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Incidence of pigment change in Werauhia (Bromeliaceae) uncorrelated with canopy cover level

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Title:
Incidence of pigment change in Werauhia (Bromeliaceae) uncorrelated with canopy cover level
Translated Title:
La incidencia del cambio del pigmento en Werauhia (Bromeliaceae) correlacionado con el nivel de cobertura del dosel ( )
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English
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Sloss, Rachelle
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Plants--Adaptation   ( lcsh )
Plant pigments   ( lcsh )
Fragmented landscapes   ( lcsh )
Monteverde Biological Station (Costa Rica)   ( lcsh )
Plantas--Adaptación
Pigmentos vegetales
Paisajes fragmentados
Estación Biológica de Monteverde (Costa Rica)
Tropical Ecology Fall 2009
Ecología Tropical Otoño 2009
Genre:
Reports   ( lcsh )
Reports

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Abstract:
The study of plant adaptation to changing habitat is particularly important in the face of continuing habitat destruction in the tropics. Habitat fragmentation will likely increase light intensity for tropical flora in these areas. Plants often respond to such changes by increasing concentrations of secondary pigment, like anthocyanins and carotenoids, as a photoprotective mechanism. Plants may also increase the ratio of chlorophyll a:b in high light because they have less need for shade-adapted accessory pigments like chlorophyll b. However, in the Werauhia bromeliads studied, average anthocyanin and carotenoid concentrations decreased while chlorophyll a:b ratio increased. The decrease in anthocyanins and carotenoids suggests a decrease in light intensity, while an increase in chlorophyll a:b ratio indicates an increase in light intensity. Due to local climate observations of the study site and the indication of the anthocyanin and carotenoid decrease, I believe that the plants experienced an overall decrease in light intensity. This suggests that the increase in chlorophyll a:b ratio found was due to other environmental factors. However, these contradictory findings suggest that further study is needed to determine effects of light intensity on Werauhia.
Abstract:
Estudios de las adaptaciones de plantas al cambio de hábitat son de particular importancia para enfrentar la continua destrucción del hábitat en los trópicos. La fragmentación del hábitat puede incrementar la intensidad de luz para la flora tropical en estas áreas. Las plantas generalmente responden a dichos cambios incrementando la concentración de pigmentos secundarios, como antocianinas y carotenoides, como un mecanismo fotoprospectivo. Las plantas también pueden incrementar la proporción de clorofila a:b en altas concentraciones de luz debido a que tienen una menor necesidad por pigmentos adaptados para la sombra como la clorofila b. Sin embargo en bromelias del género Werauhia, el promedio de antocianinas y carotenoides decrecen mientras que la proporción de clorofila a:b aumenta. La disminución en antocianinas y carotenoides sugiere una disminución en la intensidad luminosa, mientras que el aumento en la proporción de clorofila a:b indica un aumento en la misma. Debido a observaciones del clima local en el sitio de estudio y al hecho de que las antocianinas y carotenoides decrecen, yo creo que las plantas experimentan una disminución en la intensidad lumínica. Esto sugiere que el aumento en la proporción de clorofila a:b encontrado se debe a factores ambientales. Sin embargo, estos resultados contradictorios sugieren que estudios futuros son necesarios para determinar el efecto de la intensidad lumínica en Werauhia.
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Text in English.
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The study of plant adaptation to changing habitat is particularly important in the face of continuing habitat destruction in the tropics. Habitat fragmentation will likely increase light intensity for tropical flora in these areas. Plants often respond to such changes by increasing concentrations of secondary pigment, like anthocyanins and
carotenoids, as a photoprotective mechanism. Plants may also increase the ratio of chlorophyll a:b in high light because they have less need for shade-adapted accessory pigments like chlorophyll b. However, in the Werauhia bromeliads studied, average anthocyanin and carotenoid concentrations decreased while chlorophyll a:b ratio
increased. The decrease in anthocyanins and carotenoids suggests a decrease in light intensity, while an increase in chlorophyll a:b ratio indicates an increase in light intensity. Due to local climate observations of the study site and the indication of the anthocyanin and carotenoid decrease, I believe that the plants experienced an overall decrease in light intensity. This suggests that the increase in chlorophyll a:b ratio found was due to other environmental
factors. However, these contradictory findings suggest that further study is needed to determine effects of light intensity on Werauhia.
