Abaxial Anthocyanin Layer in Leaves of Tropical Rain Forest Plants: Enhancer of Light Capture in Deep Shade

The permanent pigmentation of the leaves of tropical rain forest herbs with anthocyanin has traditionally been viewed as a mechanism for enhancing transpiration by increased heat absorption. We report measurements to ?+0.1?0C on four Indo-mal- esian forest species polymorphic with respect to color. There were no detectable differences in temperature between cyanic and green leaves. In deeply shaded habitats, any temperature difference would arise from black-body infrared radiation which all leaves absorb and to which anthocyanins are transparent. Reflectance spectra of the lower leaf surfaces of these species re- vealed increased reflectance around 650-750 nm for cyanic leaves compared with green leaves of the same species. In all spe- cies anthocyanin was located in a single layer of cells immediately below the photosynthetic tissue. These observations provide empirical evidence that the cyanic layer can improve photosynthetic energy capture by back-scattering additional light through the photosynthetic tissue.

Simulating Forest Shade to Study the Developmental Ecology of Tropical Plants: Juvenile Growth in Three Vines in India

Both light quantity and quality affect the development and autoecology of plants under shade conditions, as in the understorey of tropical forests. However, little research has been directed towards the relative contributions of lowered photosynthetic photon flux density (PPFD) versus altered spectral distributions (as indicated by quantum ratios of 660 to 730 nm, or R:FR) of radiation underneath vegetation canopies. A method for constructing shade enclosures to study the contribution of these two variables is described. Three tropical leguminous vine species (Abrus precatorius L., Caesalpinia bondicela Fleming and Mucuna pruriens (L.) DC.) were grown in two shade enclosures with 3-4% of solar PPFD with either the R:FR of sunlight (1.10) or foliage shade (0.33), and compared to plants grown in sunlight. Most species treated with low R:FR differed from those treated with high R:FR in (1) percent allocation to dry leaf weight, (2) internode length, (3) dry stem weight/length, (4) specific leaf weight, (5) leaf size, and (6) chlorophyll a/b ratios. However, these plants did not differ in chlorophyll content per leaf dry weight or area. In most cases the effects of low R:FR and PPFD were additional to those of high R:FR and low PPFD. Growth patterns varied among the three species, but both low PPFD and diminished R:FR were important cues in their developmental responses to light environments. This shadehouse system should be useful in studying the effects of light on the developmental ecology of other tropical forest plants.

Water source utilization and foliar nutrient status differs between upland and flooded plant communities in wetland tree islands

Tree islands in the Everglades wetlands are centers of biodiversity and targets of restoration, yet little is known about the pattern of water source utilization by the constituent woody plant communities: upland hammocks and flooded swamp forests. Two potential water sources exist: (1) entrapped rainwater in the vadose zone of the organic soil (referred to as upland soil water), that becomes enriched in phosphorus, and (2) phosphorus-poor groundwater/surface water (referred to as regional water). Using natural stable isotope abundance as a tracer, we observed that hammock plants used upland soil water in the wet season and shifted to regional water uptake in the dry season, while swamp forest plants used regional water throughout the year. Consistent with the previously observed phosphorus concentrations of the two water sources, hammock plants had a greater annual mean foliar phosphorus concentration over swamp forest plants, thereby supporting the idea that tree island hammocks are islands of high phosphorus concentrations in the oligotrophic Everglades. Foliar nitrogen levels in swamp forest plants were higher than those of hammock plants. Linking water sources with foliar nutrient concentrations can indicate nutrient sources and periods of nutrient uptake, thereby linking hydrology with the nutrient regimes of different plant communities in wetland ecosystems. Our results are consistent with the hypotheses that (1) over long periods, upland tree island communities incrementally increase their nutrient concentration by incorporating marsh nutrients through transpiration seasonally, and (2) small differences in micro-topography in a wetland ecosystem can lead to large differences in water and nutrient cycles.

