla:Chlb), N, chlorophyll to N ratio (Chl:N), and peroxidase (POD) quantitative activity, the latter provides the highest realistic correlation with available irradiance. Ranking irradiance treatments on a Chlmass basis, Chl:N ratio, and Chla:Chlb ratio do not consistently distinguish different irradiance levels either within or between provenances, while the ranking produced by POD was an accurate reflection of the degree of irradiance within the shade houses, especially at 10-30%.

Roles of pathogens on replacement of tree seedlings in heterogeneous light environments in a temperate forest: a reciprocal seed sowing experiment

Summary In forest communities, the Janzen–Connell (J‐C) hypothesis proposes that species diversity is maintained by non‐competitive distance‐ and/or density‐dependent seedling mortality caused by host‐specific natural enemies. However, the effects of pathogen associations from nearby conspecifics versus heterospecifics remain unknown in spatially heterogeneous light environments. Seeds of hardwood species Cornus controversa (Cornus) and Prunus grayana (Prunus) were sown beneath 6–7 Cornus and Prunus adults in both the forest understory (FU) and in gaps (Gap) created by felling all woody vegetation near the focal adults. Seedling growth, mortality, killing agents (e.g. pathogens that cause damping‐off and leaf diseases), and root infection by arbuscular mycorrhizal fungi (AMF) were investigated. We found strong habitat effects on the expression of soil fungi beneath both tree species. Seedling mortality caused by soil‐borne damping‐off pathogens was greater in FU than in Gap, and AMF infection, which enhanced relative seedling growth rate, was greater in Gap than in FU. Seedling mortality caused by damping‐off pathogens did not differ between Cornus and Prunus seedlings beneath the adults of conspecific or heterospecific adults in both FU and Gap, suggesting little distance‐dependence or host preference in the fungus. Beneath the adults of Cornus and Prunus, the most prevalent leaf diseases were zonate leaf blight and angular leaf spot caused by the airborne pathogenic fungi Haradamyces foliicola and Phaeoisariopsis pruni‐grayanae, respectively. Although these pathogens attacked the seedlings of both species, conspecific seedlings (i.e. home) showed more severe leaf damage, earlier leaf shedding and/or less defensive behaviour (cell wall defence) relative to heterospecific seedlings (i.e. away), suggesting negative distance‐dependent attack (i.e. host preference) for these leaf diseases. As a result, greater seedling mortality was observed for conspecific seedlings under both FU and Gap treatments. Synthesis. In the temperate forest, the J‐C hypothesis is largely mediated through the strong negative influence of airborne leaf diseases rather than through soil‐borne damping‐off pathogens. We found that airborne diseases demonstrated distance‐dependent host preferences, which led to greater conspecific seedling damage regardless of environmental light conditions.

Crown asymmetry in high latitude forests: disentangling the directional effects of tree competition and solar radiation

Light foraging by trees is a fundamental process shaping forest communities. In heterogeneous light environments this behavior is expressed as plasticity of tree growth and the development of structural asymmetries. We studied the relative influence of neighborhood structure and directional solar radiation on horizontal asymmetry of tree crowns in late‐successional high latitude (67–68°N) forests in northern Fennoscandia. We described crown asymmetries as crown vectors (i.e. horizontal vectors from stem center to crown center), which we obtained from canopy maps based on crown perimeter measurements in the field. To disentangle the influence of the two main determinants, inter‐tree competition and directionality of above‐canopy solar radiation at high latitudes, we applied circular statistical models, utilizing cylindrical distributions, to these data consisting of orientations and intensities of crown asymmetry. At the individual tree level, our model predicted crown asymmetry vectors from the current stand structure, and the predictions became better when the intensity of asymmetry (i.e. crown vector length) was higher. Competition was the main determinant of crown asymmetry for 2/3 of trees, and the model predictions improved when we incorporated the directionality of solar radiation. At the stand‐level, these asymmetries had resulted in a small increment of the projected canopy area and an increased regularity of spatial structure. Our circular statistical modelling approach provided a quantitative evaluation of the relative importance of directionality of solar radiation and neighborhood stand structure, showing how both of these factors play a role in formation of crown asymmetries in high latitude forests. This approach further demonstrated the applicability of circular statistical modeling in ecological studies where the response variable has both orientation and intensity.