Estudios de las adaptaciones de plantas al cambio de hbitat son de particular importancia para enfrentar la continua destruccin del hbitat en los trpicos. La fragmentacin del hbitat puede incrementar la intensidad de luz para la flora tropical en estas reas. Las plantas generalmente responden a dichos cambios incrementando la concentracin de pigmentos secundarios, como antocianinas y carotenoides, como un mecanismo fotoprospectivo. Las plantas tambin pueden incrementar la proporcin de clorofila a:b en altas concentraciones de luz debido a que tienen una menor necesidad por pigmentos adaptados para la sombra como la clorofila b. Sin embargo en bromelias del gnero Werauhia, el promedio de antocianinas y carotenoides decrecen mientras que la proporcin de clorofila a:b aumenta. La disminucin en antocianinas y carotenoides sugiere una disminucin en la intensidad luminosa, mientras que el aumento en la proporcin de clorofila a:b indica un aumento en la misma. Debido a observaciones del clima local en el sitio de estudio y al hecho de que las antocianinas y carotenoides decrecen, yo creo que las plantas experimentan una disminucin en la intensidad lumnica. Esto sugiere que el aumento en la proporcin de clorofila a:b encontrado se debe a factores ambientales. Sin embargo, estos resultados contradictorios sugieren que estudios futuros son necesarios para determinar el efecto de la intensidad lumnica en Werauhia.
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Text in English.
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Plants--Adaptation
Plant pigments
Fragmented landscapes
Monteverde Biological Station (Costa Rica)
4
Plantas--Adaptacin
Pigmentos vegetales
Paisajes fragmentados
Estacin Biolgica de Monteverde (Costa Rica)
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Tropical Ecology Fall 2009
Ecologa Tropical Otoo 2009
655
Reports
720
CIEE
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t Monteverde Institute : Tropical Ecology
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u http://digital.lib.usf.edu/?m39.329



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1 Incidence of Pigment Change in Werauhia (Bromeliaceae) Uncorrelated with Canopy Cover Level Rachelle Sloss Department of Biochemistry, Biophysics, and Molecular Biology, Whitman College ABSTRACT The study of plant adaptation to changing habitat is par ticularly important in the face of continuing habitat destruction in the tropics. Habitat fragmentation will likely increase light intensity for tropical flora in these areas. Plants often respond to such changes by increasing concentrations of secondary p igment, like anthocyanins and carotenoids, as a photoprotective mechanism. Plants may also increase the ratio of chlorophyll a:b in high light because they have less need for shade adapted accessory pigments like chlorophyll b. However, in the Werauhia bro meliads studied, average anthocyanin and carotenoid concentrations decreased while chlorophyll a:b ratio increased. The decrease in anthocyanins and carotenoids suggests a decrease in light intensity, while an increase in chlorophyll a:b ratio indicates an increase in light intensity. Due to local climate observations of the study site and the indication of the anthocyanin and carotenoid decrease, I believe that the plants experienced an overall decrease in light intensity. This suggests that the increase i n chlorophyll a:b ratio found was due to other environmental factors. However, these contradictory findings suggest that further study is needed to determine effects of light intensity on Werauhia RESUMEN Estudios de las adaptaciones de plantas al cambi o de hbitat son de particular importacin para enfrentar la continua destruccin de hbitat en los trpicos. La fragmentacin del hbitat puede incrementar la intensidad de luz para la flora tropical en estas reas. Las plantas generalmente responden a dichos cambios incrementando la concentracin de pigmentos secundarios, como antocianinas y carotenoides, como un mecanismo fotoprospectivo. Plantas tambin pueden incrementar la proporcin de clorofila a:b en altas concentraciones de luz debido a que tie nen una menor necesidad por pigmentos adaptados para la sombra como la clorofila b. Sin embargo en bromelias del gnero Werauhia el promedio de antocianinas y carotenoides decrecen mientras que la proporcin de clorofila a:b aumenta. La disminucin en a ntocianinas y carotenoides sugiere una disminucin en la intesidad luminosa, mientras que el aumento en la proporcin de clorofila a:b indica un aumento en la misma. Debido a observaciones del clima local en el sitio de estudio y al hecho de que las antoc ianinas y carotenoides decrecen, yo creo que las plantas experimentan una disminucin en la intensidad lumnica. Esto sugiere que el aumento en la proporcin de clorofila a:b encontrado se debe a factores ambientales. Sin embargo, estos resultados contra dictorios sugieren que estudios futuros son necesarios para determinar el efecto de la intensidad lumnica en Werauhia. INTRODUCTION Habitat fragmentation and UV radiation increase are two major problems facing tropical forests (Hegglin and Shepherd 200 9; Opdam and Wascher 2003). Habitat fragmentation increases the amount of forest edge, thereby increasing light levels and other environmental pressures (Debinksi and Holt 2000; Skole and Tucker 1993). Hegglin and Shepherd have estimated that UV radiation will increase by 4% in the tropics before the end of the century (2009). Therefore, it is possible that tropical ecosystems will experience increases in both light penetration and UV radiation. In order to understand how tropical ecosystems will fare in th ese changing conditions, it is important to learn how plants may be able to adapt to increases in light in the tropics.

