Journal of Environmental Science and Management 60-67 (Special Issue 1-2012) ISSN 0119-1144 60 Effects of Light Intensities on Growth Performance, Biomass Allocation and Chlorophyll Content of Five Tropical Deciduous Seedlings in Lao PDR Chanhsamone Phonguodume1, Don Koo Lee2, Silavanh Sawathvong3, Yeong Dae Park4*, Wai Mun Ho5 and Edwin A. Combalicer6 ABSTRACT Growth performance, biomass allocation and chlorophyll content of Afzelia xylocarpa Craib, Anisoptera costata Korthals, Dalbergia cochinchinensis Pierre, Dipterocarpus alatus Roxb.& G.Don and Hopea odorata Roxb. seedlings were compared under different light intensities (100%, 50-70% and 30-50%) in Lao PDR. Results showed that all species had survival rate above 94% under all light intensity treatments. Root collar diameter growth (RCD) varied significantly among the five species (F=10.91, P=0.0018). Afzelia xylocarpa had significantly higher RCD than other species while A. costata and D. cochinchinensis showed higher RCD under higher light intensity. Height was also affected significantly by light intensities among the species tested (F=4.85, P=0.0223). A. xylocarpa showed better height growth at lower light intensity. The number of leaves showed significant differences among species (F=9.191, P=0.004) but was not significantly different among light intensities (p<0.05). Biomass allocation was not significantly different within and among species (F=3.39, P=0.0665) under different light intensities (F=2.72, P=0.096). Both A. xylocarpa and D. alatus accumulated higher total biomass under 50-70% light intensity compared to other light conditions. Hopea odorata showed increasing biomass with increasing light intensity but A. costata accumulated more biomass at lower light intensity. The experiment also found that total chlorophyll content was significantly different among species (p<0.05). This study showed that each species had different adaptations to light conditions that should be considered in nursery practice for optimum growth of seedlings. Key words: biomass, chlorophyll content, growth, light intensity, tropical deciduous seedlings INTRODUCTION Tropical rainforests could be influenced by the surrounding vegetation and habitat environment, such as light and temperature, in their ecological and physiological responses (Henmann and Hugh 2010; Wright 2005). In a tropical deciduous forest, more light exists as it has a thinner canopy which produces more undergrowth. There are also seasonal changes as the light and temperature shift just enough to trigger the trees to drop their leaves (Johnson 2009). However, light environment in degraded secondary forests may be diverse because light condition changes by forest structure such as gap formation and time span after disturbance (Mesquita 2000; Whitmore 1998). Light is the main factor covered in the studies of environmental responses of dipterocarp and other tropical tree species (Norisada and Kojima 2005). Light plays an important role in the development of a plant. Processes such as photosynthesis and phototropism depend on the availability of light sources for plants (Long 2011). Plant seeks out light sources for energy and survival. Light is one of the most important environmental factors affecting plant establishment and growth in a tropical deciduous forest. Plant performance is enhanced through morphological and physiological acclimation to light environment (Kozlowski and Pallardy 1996). It is also an important factor in forest regeneration and growth. The tropical species are unique in vegetation and habitat as they adapt to varying light intensities. The role of light in the life of a plant is much more complex than the plant sitting in the sun for a whole day. Shade environment or light intensity limits the photosynthetic capacity and biomass of a plant. Growth analyses of tree seedlings under controlled conditions indicate how plants adjust to light environment (Poorter 2001; Popma and Bongers 1988). Leaves under shade conditions have higher chlorophyll content and lower leaf mass per area (LMA) so as to maintain the low light compensation point (Lambers, Chapin and Pons 1998). Plants absorb light energy and transfer it into the photosynthetic apparatus (Sims and Gamon 2002). Healthy plants capable of maximum growth are generally expected to have larger amounts of chlorophyll than unhealthy ones. A better understanding of ecological traits such as