Correlates of leaf optical properties in tropical forest sun and extreme-shade plants

Thirteens hade-adaptedr ain forest species were comparedw ith twelve sun-adaptedt ropical forest species for correlates to leaf optical properties (described previously in Amer. J. Bot. 73: 1100-1108). The two samples were similar in absorptance of quanta for photosynthesis, but the shade-adaptedt axa: 1) had significantlyl ower specificl eaf weights,i ndicatinga more metabolically efficient production of surface for quantum capture; 2) synthesized less chlorophyll per unit area; and 3) used less chlorophyll for capturing the same quanta for photosynthesis. The anatomical features that best correlate with this increased efficiency are palisade cell shape and chloroplast distribution. Palisade cells with more equal dimensions have more chloroplasts on their abaxial surfaces. This dense layer of chloroplasts maximizes the light capture efficiency limited by sieve effects. The more columnar palisade cells of sun-adapted taxa allow light to pass through the central vacuoles and spaces between cells, making chloroplasts less efficient in energy capture, but allowing light to reach chloroplasts in the spongy mesophyll. Pioneer species may be an exception to these two groups of species. Three pioneer taxa included in this study have columnar palisade cells that are extremely narrow and packed closely together. This layer allows little penetration of light, but exposure of the leaf undersurface may provide illumination of spongy mesophyll chloroplasts in these plants.

Epidermal cells functioning as lenses in leaves of tropical rain-forest shade plants

A ray tracing model has been developed to investigate the possible focusing effects of the convexly curved epidermal cell walls which characterize a number of shade-adapted plants. The model indicates that such focusing occurs, resulting in higher photosynthetic photon flux densities at certain locations within the leaf. It is postulated that there will be a corresponding increase in the rate of photosynthesis. In addition, leaf reflectance measurements indicate that this is generally less for the shade plants compared with sun species and would be advantageous in increasing the efficiency of energy capture. Either effect is important for plants which must survive at extremely low light levels.

Physical and ultrastructural basis of blue leaf iridescence in four Malaysian understory plants

Iridescent blue leaf coloration in four Malaysian rain forest understory plants, Diplazium tomentosum Bl. (Athyriaceae), Lindsaea lucida Bl. (Lindsaeaceae), Begonia pavonina Ridl. (Begoniaceae), and Phyllagathis rotundifolia Bl. (Melastomataceae) is caused by a physical effect, constructive interference of reflected blue light. The ultrastructural basis for this in D. tomentosum and L. lucida is multiple layers of cellulose microfibrils in the uppermost cell walls of the adaxial epidermis. The helicoidal arrangement of these fibrils is analogous to that which produces a similar color in arthropods. In B. pavonina and P. rotundifolia the blue-green coloration is caused by parallel lamellae in specialized plastids adjacent to the abaxial wall of the adaxial epidermis. The selective advantage of this color production, if any, is unknown.

Irradiance and spectral quality affect Asian tropical rain forest tree seedling development

Iridescent blue leaf coloration in four Malaysian rain forest understory plants, Diplazium tomentosum Bl. (Athyriaceae), Lindsaea lucida Bi. (Lindsaeaceae), Begonia pavonina Ridl. (Begoniaceae), and Phyllagathis rotundifolia Bl. (Melastoma- taceae) is caused by a physical effect, constructive interference of reflected blue light. The ultrastructural basis for this in D. tomentosum and L. lucida is multiple layers of cellulose microfibrils in the uppermost cell walls of the adaxial epidermis. The helicoidal arrangement of these fibrils is analogous to that which produces a similar color in arthropods. In B. pavonina and P. rotundifolia the blue-green coloration is caused by parallel lamellae in specialized plastids adjacent to the abaxial wall of the adaxial epidermis. The selective advantage of this color production, if any, is unknown.

Pigment dynamics and autumn leaf senescence in a New England deciduous forest, eastern USA