Adaptive loss of color polymorphism and character displacements in sympatric Mnais damselflies

A geographical survey of two Mnais damselfly species in the Kinki area of Japan showed evidence for character displacements when the two species were found in sympatry. Mnais costalis, a species that has polymorphic male mating types of orange-winged territorial and clear-winged non-territorial morphs (hereafter abbreviated to orange and clear morphs respectively) in allopatry often shifted to having monomorphic orange morphs in sympatry. The mean body size of orange morphs was consistently larger than that of clear morph in allopatry. The mean body size of the sympatric orange morphs was even larger than that of allopatric orange morphs. By contrast, Mnais pruinosa, a species that also has two morphs of large orange and small clear morphs in allopatry, shifted to having monomorphic clear morphs in sympatry. The mean body size of the sympatric clear morphs was smaller than that of allopatric clear morphs. Divergence was also detected in the preference for habitat insolation conditions between sympatric Mnais damselflies. Both species in allopatric regions prefer half-light forest habitats, while in sympatric regions they showed diversified habitat preference: M. costalis preferred sunny habitats while M. pruinosa preferred shady habitats. Multiple character displacements in signal traits and habitat preference emerged in heterogeneous forest light environments are likely to have synergistic effects on the reproductive isolation of the two species.

Adaptive Molecular Evolution of PHYE in Primulina, a Karst Cave Plant.

Limestone Karst areas possess high levels of biodiversity and endemism. Primulina is a typical component of Karst endemic floras. The high species richness and wide distribution in various Karst microenvironments make the genus an idea model for studying speciation and local adaptation. In this study, we obtained 10 full-length sequences of the phytochrome PHYE from available transcriptome resources of Primulina and amplified partial sequences of PHYE from the genomic DNA of 74 Primulina species. Then, we used maximum-likelihood approaches to explore molecular evolution of PHYE in this Karst cave plant. The results showed that PHYE was dominated by purifying selection in both data sets, and two sites were identified as potentially under positive selection. Furthermore, the ω ratio varies greatly among different functional domains of PHYE and among different species lineages. These results suggest that potential positive selection in PHYE might have played an important role in the adaption of Primulina to heterogeneous light environments in Karst regions, and different species lineages might have been subjected to different selective pressures.

Photosynthetic acclimation within individual Typha latifolia leaf segments

Growing Typha latifolia leaf segments traverse a wide range of light environments as they are pushed upward from basal meristems through sediment, water, and dense litter and leaf layers. We used light transfer experiments in a greenhouse to investigate the photosynthetic acclimation of individual Typha leaf segments. Fully expanded leaf segments exhibited strong and symmetrical acclimation to increases or decreases in light. The photosynthetic light response curves of shade grown segments that were transferred to high light acclimated within 10–15 days to match those of segments that grew and remained in high light. Acclimation was highly localized; the light response curves of shade grown segments that remained shaded did not change, even as adjacent segments acclimated to high light. Leaf segments exposed to various treatments did not differ significantly in leaf mass per area; Typha leaves are apparently morphologically preformed to function in high light and allow high photosynthetic capacity, regardless of the growth environment. Likewise, nitrogen content did not differ significantly between treatments, and photosynthetic acclimation may involve a reallocation or activation of nitrogen within a leaf segment, rather than a net nitrogen translocation into or out of a segment. We interpret these patterns as an adaptive strategy that maximizes carbon gain by monocot leaves growing in a vertically heterogeneous light environment

Impact of habitat environment onPotamogeton perfoliatusL. morphology and its within-plant variability in Lake Balaton

