Shade promotes thorn development in a tropical liana, Artabotrys hexapatalus (Annonaceae)

The liana Artabotrys hexapetalus (L.f.) Bhand., which is widely planted in the Tropics and native to African rain forests, produced new reiterations (new leader shoots) normally and after damage induced by Hurricane Andrew (August 24, 1992). In each new orthotropic shoot, there is a gradient in lateral branch structures from basal thorns, to vegetative leafy branches, to distal leafy flowering branches. We noted that reiterations developing in shade had more thorns than similar reiterations developing in full sun. Tents with clear (66% photosynthetically active radiation [PAR]) and shaded plastic film (12%–14% PAR) were placed over nodes when the axillary buds began to expand to produce reiteration shoots. After 2 mo of growth inside the tents and in the open, the types of lateral outgrowths (thorn vs. branch) were recorded. Shoots in spectrally neutral shade (red to far red of full sun) and spectrally altered shade (red to far red of canopy shade) produced significantly more thorns at the lower nodes of the shoots as compared to those in full sun. Shoots in control clear plastic tents were the same as those in full sun. We conclude that the fate of lateral bud development is controlled by irradiance (light level) but not by light quality. Increased thorn production in shade could be advantageous to plants growing in the deep shade of rain forests. Thorns in the self-shaded regions of the plant, and well below the forest canopy, could aid in protection from herbivory and in climbing by acting as hooks.

The developmental responses of papaya leaves to simulated canopy shade

The developmental responses of plants to shade underneath foliage are influenced by reductions in irradiance and shifts in spectral quality (characterized by reductions in the quantum ratio of red to far-red wavelengths, R:FR). Previous research on the influence of shadelight on leaf development has neglected the reductions in R:FR characteristic of foliage shade, and these studies have almost certainly underestimated the extent and array of developmental responses to foliage shade. We have studied the effects of reduced irradiance and R:FR on the leaf development of papaya (Carica papaya L., Caricaceae). Using experimental shadehouses, replicates of plants grown in high light conditions (0.20 of sunlight and R:FR = 0.90) were compared to low light conditions (0.02 of sunlight) with either the spectral quality of sunlight (R:FR = 0.99) or of foliage shade (F:FR = 0.26). Although many characteristics, such as leaf thickness, specific leaf weight, stomatal density, palisade parenchyma cell shape, and the ratio of mesophyll air surface/leaf surface were affected by reductions in irradiance, reduced R:FR contributed to further changes. Some characters, such as reduced chlorophyll a/b ratios, reduced lobing, and greater internode length, were affected primarily by low R:FR. The reduced R:FR of foliage shade, presumably affecting phytochrome equilibrium, strongly influences the morphology and anatomy of papaya leaves.

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.

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.

Leaf Optical Properties of Rainforest Sun and Extreme Shade Plants

The opticalp ropertieso f the leaves of twelve tropicals un speciesa nd thirteent ropicale xtreme shade species were examinedw ith an integratings pherea ttachedt o a spectroradiometerM. easurements of diffuse reflectance and transmittance allowed calculations of absorptance, 350- 1,100 nm. Althoughs ome shade species absorbedh igherp ercentageso f quantumf lux densities for photosynthesis (400-700 nm, PPFD) than the mean for the sun species, the sun and shade species as groups were not significantly different from each other: 90.2, S.D. 3.6% for shade species and 88.6, S.D. 2.4% for the sun species. The groups of species did not differ in total absorptance of energy 350-1,100 nm. Furthermore, the sun and shade species were identical in theirs hifto f absorptancea t wavelengthsb etween6 50 and 750 nm. The anthocyanicc oloration of the leaf undersurfaceso f two species polymorphicf or this characteristic( Trionela hirsuta and Ischnosciphonp ruinosus)i s correlatedw ith increaseda bsorptancea t the uppere nd of the action spectrum of photosynthesis. Although sun and shade species have similar optical properties, the energy investment (as documented by dry wt per unit area of leaf surface) is much less for the shade species.

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.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Abstract Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

Effects of irradiance and spectral quality on leaf structure and function in seedlings of two Southeast Asian Hopea species

We studied the development of leaf characters in two Southeast Asian dipterocarp forest trees under different photosynthetic photon flux densities (PFD) and spectral qualities (red to far-red, R:FR). The two species, Hopea helferi and H. odorata, are taxonomically closely related but differ in their ecological requirements; H. helferi is more drought tolerant and H. odorata more shade tolerant. Seedlings were grown in replicated shadehouse treatments of differing PFD and R:FR. We measured or calculated (1) leaf and tissue thicknesses; (2) mesophyll parenchyma, air space, and lignified tissue volumes; (3) mesophyll air volumes (Vmes/Asurf) and surfaces (Ames/Asurf); (4) palisade cell length and width; (5) chlorophyll/cm2 and a/ b; (6) leaf absorption; and (7) attenuance/absorbance at 652 and 550 nm. These characters varied in response to light conditions in both taxa. Characters were predominantly affected by PFD, and R:FR slightly influenced many characters. Leaf characters of H. odorata were more plastic in response to treatment conditions. Characters were correlated with each other in a complex fashion. Variation in leaf anatomy is most likely a consequence of increasing leaf thickness in both taxa, which may increase mechanical strength and defense against herbivory in more exposed environments. Variation in leaf optical properties was most likely affected by pigment photo-bleaching in treatments of more intense PFD and was not correlated with Amax. The greater plasticity of leaf responses in H. odorata helps explain the acclimation over the range of light conditions encountered by this shade-tolerant taxon. The dense layer of scales on the leaf undersurface and other anatomical characters in H. helferi reduced gas exchange and growth in this drought-tolerant tree.