Ph.D. Candidate, Department of Forest Sciences, Research Institute for Agriculture and Life Sciences, College of Agriculture and Life Sciences, Seoul National University and Director, Forest Research Center, National Agriculture and Forestry Research Institute, Ministry of Agriculture and Forestry, Vientiane Capital, Lao PDR P.O. Box: 7174 2 Professor, Department of Forest Sciences, Research Institute for Agriculture and Life Sciences, College of Agriculture and Life Sciences, Seoul National University 3 Director General, Department of Forestry, Ministry of Agriculture and Forestry, Vientiane Capital, Lao PDR P.O. Box: 7174 4 Research Professor, Department of Forest Sciences, Research Institute for Agriculture and Life Sciences, College of Agriculture and Life Sciences, Seoul National University, 599 Gwanangno, Gwanak-gu, Seoul, 151-921, Republic of Korea. E-mail: [email protected] (*corresponding author) 5 Researcher, Forest Research Institute Malaysia, 52109 Kepong, Selangor Malaysia 6 Assistant Professor, College of Forestry and Natural Resources, University of the Philippines Los Banos, 4031 College, Laguna, Philippines 1 Journal of Environmental Science and Management (Special Issue 1-2012) 61 strong light tolerance for the target species could improve techniques for enrichment planting in secondary forests (Appanah and Weinland 1993; Kenzo et al. 2008; Krishnapillay 2002). It is therefore necessary to consider specific species favorable to be planted in light conditions when conducting effective enrichment planting in degraded secondary forests. This will be very useful in managing mixed plantation forests and in rehabilitating degraded areas of the country. However, there are limited information about the effect of light intensities on growth and biomass allocation of tree species. Thus, the objective of this study was to determine the effects of light intensities on growth, biomass allocation and chlorophyll content of selected native species in tropical deciduous forest of Lao PDR. Biomass (roots, stem, and leaves) was determined in one-year-old seedlings of the five tropical deciduous species. Thirty healthy (30) seedlings were randomly selected for each species in every treatment for determination of fresh weight and total dry biomass. Samples were placed in paper bags for transferring to the laboratory. The root, stem and leaf samples were oven dried at 80oC for 48 to 72 hours until constant weight. MATERIALS AND METHODS Chlorophyll content Site description The chlorophyll content (a, b and total) was obtained by collecting five leaf samples in each species for every treatment. A single-hole puncher was used to get the samples. The samples were then put into vial with 10 mL 80% acetone to extract the chlorophyll. The extracts were kept in the dark at 4oC for seven days. The absorbance of extracts was read at wavelengths of 663 nm and 645 nm using Hitachi U-1000 Spectrophotometer (Shimadzu Co., Japan) in the Department of Chemistry, Faculty of Sciences, National University of Lao PDR. Arnon’s equation (Arnon 1949) was used to calculate the chlorophyll content (a, b, and total) as shown below. Unlike other methods which use solvents like Dimethyl Formamide (DMF) and Dimethyl Sulfoxide (DMSO), the use of acetone in Arnon’s method is not toxic to human as it is not easily absorbed by the skin. (1) Chlorophyll a ( g/mL-1) = 12.7 A663 – 2.59 A645 (2) Chlorophyll b ( g/mL-1) = 22.9 A645 – 4.67 A663 (3) Total chlorophyll ( g/mL-1) = 8.05 A663 + 20.29 A645, where: A 663 is the absorbance at 663 nm; and A645 is the absorbance at 645 nm. The study was conducted in the nursery of Forestry Research Centre, National Agriculture and Forestry Research Institute, Lao` PDR. It is located between 18o 15’ 32’’ N longitude and 102o 33’ 24’’ E latitude with an altitude of 195 m above sea level. The average annual precipitation is 1,990 mm from 2000 to 2008 and was moderately seasonal in distribution. The average air temperature is 27oC with the lowest in December (23.5oC) and highest in May (30.5oC). Relative humidity is lowest in February (69%) and highest in August (84%). Study species and planting materials The seeds of five tropical deciduous species, namely Afzelia xylocarpa, Anisoptera costata, Dalbergia cochinchinensis, Dipterocarpus alatus, and Hopea odorata were collected from identified natural seed sources in Lao PDR. Planting materials used were plastic pots (12 cm in