Plastic effect of environmental factors acting on an aquatic submerged plant, Potamogeton perfoliatus L. at the plant-level (nutrient availability) and the leaf-level (light intensity) at different sites in Lake Balaton was studied. Light-dependent morphological traits (foliar morphology and internode length) of P. perfoliatus were measured and analysed across the environmental gradients of the lake. The size of leaves was influenced by both trophic state and light environment: nutrient surplus increased the size of leaves by ∼29%, whereas a more heterogeneous light environment resulted in 15% larger leaves. The light environment influenced shoot morphology (internode length) to a greater extent than nutrient surplus (38% vs. 19%). Contrary to this, within-plant morphological variability was significantly higher (41%) at the nutrient limiting sites as a result of diversification effect of the leaf-level environmental factor, light. Foliar parameters and within-plant variability showed correlation only with the total N content of the sediment. Appearance of P. perfoliatus is shaped by counteracting effects: within-plant differentiation, promoted by leaf-level environmental sensitivity and within-plant homogenization triggered by perception of the surroundings at plant-level. Both light attenuation, stimulating an increase of morphological variability, and nutrient surplus, initiating the stabilization of morphological parameters, could have adaptive advantages. The variability of leaf size leads to diversification of foliar parameters, thus increasing the efficiency of light harvest at low-nutrient sites and making responses to changes in the light environment more dynamic. These results suggest that leaf-level-induced diversification is counteracted by the standardization effect triggered by plant-level environmental factors.

Concentrative nitrogen allocation to sun-lit branches and the effects on whole-plant growth under heterogeneous light environments

We investigated the nitrogen and carbohydrate allocation patterns of trees under heterogeneous light environments using saplings of the devil maple tree (Acer diabolicum) with Y-shaped branches. Different branch groups were created: all branches of a sapling exposed to full light (L-branches), all branches exposed to full shade (S-branches), and half of the branches of a sapling exposed to light (HL-branches) and the other half exposed to shade (HS-branches). Throughout the growth period, nitrogen was preferentially allocated to HL-branches, whereas nitrogen allocation to HS-branches was suppressed compared to L- and S-branches. HL-branches with the highest leaf nitrogen content (N(area)) also had the highest rates of growth, and HS-branches with the lowest N(area) had the lowest observed growth rates. In addition, net nitrogen assimilation, estimated using a photosynthesis model, was strongly correlated with branch growth and whole-plant growth. In contrast, patterns of photosynthate allocation to branches and roots were not affected by the light conditions of the other branch. These observations suggest that tree canopies develop as a result of resource allocation patterns, where the growth of sun-lit branches is favoured over shaded branches, which leads to enhanced whole-plant growth in heterogeneous light environments. Our results indicate that whole-plant growth is enhanced by the resource allocation patterns created for saplings in heterogeneous light environments.

Light gradients and optical microniches in coral tissues

Light quantity and quality are among the most important factors determining the physiology and stress response of zooxanthellate corals. Yet, almost nothing is known about the light field that Symbiodinium experiences within their coral host, and the basic optical properties of coral tissue are unknown. We used scalar irradiance microprobes to characterize vertical and lateral light gradients within and across tissues of several coral species. Our results revealed the presence of steep light gradients with photosynthetically available radiation decreasing by about one order of magnitude from the tissue surface to the coral skeleton. Surface scalar irradiance was consistently higher over polyp tissue than over coenosarc tissue in faviid corals. Coral bleaching increased surface scalar irradiance by ~150% (between 500 and 700 nm) relative to a healthy coral. Photosynthesis peaked around 300 μm within the tissue, which corresponded to a zone exhibiting strongest depletion of scalar irradiance. Deeper coral tissue layers, e.g., ~1000 μm into aboral polyp tissues, harbor optical microniches, where only ~10% of the incident irradiance remains. We conclude that the optical microenvironment of corals exhibits strong lateral and vertical gradients of scalar irradiance, which are affected by both tissue and skeleton optical properties. Our results imply that zooxanthellae populations inhabit a strongly heterogeneous light environment and highlight the presence of different optical microniches in corals; an important finding for understanding the photobiology, stress response, as well as the phenotypic and genotypic plasticity of coral symbionts.