The leaves of woody plants at Harvard Forest in Central Massachusetts, USA, changed color during senescence; 70% (62/89) of the woody species examined anatomically contained anthocyanins during senescence. Anthocyanins were not present in summer green leaves, and appeared primarily in the vacuoles of palisade parenchyma cells. Yellow coloration was a result of the unmasking of xanthophyll pigments in senescing chloroplasts. In nine red-senescing species, anthocyanins were not detectable in mature leaves, and were synthesized de novo in senescence, with less than 20 m g cm - 2 of chlorophyll remaining. Xanthophyll concentrations declined in relation to chlorophyll to the same extent in both yellow- and red-leaved taxa. Declines in the maximum photosystem II quantum yield of leaves collected prior to dawn were only slightly less in the red-senescing species, indicating no long-term protective activity. Red-leaved species had significantly greater mass/area and lower chlorophyll a / b ratios during senescence. Nitrogen tissue concentrations in mature and senescent leaves negatively correlated to anthocyanin concentrations in senescent leaves, weak evidence for more efficient nitrogen resorption in anthocyanic species. Shading retarded both chlorophyll loss and anthocyanin production in Cornus alternifolia , Acer rubrum , Acer saccharum , Quercus rubra and Viburnum alnifolium . It promoted chlorophyll loss in yellow-senescing Fagus grandifolia . A reduced red : far-red ratio did not affect this process. Anthocyanins did not increase leaf temperatures in Q. rubra and Vaccinium corymbosum on cold and sunny days. The timing of leaf-fall was remarkably constant from year to year, and the order of senescence of individual species was consistent.

Human disturbance and forest diversity in the Tansa Valley, India

We assessed the diversity of woody plants at 15 forested sites in the Tansa Valley of Thane District, in Maharashtra, India. The fewest species (11) were seen at a degraded mangrove site near the river mouth, and the greatest number (150) in the rich semi-evergreen forest on Tungar Hill. For all sites there were 141 tree, 25 shrub and 15 liana species, a total of 181 species. Excluding the mangrove site, which had no species in common with the other 14 sites, we analyzed the species distributions in detail. 2 These sites ranged in area from 4 to 30 km each, had woody floras of 89 6 6 species, and varied in intensity of human impact. Despite a history of exploitation and substantial reduction in biomass from firewood collecting, set fires and illicit tree felling, considerable plant diversity remains in the area.We found a modest increase in species richness in transects away from two villages. We observed the exploitation of the forest by the principal users, primarily of the Warli Tribe. They exploited a wide variety of forest resources (92 species), for medicines, foods, construction materials, household goods, manure and other purposes. They collected 15 items for sale. By far the single most important item collected was firewood, which dramatically reduced forest biomass within 2 km of villages. The species distributions in these forest remnants are strongly nested, mostly due to varying degrees of disturbance at individual sites. The high species diversity on Tungar Hill is most likely a relict of the earlier character of forests throughout much of the valley. It merits the highest priorities for preservation, as a refuge for Western Ghat species at the northern limits of their distributions.

Factors affecting the nodulation and growth of tropical woody legume seedlings

This study surveys the occurrence of nodulation in woody legume species in Panamá and Costa Rica, describes nodule and root characteristics, and researches host-bacteria specificity, nodulation potential of soils, and the effects of light, added nitrogen, and rhizobia and VA mycorrhizal fungi inoculation on seedling growth. I examined 83 species in 37 genera and found 80% to be nodulated. Percent nodulated species in the Caesalpinioideae, Mimosoideae, and Papilionoideae was 17, 95, and 86, respectively, with no correlation between nodule morphology and tribal classification. Nodules formed mainly at root branch points which supports epidermal breaks as an important rhizobia infection route. More non-nodulated than nodulated species had root hairs. Several species emitted volatile sulfur-containing compounds, including the toxic compound ethylmercaptan, from roots, germinating seeds, and other tissues. These emissions may have an allelopathic action against pathogens, predators, or other plants. In contrast to the general non-specificity of most legumes for rhizobia, Mimosa pigra L. was highly specific and only nodulated in flooded soils. This species' specificity, combined with a limited occurrence of its root nodule bacteria may limit its natural distribution, but its spread as an invasive weed is facilitated when fill material from rivers is deposited in other areas. ^ An experimental light level of 1.5% of full sun completely inhibited seedling nodulation, as do similar naturally low levels in forest understory. In the forest, trees and seedlings were not nodulated. in some soils with suspected high N content. For six experimental species, added N progressively increased seedling growth while decreasing nodule biomass; at the highest level of added N nodulation was completely suppressed. Species and individuals showed variation in nodule biomass at high N applications which may indicate an opportunity for genetic selection for optimal N acquisition. Rhizobia inoculation had a small positive effect on seedling shoot growth, but VA mycorrhiza inoculation overwhelmingly increased seedling size, biomass, and leaf mineral concentration. In lowland tropical forest, VA mycorrhizal colonization appears indispensable for legume nodulation because of the fungus' ability to supply P in deficient soils. This requirement makes the legume-rhizobia-mycorrhiza association obligately tripartite. ^