diameter x 18 cm in height) and mixture of planting medium with a ratio of 2:1:2:2 for soil, sand, rice husk (biochar) and manure, respectively. Seedlings were transplanted from seed germination bed to plastic bag containers two weeks after germination. Seeds were germinated based on the recommended germination procedure for each species. Seedlings were nursed for another two weeks by removing weeds and constant application of water to ensure seedlings survival before treatment application in May 2007. A Completely Randomized Design (CRD) was employed for this study with five species and three treatments of light intensity at 100% (control), 50-70% and 30-50%. A total of 1,500 (100 seedlings per treatment for each species) were used for the study. Growth performance The growth performance of seedlings was assessed using survival rate (%), root collar diameter (RCD, mm), total height (cm) and number of leaves. These parameters were measured every three months from June 2007 up to June 2008. Biomass Data analysis The mean values and standard errors of growth parameters, biomass, and chlorophyll content were analyzed using MS Excel 2007 and SAS 9.1.3 for Windows 2007 (SAS Institute Inc., USA). Duncan’s Multiple Range Test (DMRT) was used for multiple comparisons of seedling performance under different light intensities. The significance for all analysis was determined at p<0.05. RESULTS AND DISCUSSION Tree species used in the study Two legumes (Afzelia xylocarpa Craib and Dalbergia cochinchinensis Pierre.) and three dipterocarp (Anisoptera cossta Korth., Dipterocarpus alatus Roxb., and Hopea Effects of light intensities 62 odorata Roxb.) were used for the study. The selected species have wide ranges of ecological adaptations. For instance, most dipterocarp species are shade tolerant while the legumes are light demanding. The species in this experiment are also highly known for their high quality timber and non-timber products (Table 1). Growth performance Survival rate Most of the species showed high survival rates of over 94% in all species and light treatments. Survival of all the one-year-old seedlings within and among the species was not affected by different light intensity treatments (Table 2). This indicated that both legumes species (A. xylocarpa and D. cochinchinensis) and dipterocarp species (Dipterocarpus alatus, H. odorata and A. costata) performed similarly (p>0.05) in terms of survival under the given light conditions. In a case study at Barro Colorado Island of Panama, however, different species (e.g. pioneer, shade tolerant, and intolerant, evergreen or deciduous tree species) showed different responses to light or gap dynamics (Kitajima 1994). In contrast, this study showed that there were no significant differences in seedlings survival grown under different light intensities. Figure 1 shows that different light intensities resulted in statistically different RCD among the species (F=10.91, P=0.0018). Means of RCD showed significantly higher value (p<0.05) in A. xylocarpa compared to other species probably due to its ability, as a leguminous species, in nitrogenfixation that promoted better growth. This is followed by A. costata and D. alatus. Dalbergia cochinchinensis recorded the smallest mean with lower RCD at reduced light intensity. The root collar diameter (RCD) of A. xylocarpa was significantly higher than the other four species at all light treatments. This indicates its suitability for planting in open or under shade. The RCD of A. costata and D. cochinchinensis were large under higher light intensity and decreased with decrease in light intensity level. On the other hand, the RCD for H. odorata and D. alatus were significantly higher (p<0.05) under the moderate light condition (5070%). Among the five species studied, A. xylocarpa had significantly thicker RCD (9.7-10.5 mm) while H. odorata and D. cochinchinensis had thinner RCD (5.7-6.4 mm). The maximum growth of tropical rainforest seedlings in Africa was reported at 10-44% sunlight (Agyeman, Swaine, and Thomption 1999) while another study showed 50-100% sunlight for fast growing species and 25-50% shade for slow growing species in Bolivia (Poorter 1999). Height Height was affected significantly by different light intensities among the species tested (F=4.85, P=0.0223). A. xylocarpa showed significantly (p<0.05) higher growth at 79.40 cm and 78.20 cm in 30-50% and 50-70% light