Costs versus risks: Architectural changes with changing light quantity and quality in saplings of temperate rainforest trees of different shade tolerance

AbstractLight requirements and functional strategies of plants to cope with light heterogeneity in the field have a strong influence on community structure and dynamics. Shade intolerant plants often show a shade avoidance strategy involving a phytochrome‐mediated stem elongation in response to changes in red : far red ratio, while shade‐tolerant plants typically harvest light very efficiently. We measured plant size, stem diameter, internode and leaf lengths in randomly chosen saplings of 11 woody species differing in their shade tolerance in both a secondary forest and an old‐growth temperate evergreen rainforest in southern Chile. We also recorded the irradiance spectrum and the diffuse and direct light availabilities at each sampling point. Significant differences were found for the mean light environment of the saplings of each species, which also differed in basal stem diameter, internode length and leaf length, but not in plant height. Both plant slenderness (plant height/stem diameter) and mean internode length increased with increasing light availability, but no relationship was found between any of these two traits and red : far red ratio. The change in plant slenderness with light availability was of lesser magnitude with increasing shade tolerance of the species, while internode change with light availability increased with increasing shade tolerance of the species. Shade tolerators afford higher costs (thicker stems and plants), which render more biomechanically robust plants, and respond more to the light environment in a trait strongly influencing light interception (internode length) than shade intolerant species. By contrast, less shade‐tolerant plants afforded higher risks with a plastic response to escape from the understorey by making thinner plants that were biomechanically weaker and poorer light interceptors. Thus, species differing in their shade tolerances do differ in their plastic responses to light. Our results contribute to explain plant coexistence in heterogeneous light environments by improving our mechanistic understanding of species responses to light.

Self-shading in cork oak seedlings: Functional implications in heterogeneous light environments

The high self-shading found in Quercus suber seedlings has been interpreted as a feature common for plants growing in high light environments. But many studies reveal that Q. suber has high survival rates under low-light conditions, so a high degree of self-shading could be the consequence of a foliage composed of many small leaves, with no drawbacks for coping with low light. A characterization of the light environment in a Q. suber stand together with a study of photosynthetic parameters of full sunlight-exposed (FSLE) and self-shaded (SS) leaves were carried out to tackle this apparent contradiction. Although the number of sunflecks longer than 120 min during the 3 months of measurements was low, the occurrence of at least one sunfleck longer than 120 min per day in the understory of the forest studied was very common. Sunflecks shorter than 30 min promoted an increase in net photosynthesis ( A) in FSLE leaves, but not in SS leaves. However, sunflecks longer than 60 min led to a very strong decrease in A and in actual photosystem II efficiency (Φ PSII) in FSLE leaves, when compared to sunflecks shorter than 30 min. In SS leaves, changes were, again, negligible. The multi-layered foliage of Q. suber seedlings allowed i) FSLE leaves to obtain the maximum photosynthetic yield for short sunflecks, and ii) SS leaves to increase their contribution to the photosynthesis of the whole plant for long sunflecks, thus, optimizing the use of light by FSLE and SS leaves during short and long sunflecks respectively. Therefore, shoot architecture of Q. suber seedlings involving high levels of self-shading allows to adequately cope with the low but highly heterogeneous light conditions of the understory, particularly when sunflecks of contrasting durations take place as it is frequently the case for evergreen Mediterranean forests.

Effects of forest fragmentation on the seedling recruitment of a tropical herb: assessing seed vs. safe‐site limitation

Studies simultaneously evaluating the importance of safe-site and seed limitation for plant establishment are rare, particularly in human-modified landscapes. We used spatially explicit neighborhood models together with data from 10 0.5-ha mapped census plots in a fragmented landscape spanning 1000 km2 to (1) evaluate the relative importance of seed production, dispersal, and safe-site limitation for the recruitment of the understory herb Heliconia acuminata; and (2) determine how these processes differ between fragments and continuous forests. Our analyses demonstrated a large degree of variation in seed production, dispersal, and establishment among and within the 10 study plots. Seed production limitation was strong but only at small spatial scales. Average dispersal distance was less than 4 m, leading to severe dispersal limitation at most sites. Overall, safe-site limitation was the most important constraint on seedling establishment. Fragmentation led to a more heterogeneous light environment with negative consequences for seedling establishment but had little effect on seed production or dispersal. These results suggest that the effects of fragmentation on abiotic processes may be more important than the disruption of biotic interactions in driving biodiversity loss in tropical forests, at least for some functional groups. These effects may be common when the matrix surrounding fragments contains enough tree cover to enable movement of dispersers and pollinators.