Using bottles to study shade responses of seedlings and other plants

We briefly review the nature of light and its effects on plants, and then describe an inexpensive experimental system for studying the effects of shade, specifically the contributions of reduced intensity ("quantity") and the altered spectral distribution of foliage shade ("quantity") on the development of seedlings and other plants. This system has been devised to be safe to construct, inexpensive in its use of readily available materials, and appropriate for a range of student grade levels, from ~grade six to university courses in botany. We conclude by suggesting a range of experiments this system will allow. An advantage of this system is that it promotes the study of the responses of a large range of plants, most completely unstudied for these responses.

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.

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Two species of mangrove trees of Indo- Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year-1. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximumnumber of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m2 of mangrove forest with stem densities of 24,735 ha-1. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year-1. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.

The effects of climate on the growth and physiology of tropical rainforest trees

Tropical rainforests account for more than a third of global net primary production and contain more than half of the global forest carbon. Though these forests are a disproportionately important component of the global carbon cycle, the relationship between rainforest productivity and climate remains poorly understood. Understanding the link between current climate and rainforest tree stem diameter increment, a major constituent of forest productivity, will be crucial to efforts at modeling future climate and rainforest response to climate change. This work reports the physiological and stem growth responses to micrometeorological and phenological states of ten species of canopy trees in a Costa Rican wet tropical forest at sub-annual time intervals. I measured tree growth using band dendrometers and estimated leaf and reproductive phenological states monthly. Electronic data loggers recorded xylem sap flow (an indicator of photosynthetic rate) and weather at half-hour intervals. An analysis of xylem sap flow showed that physiological responses were independent of species, which allowed me to construct a general model of weather driven sap flow rates. This model predicted more than eighty percent of climate driven sap flow variation. Leaf phenology influenced growth in three of the ten species, with two of these species showing a link between leaf phenology and weather. A combination of rainfall, air temperature, and irradiance likely provided the cues that triggered leaf drop in Dipteryx panamensis and Lecythis ampla. Combining the results of the sap flow model, growth, and the climate measures showed tree growth was correlated to climate, though the majority of growth variation remained unexplained. Low variance in the environmental variables and growth rates likely contributed to the large amount of unexplained variation. A simple model that included previous growth increment and three meteorological variables explained from four to nearly fifty percent of the growth variation. Significant growth carryover existed in six of the ten species, and rainfall was positively correlated to growth in eight of the ten species. Minimum nighttime temperature was also correlated to higher growth rates in five of the species and irradiance in two species. These results indicate that tropical rainforest tree trunks could act as carbon sinks if future climate becomes wetter and slightly warmer. ^

The isolation, characterization, and application of WRKY genes as useful molecular markers in tropical trees

The improvement of tropical tree crops using conventional breeding methods faces challenges due to the length of time involved. Thus, like most crops, there is an effort to utilize molecular genetic markers in breeding programs to select for desirable agronomic traits. Known as marker assisted breeding or marker assisted selection, genetic markers associated with a phenotype of interest are used to screen and select material reducing the time necessary to evaluate candidates. As the focus of this research was improving disease resistance in tropical trees, the usefulness of the WRKY gene superfamily was investigated as candidates for generating useful molecular genetic markers. WRKY genes encode plant-specific transcriptional factors associated with regulating plants' responses to both biotic and abiotic stress. ^ One pair of degenerate primers amplified 48 WRKY gene fragments from three taxonomically distinct, economically important, tropical tree crop species: 18 from Theobroma cacao L., 21 from Cocos nucifera L. and 9 from Persea americana Mill. Several loci from each species were polymorphic because of single nucleotide substitutions present within a putative non-coding region of the loci. Capillary array electrophoresis-single strand conformational polymorphism (CAE-SSCP) mapped four WRKY loci onto a genetic linkage map of a T. cacao F2 population segregating for resistance to witches' broom disease. Additionally, PCR primers specific for four T. cacao loci successfully amplified WRKY loci from 15 members of the Byttneriae tribe. A method was devised to allow the reliable discrimination of alleles by CAE-SSCP using only the mobility assigned to the sample peaks. Once this method was validated, the diversity of three WRKY loci was evaluated in a germplasm collection of T. cacao . One locus displayed high diversity in the collection, with at least 18 alleles detected from mobility differences of the product peaks. The number of WRKY loci available within the genome, ease of isolation by degenerate PCR, codominant segregation demonstrated in the F2 population, and usefulness for screening germplasm collections and closely related wild species demonstrates that the WRKY superfamily of genes are excellent candidates for developing a number of genetic molecular markers for breeding purposes in tropical trees. ^

Wayside Trees of Tropical Florida : a guide to the native and exotic trees and palms of Miami and tropical south Florida

Wayside Trees is an beautifully illustrated guide to Florida trees growing south of Lake Okeechobee. It covers both native and exotic species in the areas of Miami to Palm Beach on the east coast, and Naples to Fort Meyers on the west. The introduction describes environmental, cultural and economic importance of trees, while a non-technical key provides a means for even non-specialists to identify the 167 most common species. The bulk of the book consists of illustrated descriptions of the trees, arranged by plant family, and includes ecological and cultural information on each species. Lavishly illustrated with over 1200 color photographs and diagrams, the book is designed to serve homeowners, gardeners, teachers and students, as well as environmental professionals. It is also a useful guide to urban tropical trees growing outside south Florida. The authors, a botanist and a graphic artist, have 70 collective years of experience living, working, and loving the trees of south Florida.