Seed dispersal and reproduction patterns among Everglades plants

In this study three aspects of sexual reproduction in Everglades plants were examined to more clearly understand seed dispersal and the allocation of resources to sexual reproduction—spatial dispersal process, temporal dispersal of seeds (seedbank), and germination patterns in the dominant species, sawgrass (Cladium jamaicense). Community assembly rules for fruit dispersal were deduced by analysis of functional traits associated with this process. Seedbank ecology was investigated by monitoring emergence of germinants from sawgrass soil samples held under varying water depths to determine the fate of dispersed seeds. Fine-scale study of sawgrass fruits yielded information on contributions to variation in sexually produced propagules in this species, which primarily reproduces vegetatively. It was hypothesized that Everglades plants possess a set of functional traits that enhance diaspore dispersal. To test this, 14 traits were evaluated among 51 species by factor analysis. The factorial plot of this analysis generated groups of related traits, with four suites of traits forming dispersal syndromes. Hydrochory traits were categorized by buoyancy and appendages enhancing buoyancy. Anemochory traits were categorized by diaspore size and appendages enhancing air movement. Epizoochory traits were categorized by diaspore size, buoyancy, and appendages allowing for attachment. Endozoochory traits were categorized by diaspore size, buoyancy, and appendages aiding diaspore presentation. These patterns/trends of functional trait organization also represent dispersal community assembly rules. Seeds dispersed by hydrochory were hypothesized to be caught most often in the edge of the north side of sawgrass patches. Patterns of germination and dispersal mode of all hydrochorous macrophytes with propagules in the seedbank were elucidated by germination analysis from 90 soil samples collected from 10 sawgrass patches. Mean site seed density was 486 seeds/m2 from 13 species. Most seeds collected at the north side of patches and significantly in the outer one meter of the patch edge (p = 0.013). Sawgrass seed germination was hypothesized to vary by site, among individual plants, and within different locations of a plant’s infructescence. An analysis of sawgrass fruits with nested ANOVAs found that collection site and interaction of site x individual plant significantly affect germination ability, seed viability, and fruit size (p ≤ 0.050). Fruit location within a plant’s infructescence did not significantly affect germination. As for allocation of resources to sexual reproduction, only 17.9% of sawgrass seeds germinated and only 4.8% of ungerminated seeds with fleshy endosperm were presumed viable, but dormant. Collectively, only 22% of all sawgrass seeds produced were viable.

Effect of leaf-cutting ant nests on plant growth in an oligotrophic Amazon rain forest

This study examined whether high nutrient concentrations associated with leaf-cutting ant nests influence plant growth and plant water relations in Amazon rain forests. Three nests of Atta cephalotes were selected along with 31 Amaioua guianensis and Protium sp. trees that were grouped into trees near and distant (>10 m) from nests. A 15N leaf-labelling experiment confirmed that trees located near nests accessed nutrients from nests. Trees near nests exhibited higher relative growth rates (based on stem diameter increases) on average compared with trees further away; however this was significant for A. guianensis (near nest 0.224 y−1 and far from nest 0.036 y−1) but not so for Protium sp. (0.146 y−1 and 0.114 y−1 respectively). Water relations were similarly species-specific; for A. guianensis, near-nest individuals showed significantly higher sap flow rates (16 vs. 5 cm h−1), higher predawn/midday water potentials (−0.66 vs. −0.98 MPa) and lower foliar δ13C than trees further away indicating greater water uptake in proximity to the nests while the Protium sp. showed no significant difference except for carbon isotopes. This study thus shows that plant response to high nutrient concentrations in an oligotrophic ecosystem varies with species. Lower seedling abundance and species richness on nests as compared with further away suggests that while adult plants access subterranean nutrient pools, the nest surfaces themselves do not encourage plant establishment and growth.