Biomass allocation and photosynthetic responses of lianas and pioneer tree seedlings to light

Lianas are frequently considered as light demanding plants due to their proliferation in gaps and forest edges. Since lianas are exposed to a very heterogeneous light environment, they could be expected to express morphological and physiological plasticity in response to changes in the light environment, as high as that found in pioneer trees. We compared the biomass allocation and photosynthetic responses of seedlings of three species of lianas and two species of pioneer trees to increased light availability. Seedlings were transferred from medium (4–5 mol m −2 d −1 ) to high irradiance (12–15 mol m −2 d −1 ) in a controlled environment. In general the three liana species allocated fewer resources to the stem in comparison with the trees. The difference in the response between irradiance regimes was similar among the species, with no strong differences between trees and lianas probably due the early stage of the plants. With increase in irradiance plants accumulated more biomass, allocated more resources to the roots and less to the leaves, reduced the leaf area ratio (LAR) and specific leaf area (SLA). The photosynthetic rates recorded were not related to the rates of growth as measured by the increase in dry biomass (RGRm). Regardless of the life form, plants under higher irradiance increased their light compensation point (Lcp) and attained light saturation (Lsp) at higher levels of irradiance, while the saturated photosynthetic rate (A max ) did not show a clear pattern, and dark respiration (R d ) and quantum yield (Q) were not affected by the transference. The understanding of liana and tree seedlings responses to the light environment may have important implications in the dynamics of tropical forest regeneration.

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments.

The influence of light on the growth of watercress (Nasturtium officinale R. Br.)

In stream ecosystems, the growth of aquatic primary producers is affected by spatial and temporal variations in the riparian canopy, which can influence the availability of light resources. Aquatic plants can acclimate to low light environments by employing a suite of morphological or physiological mechanisms to increase light capture or photosynthetic efficiency. Some species may also use alternate types of propagules to colonize environments with heterogeneous light environments. In a greenhouse experiment we examined the morphological and physiological response of watercress (Nasturtium officinale R. Br.) to a gradient of increasing light levels, which ranged from 7% ambient light to full sunlight. We also determined if watercress seedlings and vegetative fragments differed in their growth response to increasing light levels. Total biomass and root biomass of seedlings and vegetative fragments decreased with decreasing light levels. The difference in plant biomass across treatments was due to morphological changes in total canopy area and leaf area, both of which increased with decreasing light levels. Seedlings and vegetative fragments did not differ in their response to light availability, but vegetative fragments had higher final biomass as a result of higher initial biomass. Physiological acclimation to low light levels appears to be of secondary importance for watercress as the concentrations of total chlorophyll, chlorophyll a, chlorophyll b, and chlorophyll a:b did not differ among light levels or between seedlings and vegetative fragments. Seedlings and vegetative fragments grown under high light levels had a greater percentage of carbon and a lower percentage of nitrogen than plants grown under low light conditions. The results of this study indicate that watercress displays considerable morphological plasticity and acclimates to low light conditions primarily by increasing leaf area and canopy surface area. There is no evidence that the type of watercress propagule (seedling vs. vegetative fragment) imparts any growth advantage in low light environments and watercress grown from either type of propagule showed no differences in their morphological or physiological responses to varying light regimes.