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2 It is well known that changes in light intensity cause pigment alterations in many plants (Ulrich 2008). In low light, plants need acces sory pigments to absorb additional wavelengths of light to increase photosynthetic efficiency (Chazdon et al. 1996). Therefore, plants adapted to shade often have extra chlorophyll b, which absorbs different light wavelengths than the standard chlorophyll a. Due to this extra chlorophyll b, shade plants have lower chlorophyll a:b ratios than sun plants (Chazdon 1996; Taiz and Zieger 1991; Ulrich 2008). In high light, extra pigments are needed to absorb the extra light energy entering plant cells. If this su rplus light energy is not absorbed, it can create a free radical from oxygen gas that can damage many cellular structures (Hopkins 1995). To prevent this cellular damage, plants in high light often have high carotenoid and anthocyanin concentrations. Both carotenoids and anthocyanins are photoprotective pigments that absorb the extra light energy (Rosevear et al. 2001; Taiz and Zieger 1991). Though the concentrations of all of these pigments are most affected by light levels, they can also be influenced by temperature, water, and nutrient availability (Chazdon 1996; Sullivan 1998). Plants of the Bromeliaceae family respond especially well to light intensity changes with pigment alterations (Luther 2000; Ulrich 2008). In tropical forests, bromeliads are an im portant part of the epiphyte community. They cycle nutrients and provide food and shelter for many tree dwelling animals (Luther 2000). Werauhia a genus of epiphytic bromeliad, was found to be common in the Monteverde area and was chosen to study pigment alterations due to changes in canopy cover, which I expect correlates with light intensity. Werauhia moved to areas of low canopy cover are expected to increase anthocyanin and carotenoid concentrations as well as chlorophyll a:b ratio. Plants moved to h igh canopy cover areas are expected to decrease anthocyanin and carotenoids concentrations as well as chlorophyll a:b ratio. MATERIALS AND METHODS This study was conducted in and around the Estacin Biolgica (1500 m), Monteverde, Costa Rica. Fifty five Werauhia bromeliads were collected on their branches from San Luis (1000 m) in the Monteverde area. Groups of eleven plants were placed in five treatments with varying levels of canopy cover: 0, 25, 50, 75, and 100%. Canopy cover was measured with a Robert E. Lemmon Model C spherical densitometer. Percent canopy cover was calculated at each location facing North, South, East and West, and averaged for a total canopy cover value. Initial leaf samples were collected from each plant the morning after placemen t in treatment locations to get baseline pigment concentrations. One leaf from each bromeliad was cut at the base without damaging the rest of the plant. A 4.5 cm 2 rectangle was cut from each leaf using a razor blade and a cardboard template. The mass of t he leaf rectangle was recorded using a FisherScientific T top loading balance. The leaf sample was then cut into fine leaf fragments with the razor and transferred to a labeled test tube. 2.5 ml of a mixed solution containing 2 ml of 100% acetone and .5 ml phosphate buffer (pH 6.5) was added to each test tube. The phosphate buffer was used to maintain the natural pH of chloroplasts (Wallentine 2006). The leaf acetone mixture was allowed to sit for approximately 30 minutes with mild shaking to encourage prec ipitation of photosynthetic pigments. Each mixture was centrifuged for 1 minute with a Premiere XC 1000 centrifuge to separate the leaf fragments from the pigment acetone solution. The solution was then decanted into cuvettes, which were filled by adding 1 .5 ml of 100% acetone. The pigment concentrations of each sample were measured in a Sequoia Turner Model 340 spectrophotometer at light wavelengths of 663 (chlorophyll a), 645 (chlorophyll b), 545

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3 (anthocyanins), and 480 nm (carotenoids). These samples wer e measured against a control cuvette with 100% acetone. The above methodology was repeated after 14 days. The concentrations of pigments were calculated using the following equations from Lichtenthaler and Buschmann for pigments dissolved in 100% acetone (2001). Chlorophyll a (g/g) = [11.24*Abs 663 2.04*Abs 645 ] [Purified Volume (ml)] [Mass of Leaf (g)] Chlorophyll b (g/g) = [20.13*Abs 645 4.19*Abs 663 ] [Purified Volume (ml)] [Mass of Leaf (g)] Carotenoids (g/g) = [1000*Abs 480 1 .90*Abs 645 63.14*Abs 663 ] [Purified Volume (ml)] 214*[Mass of Leaf (g)] Because no Anthocyanin equations for 100% acetone were available, one calculated for pigment in 80% aqueous acetone was used (Sims and Gamon 2002). However, absorbance value s at 537 nm and 647 nm based