intensities, respectively. However, H. odorata and A. costata had taller height under higher light intensity and are therefore suitable to be planted in the open. D. cochinchinensis had almost similar height at all light intensity levels. On the other hand D. alatus had taller height (39 cm) under medium light intensity (Figure 2). In a study by Kenzo et al. (2008) and Sack and Grubb (2002), they found that light had effects on wood density and height of seedlings. They observed that 35% of light condition was suitable for seedling growth. Mesquita (2000) conducted a research to compare five light intensities (100%, 89-90%, 50-60%, 2030% and 0% control). It was reported that in natural forests, seedlings height grew to more than 25 cm under 50% light intensity and one-fourth of the species grew to more than 25 cm year-1. Number of leaves The result also agrees with Agyeman, Swaine and Thomption (1999) who showed that the number of leaves showed strong relationships with species characteristics and light. Table 3 shows the mean numbers of the leaves of the five species grown under different light intensities. Dalbergia cochinchinensis and A. xylocarpa showed significantly higher number of leaves under all light intensities. Further, D. cochinchinensis and A. xylocarpa had highest number of leaves, 35 and 23, respectively, when grown at high light intensity levels. Biomass allocation Light was reported to affect biomass allocation (Poorter 1999). Most of the studies had focused on growth rather than biomass (Fetcher, Strain and Oberbauer 1983; King 1991; Oberbauer and Strain 1985; Osunkoya et al. 1993; Popma and Bongers 1991; Rincon and Huante 1993). AKECOP (2010a) reported that total biomass (roots, stems and leaves) of two-year-old Pterocarpus indicus were significantly different among light intensities with 120 g in 100%, 63 g in 65% and 59 g in 30%, respectively. In addition, biomass allocation was found to be in order of leaves > stems > roots. In this study, biomass allocation was not statistically different at 5% level within and among the species (F=3.39, P=0.0665) and among under different light intensities with (F=2.72, P=0.096). Under 100% light intensity, Dipterocarpus alatus had the highest accumulation of total biomass (p<0.05) followed by H. odorata (Table 4). These two species have thus shown their possibility to grow under direct sunlight, for instance, in the open areas of degraded sites for rehabilitation purposes. Nonetheless, D. cochinchinensis and A. costata Journal of Environmental Science and Management (Special Issue 1-2012) 63 Table 1. Ecological distribution, growth habit and economic importance of the species used in the study. Family Leguminosae Dipterocarpaceae Leguminosae Species Afzelia xylocarpa Ecological distribution Native to Cambodia, Lao PDR, Myanmar, Thailand and Vietnam; occurs in evergreen and deciduous forests at 100-650 m altitudes Native to Cambodia, Lao PDR, Malaysia, Anisoptera Thailand and Vietnam; occurs in evergreen, costata moist or slightly dry forests, sometimes gregariously growing in a pure stand at altitudes up to 1000 m Occurs in dry evergreen forests of Dalbergia cochinchinensis Cambodia, Lao PDR, Thailand and Vietnam at altitudes up to 1000 m Dipterocarpaceae Dipterocarpus alatus Dipterocarpaceae Hopea odorata Growth habit Tree up to 30 m tall and 150 cm in diameter; lightdemanding; deciduous; endangered species Large tree up to 30-40 m and 50-100 cm in diameter; endangered species Deciduous tree up to 30 m tall, 60-80 cm in diameter; drought-tolerant but slow growing; shade-tolerant when young; vulnerable species Native to evergreen and deciduous forests Large tree up to 40 m tall of Cambodia, Lao PDR, Myanmar, and 150 cm in diameter; Philippines, Thailand, Vietnam, Bangladesh, shade-tolerant and fast Andaman islands; occurs gregariously along growing; endangered rivers at altitudes up to 500 m species Distributed in Cambodia, India, Lao PDR, Large tree up to 30-40 m Malaysia, Thailand, Vietnam; occurs and 60-80 cm in diameter; it gregariously in dense tropical evergreen demands wet and deep soil; forests, but after a long period of selective natural regeneration is good logging, it is only found in small groups or under thin forest-cover, in solitude at altitudes up to 900 m endangered species Uses Timber, food, tanning, soil improvement Timber, resin Timber, soil improvement Timber, oleoresins, medicine, soil improvement Timber, resins