Printing Light Harvesting Biological “Devices” and other Functional Materials Applicable to Organic Photovoltaics

Many processing steps in the manufacturing of organic photovoltaics require solution processing. Inkjet printing is a facile method for depositing solutions into patterned thin films. Piezoelectric inkjet printing is a thermally consistent process, thereby protecting the tertiary structure of organic molecules that convert optical power to electrical power. We have taken several steps back from device manufacturing and worked out the details of inkjet printing light harvesting bacteria, more specifically photosynthetic cyanobacteria. Each individual single-celled organism is a functioning biological photovoltaic device that works with only the input of water as an electron donor for the photosynthetic reaction. In addition, these cells harvest photons from a variety of wavelengths so that they function more efficiently in heterogeneous light environments. An added bonus is that their byproduct is pure oxygen and conversely, they use C02 as a food source, thus removing carbon dioxide from the atmosphere. In these biological organic photovoltaic devices, a series of photochemical reactions occur through a thylakoid-containing internal membrane structure underneath their gelatinous cell wall. We will show the successful thin film patterning of these cells, and we will also show their light emission characteristics. In addition, we will show their carbohydrate production yields highlighting their ability to use light to form energy that can be converted easily into electricity. Finally, we will show other patterned thinfilms of organic photovoltaic relevant materials including conductive silver used in cathodes, carbon nanotubes used as transparent conductors, conjugated polymers used in the active layer and quantum dots, band gap acceptor materials that also function in non-white light environments. Finally, this talk will discuss the impact of inkjet printing on the photovoltaic market and highlight the research efforts of leaders in this field.

Estimation of light interception in research environments: a joint approach using directional light sensors and 3D virtual plants applied to sunflower (Helianthus annuus) and Arabidopsis thaliana in natural and artificial conditions.

Light interception is a major factor influencing plant development and biomass production. Several methods have been proposed to determine this variable, but its calculation remains difficult in artificial environments with heterogeneous light. We propose a method that uses 3D virtual plant modelling and directional light characterisation to estimate light interception in highly heterogeneous light environments such as growth chambers and glasshouses. Intercepted light was estimated by coupling an architectural model and a light model for different genotypes of the rosette species Arabidopsis thaliana (L.) Heynh and a sunflower crop. The model was applied to plants of contrasting architectures, cultivated in isolation or in canopy, in natural or artificial environments, and under contrasting light conditions. The model gave satisfactory results when compared with observed data and enabled calculation of light interception in situations where direct measurements or classical methods were inefficient, such as young crops, isolated plants or artificial conditions. Furthermore, the model revealed that A. thaliana increased its light interception efficiency when shaded. To conclude, the method can be used to calculate intercepted light at organ, plant and plot levels, in natural and artificial environments, and should be useful in the investigation of genotype-environment interactions for plant architecture and light interception efficiency.

Trade-offs Between Seedling Growth and Survival in Deciduous Broadleaved Trees in a Temperate Forest

In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest. The relationship between seedling survival and seedling relative growth rate in mass (RGR(M)) or height (RGR(H)) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes. Within each species, both RGR(M) and RGR(H) usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGR(M) and RGR(H) in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG. The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.

Natural selection on ecophysiological traits of a fern species in a temperate rainforest

Unlike other species of the genus Blechnum, the fern Blechnum chilense occurs in a wide range of habitats in Chilean temperate rainforest, from shaded forest understories to abandoned clearings and large gaps. We asked if contrasting light environments can exert differential selection on ecophysiological traits of B. chilense. We measured phenotypic selection on functional traits related to carbon gain: photosynthetic capacity (Amax), dark respiration rate (Rd), water use efficiency (WUE), leaf size and leaf thickness in populations growing in gaps and understorey environments. We assessed survival until reproductive stage and fecundity (spo- rangia production) as fitness components. In order to determine the potential evo- lutionary response of traits under selection, we estimated the genetic variation of these traits from clonally propagated individuals in common garden experiments. In gaps, survival of B. chilense was positively correlated with WUE and negatively correlated with leaf size. In contrast, survival in shaded understories was positively correlated with leaf size. We found positive directional fecundity selection on WUE in gaps population. In understories, ferns of lower Rd and greater leaf size showed greater fecundity. Thus, whereas control of water loss was optimized in gaps, light capture and net carbon balance were optimized in shaded understories. We found a significant genetic component of variation in WUE, Rd and leaf size. This study shows the potential for evolutionary responses to heterogeneous light environments in functional traits of B. chilense, a unique fern species able to occupy a broad successional niche in Chilean temperate rainforest.