Population viability analyses of Chamaecrista keyensis (Leguminosae: Caesalpinioideae), a narrowly endemic herb of the lower Florida Keys: Effects of seasonal timing of fires and the urban -wildland interface

This research first evaluated the effects of urban wildland interface on reproductive biology of the Big Pine Partridge Pea, Chamaecrista keyensis, an understory herb that is endemic to Big Pine Key, Florida. I found that C. keyensis was self-compatible, but depended on bees for seed set. Furthermore, individuals of C. keyensis in urban habitats suffered higher seed predation and therefore set fewer seeds than forest interior plants. ^ I then focused on the effects of fire at different times of the year, summer (wet) and winter (dry), on the population dynamics and population viability of C. keyensis. I found that C. keyensis population recovered faster after winter burns and early summer burns (May–June) than after late summer burns (July–September) due to better survival and seedling recruitment following former fires. Fire intensity had positive effects on reproduction of C. keyensis. In contrast, no significant fire intensity effects were found on survival, growth, and seedling recruitment. This indicated that better survival and seedling recruitment following winter and early summer burns (compared with late summer burns) were due to the reproductive phenology of the plant in relation to fires rather than differences in fire intensity. Deterministic population modeling showed that time since fire significantly affected the finite population growth rates (λ). Particularly, recently burned plots had the largest λ. In addition, effects of timing of fires on λ were most pronounced the year of burn, but not the subsequent years. The elasticity analyses suggested that maximizing survival is an effective way to minimize the reduction in finite population growth rate the year of burn. Early summer fires or dry-season fires may achieve this objective. Finally, stochastic simulations indicated that the C. keyensis population had lower extinction risk and population decline probability if burned in the winter than in the late summer. A fire frequency of approximately 7 years would create the lowest extinction probability for C. keyensis. A fire management regime including a wide range of burning seasons may be essential for the continued existence of C. keyensis and other endemic species of pine rockland on Big Pine Key. ^

Seed Dispersal and Reproduction Patterns Among Everglades Plants

In this study three aspects of sexual reproduction in Everglades plants were examined to more clearly understand seed dispersal and the allocation of resources to sexual reproduction— spatial dispersal process, temporal dispersal of seeds (seedbank), and germination patterns in the dominant species, sawgrass (Cladium jamaicense). Community assembly rules for fruit dispersal were deduced by analysis of functional traits associated with this process. Seedbank ecology was investigated by monitoring emergence of germinants from sawgrass soil samples held under varying water depths to determine the fate of dispersed seeds. Fine-scale study of sawgrass fruits yielded information on contributions to variation in sexually produced propagules in this species, which primarily reproduces vegetatively. It was hypothesized that Everglades plants possess a set of functional traits that enhance diaspore dispersal. To test this, 14 traits were evaluated among 51 species by factor analysis. The factorial plot of this analysis generated groups of related traits, with four suites of traits forming dispersal syndromes. Hydrochory traits were categorized by buoyancy and appendages enhancing buoyancy. Anemochory traits were categorized by diaspore size and appendages enhancing air movement. Epizoochory traits were categorized by diaspore size, buoyancy, and appendages allowing for attachment. Endozoochory traits were categorized by diaspore size, buoyancy, and appendages aiding diaspore presentation. These patterns/trends of functional trait organization also represent dispersal community assembly rules. Seeds dispersed by hydrochory were hypothesized to be caught most often in the edge of the north side of sawgrass patches. Patterns of germination and dispersal mode of all hydrochorous macrophytes with propagules in the seedbank were elucidated by germination analysis from 90 soil samples collected from 10 sawgrass patches. Mean site seed density was 486 seeds/m2 from 13 species. Most seeds collected at the north side of patches and significantly in the outer one meter of the patch edge (p = 0.013). Sawgrass seed germination was hypothesized to vary by site, among individual plants, and within different locations of a plant’s infructescence. An analysis of sawgrass fruits with nested ANOVAs found that collection site and interaction of site x individual plant significantly affect germination ability, seed viability, and fruit size (p < 0.050). Fruit location within a plant’s infructescence did not significantly affect germination. As for allocation of resources to sexual reproduction, only 17.9% of sawgrass seeds germinated and only 4.8% of ungerminated seeds with fleshy endosperm were presumed viable, but dormant. Collectively, only 22% of all sawgrass seeds produced were viable.