and gums Sources: Dung 1996; Lao Tree Seed Project 2003. Figure 1. Root collar diameter of Afzelia xylocarpa, Anisoptera costata, Dipterorpus alatus, Hopea odorata and Dalbergia cochinchinensis under different light intensities (the same letters are not significantly different by using DMRT p<0.05). Effects of light intensities 64 Figure 2. Height of Afzelia xylocarpa, Hopea odorata, Anisoptera costata, Dalbergia cochinchinensis and Dipterocarpus alatus under different light intensities (the same letter are not significant difference among height using DMRT p<0.05). Table 2. Survival rate of seedling under different light intensities (% of survival). Species Afzelia xylocarpa Anisoptera costata Dalbergia cochinchinensis Dipterocarpus alatus Hopea odorata Overall mean of treatments Light treatment 100% 50-70% 30-50% 96 100 100 96 100 100 98 100 99 Overall mean of species 96.7 100.0 99.7 94 96 97.2 96 97 97.8 94 99 98.0 94.7 97.3 97.7 Table 3. Number of leaves in one-year-old tropical deciduous seedlings in the nursery. Species Afzelia xylocarpa Anisoptera costata Dalbergia cochinchinensis Dipterocarpus alatus Hopea odorata 100% 23 (5)a 12 (3)bc 35 (6)a 7 (1)c 20 (7)b Light treatment 50-70% 30-50% 27 (9)a 25 (9)a 12 (2)bc 12 (2)bc 24 (2)a 26 (4)a 9 (2)c 9 (1)c 17 (4)b 15 (4)b Notes: Value in parenthesis indicates standard errors; Different letters in columns indicate significant difference among number of leaf using DMRT (p<0.05). performed poorly in the strongest light intensity treatment. Afzelia xylocarpa showed significantly higher (p<0.05) total biomass under 50-70% light intensity among the five species. Under this light treatment, A. xylocarpa as well as D. alatus accumulated the highest biomass compared to other light condition. Dalbergia cochinchinensis and A. costata recorded the lowest biomass again under 50-70% light intensity. Under the lowest light intensity treatment (30-50%) A. xylocarpa accumulated significantly higher (p<0.05) total biomass followed by D. alatus. This indicated that both species preferred low to moderate light intensity for optimum biomass production. Hopea odorata had the lowest total biomass in 30-50% light intensity but showed and increasing trend with higher light intensity. Under 100% light intensity, H. odorata only showed higher root biomass, stem and leaf. Dalbergia cochinchinensis in 30-50% light intensity, Dipterocarpus alatus and A. xylocarpa showed higher root, stems and leaves. While A. costata showed different root biomass at 30-50%, stem at 100% and leaf at 50-70% light intensities. However, species characteristic are optimum growth and biomass allocation with different light intensities. In a study by Montgomery and Chazdon (2002), 14month-old seedlings of three tropical species showed a strong Journal of Environmental Science and Management (Special Issue 1-2012) 65 Table 4. Biomass allocation of one-year-old Hopea odorata (Hop), Dalbergia cochinchinensis (Dal), Dipterocarpus alatus (Dip), Anisoptera costata (Ani) and Afzelia xylocarpa (Afz) grow in nursery under different light intensities. Light treatment 100% 50-70% 30-50% Component Root Stem Leaf Total Root Stem Leaf Total Root Stem Leaf Total Ani 5.70 (1.46)c 5.60 (1.91)c 5.23 (0.81)b 16.54 (1.39)d 6.13 (0.83)c 5.51 (0.66)d 8.13 (0.85)b 19.77 (0.78) 7.01 (1.26)b 3.95 (2.84)d 6.89 (0.61)b 17.85 (1.57)c Afz 6.72 (1.40)a 10.93 (2.16)b 3.18 (1.15)c 20.82 (1.57)c 15.99 (0.34)a 38.55 (0.61)a 6.27 (0.49)c 61.32 (0.48)a 12.16 (1.50)a 23.22 (1.86)a 4.98 (1.04)bc 40.36 (1.46)a Dal 6.05 (0.76)c 6.34 (1.07)c 4.37 (0.45)b 16.76 (0.76)d 5.31 (0.61)c 6.17 (0.90)d 2.54 (0.33)d 14.02 (0.61) 6.28 (0.43)bc 7.69 (1.67)c 5.62 (0.81)b 19.59 (0.91)c Dip 10.16 (0.60)b 15.35 (0.75)a 9.78 (0.43)a 35.29 (0.60)a 11.44 (1.88)b 22.48 (2.80)b 16.26 (1.20)a 50.18 (1.96)b 7.51 (0.51)b 10.23 (0.77)b 9.95 (0.82)a 27.67 (0.70)b Hop 9.29 (2.27)c 12.01 (5.88)b 8.08 (1.04)ab 29.38 (3.06)b 4.29 (0.49)c 10.19 (0.68)c 7.55 (1.27)c 22.03 (0.81)c 3.37 (1.13)c 5.26 (1.05)cd 4.59 (0.62)bc 13.22 (0.93)d Notes: Value in parenthesis indicates standard errors; Different letters in rows indicate significant difference among number of leaf using DMRT (p<0.05). increasing in total biomass as light increased. However, the strength of the growth response differed among species causing a change in the rank order of species growth rates as light availability increase. Poorter (2001) reported that 17% of six tropical