on prior research were used instead of 545 and 645 nm. Because of these discrepancies, the anthocyanin concentrations should be considered estimates. Anthocyanin (g/g) = {[.08173*Abs 545 .00697*Abs 645 .002228* Abs 663 ]* [Pu rified Volume (ml)]*[Molecular Weight of Anthocyanin (595 mol/g)]}/ [Mass of Leaf (g)] Using these calculated pigment concentrations, significance of pigment changes (as a percentage of total pigment after 14 days) at each treatment were determine d, as well as significance of change in chlorophyll a:b ratio. One way ANOVA tests were used to determine significance of changes for each pigment at each treatment as well as overall pigment changes. RESULTS Percent canopy cover was not found to affect pigment change for any pigment. By this, I mean that the bromeliads at 0% canopy cover did not have significantly different changes in pigment concentrations than bromeliads at 100% canopy cover. However, some overall trends in pigment change were found b etween initial and day 14 measurements when concentrations were averaged concentrations in San Luis to their altered pigment concentrations at the biological st ation.

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4 Overall, anthocyanin concentrations decreased (from an average of 508.98 to 330.97 g/g) at all canopy cover treatments between the initial sample and 14 days (Fig. 1). Carotenoid concentrations also decreased (from an average of 82.32 to 69.51 g/g) between the initial sample and after 14 days (Fig. 2). When separated by canopy cover treatment, however, decreases of carotenoids were found in plants at 25, 75 and 100%, but the plants at 0 and 50% canopy cover locations showed no change in carotenoid levels. Concentrations of total chlorophylls (a+b) decreased ( from an average of 327.96 to 256.79 g/g) from initial measurement to day 14 (Fig. 3). Chlorophyll a:b ratios increased ( from an average of 0.96 to 1.16 g/g ) from ini tial measurement to day 14 (Fig. 4). Like the changes in carotenoids, however, plants at two of the five locations showed no significant change. Plants at 0, 75, and FIG URE 1: Change in average anthocyanin concentration (g/g) from initial to day 14 measurements for all plants. One way ANOVA: F 1,108 = 61.89, p < 0.0001 FIGURE 3: Change in total chlorophyll concentration (g/g) from initial to day 14 for all plants. One way ANOVA: F 1,218 = 43.27, p < 0.0001 FIGURE 4: Change in chlorophyll a:b ratio from initial to d ay 14 for all plants One way ANOVA: F 1,108 = 30.49, p < 0.0001 FIGURE 2: Change in average carotenoid concentration (g/g) from initial to day 14 measurements for all plants. One way ANOVA: F 1,108 = 16.54, p < 0.0001

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5 100% canopy cover showed an increase in chlorophyll a:b ratio, while plants at 25 and 50% canopy cover showed no change. DISCUSSION Pigment concentrations were not affected by percent canopy cover. However, the overall pigment decreases indicate that some other factor influenced pigment concentrations at all treatments. I believe that the in fluencing factor is some environmental factor that differs between the experiment site (Biological Station) and the collection site (San Luis). Low wind driven precipitation and cloud immersion during transitional seasons in San Luis mean that the collecti on site was likely drier with less cloud cover than the experimental site during the study (Clark et al. 2000). The bromeliads may have been affected more by the change to a wetter, cloudier, and cooler habitat than by amount of canopy cover at each treatm ent (Polle 1992). Anthocyanins Anthocyanin increase is well known as a protective response to increases in UV radiation. Therefore, an anthocyanin decrease may indicate a decrease in UV radiation (Sullivan 1998). However, plants may also change anthocya nin concentrations in response to stress from other environmental factors, such as temperature extremes, desiccation, or nutrient availability (Dixon et al. 2001; Mori et al. 2005). Dixon et al. found that anthocyanin increases in Impatiens seemed to resul t from stressful conditions (2001). Mori et al. found that higher night temperatures decrease anthocyanin concentrations (2005). These studies suggest that the decrease in anthocyanins found across treatments could be due to lower UV radiation, less stress ful conditions, higher night temperatures, or other factors such as temperature, water, or nutrient availability. Although many factors could have influenced anthocyanins, I believe that overall lower light levels at the experiment locations caused the rec orded anthocyanin decreases. Future experiments should test this hypothesis in relation to the possible increase in cloud cover that the bromeliads experienced when relocated from sunny San Luis to the cloud forest. Carotenoids