rain forest saplings had negative relative biomass growth rates, although they occurred in fairly bright conditions. Chlorophyll content Table 5 shows the mean total chlorophyll content of one-year-old tropical deciduous seedlings in different leaf positions under 100% light intensity. In this study, total chlorophyll content was significantly different (p<0.05) among species. Dipterocarpus alatus showed highest chlorophyll contents in all respective leaf positions among the five species studied. The highest chlorophyll content in this species was found in the middle leaf position. Meanwhile, H. odorata and D. cochinchinensis had significantly higher total chlorophyll contents in the lower position. In A. xylocarpa the total chlorophyll content was highest in the middle leaf position (1.06 g mL-1). The total chlorophyll content is more depedent on physiological characteristics of species than on light or shade condition (Minotta and Pinzauti 1996). Cho et al. (2008) reported that chlorophyll contents of five hardwood species in central temperate zone of Korea were highest in 7-12% light intensity, while lowest under 100% light intensity. Sun leaves (upper leaf) grown under high irradiance possess higher photosynthetic rates than shade leaves (Boardman 1977; Lichtenthaler and Akoyunoglou 1981) but most of the leaves in shading conditions have a high leaf area index. AKECOP (2010b) reported that chlorophyll contents of different leaf positions were significantly different in three one-year-old tropical species in the Philippines with highest in lower leaf position of Acacia auriculiformis, (11.5 g/mL-1) and A. mangium (12.2 g/mL-1) and, in middle position of Pterocarpus indicus (18.3 g/mL-1). Lee (2006) reported that chlorophyll contents of four five-year-old species planted in the Philippines were significantly different with different spacing (2 x 2 m, 4 x 4 m, 6 x 6 m) The results showed highest chlorophyll contents in Erythrina orientalis (5.5 g/mL-1) and P. indicus (3.6 g/ mL-1) for 4 x 4 m, in Bischofia javanica (2.3 g/mL-1) for 6 x 6 m, and in Dracontamelon dao (2.2 g/mL-1) for 2 x 2 m. Table 5.Total chlorophyll content of five species after oneyear of planting in nursery. Species Anisoptera costata Afzelia xylocarpa Dalbergia cochinchinensis Dipterocarpus alatus Hopea odorata Upper 2.678 (0.84)a 0.994 (0.02)ab 1.086 (0.08)b 2.824 (0.78)b 1.067 (0.13)c Leaf positions Middle 1.738 (0.91)b 1.055 (0.07)a 1.044 (0.09)b 3.801 (0.06)a 1.249 (0.01)b Lower 1.090 (0.07)c 0.692 (0.12)b 1.976 (1.15)a 2.842 (0.95)b 2.827 (0.90)a Notes: Value in parenthesis indicates standard errors; Different letters in rows indicate significant difference using DMRT (p<0.05). Table 6 summarizes the parameters used in the five study species under different light intensities. Among the light intensity treatments, most of the species performed better in 50-70% light intensity such as H. odorata, D. alatus and A. xylocarpa. Only D. cochinchinensis had good performance in 30-50% light intensity. This means that 50-70% light intensity is the optimum amount needed by the plant species to survive and to obtain good performance in the field. The presence of optimum light can have a direct bearing on the plant’s rate of growth. A plant’s most natural habitat provides the intensity of light needed for optimal growth. As a result, different plant types may require different light intensities. Effects of light intensities 66 Table 6. Summary of parameters used in the five species by score. Species SR RCD 3 3 3 2 3 3 3 2 3 1 Ani Afz Dal Dip Hop Legend: SR RCD H Survival rate Root collar diameter Total height L W Ch 100% 50-70% 30-50% H L W Ch Total SR RCD H L W Ch Total SR RCD H L W Ch Total 3 3 1 3 16 3 2 1 3 3 2 14 3 2 2 3 2 1 13 1 1 1 2 10 3 3 2 3 3 3 17 3 3 3 2 2 1 14 2 3 2 2 15 3 2 2 1 1 1 10 3 1 3 2 3 3 15 2 2 2 1 12 3 3 3 3 3 3 18 3 1 2 3 1 2 12 2 3 3 1 13 3 3 3 2 2 2 15 3 2 1 1 1 3 11 Number of leaves Biomass Chlorophyll content 1 2 3 Detailed physiological effects of varying light intensities were not determined in this study except for chlorophyll contents. However, it can be inferred that light intensity contributes greatly to plant physiological processes since plant growth relies on a series of interactions that involve the presence of light. For example, photosynthesis enables plant metabolism processes to take place and provides the energy that fuels these processes. Light intensity levels