Like anthocyanins, caroten oid increase has been shown to be a response to excess light. Therefore, the overall decrease in carotenoids found may indicate that all bromeliads experienced a change to a location with less light (McKinnon and Mitchell 2003). This conclusion is in agree ment with that found by the overall decrease in anthocyanins. There is some speculation that low concentrations of carotenoids in shade plants may function as accessory pigments to maximize photosynthesis (Wallentine 2006, Taiz and Zieger 1991). More resea rch should be done to explore this biological role of carotenoids. Total Chlorophyll Nutrient deficiency or temperature change could have decreased total chlorophyll concentrations (Hagen, et al. 1993; Polle 1992). Hagen et al. showed that both nutrient limitation and strong light conditions can cause chlorophyll degradation (1993). Polle showed that overall chlorophyll content of spruce needles decreased with increasing altitude. Lower temperatures in combination with higher light levels at higher altit udes led to photo oxidative stress which reduced chlorophyll concentrations (1992). Though it seems (as shown by decreases in anthocyanins and carotenoids) that higher light levels did not occur at the experiment locations, nutrient limitation and/or lower temperatures could have initiated overall chlorophyll degradation. The

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6 experimental location (Biological Station) is 500 m higher in elevation than the collection location (San Luis), so it seems likely that lower temperatures at the higher altitude may h ave caused chlorophyll degradation (Clark 2000). Further research is needed to determine if removal from their natural habitat could have caused nutrient deficiencies in the Werauhia that decreased chlorophyll concentrations. Chlorophyll a:b ratio Decre ases in chlorophyll a:b ratios are a well known response to shade (Fetene 1990; Lichtenthaler 2001; Thomas 1997). Therefore, one might expect that an increase in chlorophyll a:b ratio would occur in high light. This would indicate that overall increase in a:b ratio found was induced by higher than normal light levels. However, this conclusion contradicts the one deduced from anthocyanin and carotenoid decreases. More studies must be done to determine the range of environmental factors that could affect the chlorophyll a:b ratio in a plant. Another explanation for the contradictory results could be that canopy cover is not an adequate measurement of light irradiance on an individual plant level. Effects of sun flecks or exact position could be more influent ial to a plant than canopy cover (Chazdon 1996). Further research should determine if canopy cover is an accurate estimate of light intensity for individual plants. However, since overall pigment changes did occur, it seems that drastic environmental alte rations do significantly affect Werauhia If changes in light intensity did not affect pigments in the Werauhia studied, some other environmental factor must be responsible. Perhaps rainfall, temperature, and nutrient availability are more influential to b romeliads than previously thought. Though these speculations hint at adaptive abilities of bromeliads to habitat alterations, more research must be done in order to determine how bromeliads will fare in the light of continued habitat destruction. ACKNOW LEDGMENTS Thanks to Anjali Kumar for all her invaluable help, from collecting bromeliads to analyzing data. Thanks to Alan Masters and Pablo Allen for their brainstorming help and support along the way, from teaching me how to use equipment to helping pla n my experiment design. Thanks to Moncho and Yimen for always being supportive and opening the closet countless times to provide equipment. Thanks to Willow Zuchowski and Bill Haber for identifying my Werauhia LITERATURE CITED Anderson, R. C., O. L. Lo ucks and A. M. Swain. 1969. Herbaceous response to canopy cover, light intensity, and throughfall precipitation in conifereous forests. Ecology 50(2): 255 263. Chazdon, R., R. Pearcy, D. Lee, and N. Fetcher. 1996. Photosynthetic responses of tropical fores t plants to contrasting light environments. In S. Mulkey, R. Chazdon, A. Smith. 1996. Tropical Forest Plant Ecophysiology New York: Chapman & Hall. pp. 15 44. Clark, K., R. Lawton, and P. Butler. 2000. The Physical Environment. In Nadkarni, N. and N. Whe elwrite. 2000. Monteverde Oxford University Press: New York. Debinski, D. and R. Holt. 2000. A survey and overview of habitat fragmentation experiments. Conservation Biology 14: 342 355. Dixon, P., C. Weinig, and J. Schmitt. 2001. Susceptibility to UV da mage in Impatiens capensis (Balsaminaceae): Testing for opportunity costs to shade avoidance and population differentiation. American Journal of Botany 88(8): 1401 1408.

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7 Fetene, M., H. S. J. Lee, and U. Luttge. 1990. Photosynthetic acclimation in a terrest rial CAM bromeliad, Bromelia humilis Jacq. New Phytologist 114(3): 399 406. Hagen, C., et al. 1993. Functional aspects of secondary carotenoids in Haematococcus lacustris (Girod) Rostafinski (Volvocales). I. The accumulation period as an active metaboli pr ocess. New Phytologoist 125(3): 625 633. Hegglin, M. and T. Shepherd. 2009. Large climate induced changes in ultraviolet index and stratosphere to troposphere ozone flux. Nature Geoscience. Hopkins, W. 1995. Bioenergetics and the light dependent reactions of photosynthesis. Introduction to Plant Physiology New York: John Wiley and Sons. pp. 163 183 Lichtenthaler, H. and C. Buschmann. 2001Chlorophylls and carotenoids; measurement and characterization by UV VIS Spectroscopy. Current Protocols in Food Analyt ical Chemistry F4(3): 1 8. Luther, H. 2000. Bromeliads. In Nadkarni, N. and N. Wheelwrite. 2000. Monteverde Oxford University Press: New York. pp. 73 74 McKinnon, L. M., and A. K. Mitchell. 2003. Photoprotection, not increased growth, characterizes the re sponse of Engelmann Spruce (Picea engelmannii) seedlings to high light, even when resources are plentiful. New Phytologist 160(1): 69 79. Mori, K., S. Sugaya, and H. Gemma. 2005. Decreased anthocyanin biosynthesis in grape berries grown under elevated nigh t temperature condition. Scientia Horticulturae 105: 319 330. Opdam, P. and D Wascher. 2003. Climate change meets habitat fragmentation: linking landscape and biogeographical scale levels in research and conservation. Biological Conservation 117: 285 297 P olle, A., et al. 1992. Field studies on Norway Spruce trees at high altitudes. I. Mineral, pigment and soluble protein contents of needles as affected by climate and pollution. New Phytologist 121(1): 89 99. Rosevear, M. J., A. J. Young, and G. N. Johnson. 2001. Growth conditions are more important than species origin in determining leaf pigment content of British plant species. Functional Ecology 15(4): 474 480. Sims, D. and J. Gamon. 2002. Relationships between leaf pigment content and spectral reflectanc e across a wide range of species, leaf structures and developmental stages. Remote Sensing of Environment 81: 337 354. Skole, D. and C. Tucker. 1993. Tropical deforestation and habitat fragmentation in the Amazon: Satellite data from 1978 to 1988. Science 260: 1905 1910. Sullivan, J. 1998. Anthocyanin. For The Carnivorous Plant Newsletter. < http://bromeliads. home.att.net/Anthocyanin.htm> Taiz, L. and E. Zieger. 1991. Plant physiology The Benjamin/Cummings Publishing Company, Inc. Redwood City, Californ ia. pp. 335 336. Thomas, H. 1997. Tansley Review No. 92 Chlorophyll: A symptom and a regulator of plastid development. New Phytologist 136(2): 163 181. Ulrich, L. 2008. Physiological Ecology of Tropical Plants Springer, Berlin. pp 107 125. Wallentine, B. 2006. Tropical cloud forest canopy and subcanopy adapt to different light environments by regulating photosynthetic pigments. Tropical Ecology and Conservation Fall 2006. CIEE Monteverde, Costa Rica. pp. 31 42.