can have a significant effect on photosynthesis rates, which are directly related to a plant’s ability to grow. According to Jeanty (2008), light intensity has something to do with the amount of light energy made available to a plant, which can vary according to color and the actual strength of the light. In addition, plants are typically most responsive to light that falls within the blue and red light ranges. In effect, higher light intensities make more energy available for plant photosynthesis processes to take place. CONCLUSION Light is an important factor in forest regeneration and growth. Tropical deciduous tree species are unique in vegetation and habitat and therefore light affects the shade-tolerant species physiologically and biochemically. Understanding light requirement is important to plant physiology with regard to stress. In conclusion, the survival was not significantly different within and among the species in different light intensities, while the number of leaves, root collar diameter, height and biomass allocation of seedlings vary with light and species. The chlorophyll content of each species in this study was also dependent on different leaf positions (upper, middle and lower positions). Light intensity between 50-100% was suitable for the growth of one-year-old tropical deciduous in the nursery. The results of this study can be applied in the nursery practice to produce healthy seedlings for successful rehabilitation of degraded tropical forests in Lao PDR. Low Medium High the derivation of light response index.” Journal of Ecology 87: 815-827. AKECOP. 2010a. “Photosynthetic response to different light intensities on Narra (Pterocarpus indicus) in La Mesa Watershed, Philippines.” In: Restoration of Degraded Terrestrial and Mangrove Forest Ecosystems in ASEAN Region (Ed. D.K. Lee). Seoul National University, Republic of Korea. pp. 109-118. AKECOP. 2010b. “Physiological characteristics and nutrient productivity of Acacia auriculiformis, Acacia mangium, in Mt. Makiling and Pterocarpus indicus in La Mesa Watershed, Philippines.” In: Restoration of Degraded Terrestrial and Mangrove Forest Ecosystems in ASEAN Region (Ed. D.K. Lee). Seoul National University, Republic of Korea. pp. 95-108. Appanah, S. and Weinland, G. 1993. Planting Quality Timber Trees in Peninsular Malaysia. a Review. Malayan Forest Records No. 38. FRIM, Kuala Lumpur, Malaysia. 247 pp. Arnon, D.I. 1949. “Copper enzymes in isolated chloroplasts, polyphenol-oxidase in Betula vulgaris.” Plant Physiology 24:1-15. Boardman, N.K. 1977. “Comparative photosynthesis of sun and shade plants.” Annual Review Plant Physiology 28: 355-377. Cho, M.S., K.W. Kwon, G.N. Kim, and S.Y. Woo. 2008. “Chlorophyll contents and growth performances of the five deciduous hardwood species growing under different shade treatments.” Korean Journal of Agriculture and Forest Meteorology 10: 149-157 (in Korean with English Abstract). Dung, V.V. 1996. Vietnam Forest Trees. Forest Inventory and Planning Institute of Vietnam. Agricultural Publishing House, Hanoi, Vietnam. REFERENCES Fetcher, N., B.R. Strain, and S.F. Oberbauer. 1983. “Effects of light regime on the growth, leaf morphology, and water relations of seedlings of two species of tropical trees.” Oecologia 58: 314-319. Agyeman, V.K., Swaine, M.D. and Thomption, J. 1999. “Responses of tropical forest tree seedlings to irradiance and Henmann, B. and D.S. Hugh. 2010. “Late-glacial and Holocene vegetation, climate and fire dynamics in the Serra dos Journal of Environmental Science and Management (Special Issue 1-2012) 67 Orgaos, Rio de Janeiro State, southeastern Brazil.” Global Change Biology 16: 1661-1671. partitioning by tropical tree seedlings in the absence of canopy gaps.” Oecologia 131: 165-174. Jeanty, J. 2008. The Effect of Light Intensity on Plant Growth. Access: http://www.ehow.com/about_6671823_effect-lightintensity-plant-growth.html. Norisada, M. and K. Kojima. 2005. “Nitrogen form preference of six Dipterocarp species.” Forest Ecology and Management 216: 175-186. Johnson, J. 2009. What is a Tropical Deciduous Forest? Access: http://www.helium.com/items/1439226-tropical-rainforestdefinition-what-is-a-tropical-rainforest-tropical-deciduousrainforest. Oberbauer, S.F. and B.R. Strain. 1985. “Effects of light regime on the growth and physiology of Pentaclethra macroloba (Mimosaceae) in Costa Rica.” Journal of Tropical Ecology 1: 303-320. Kenzo, T., R. Yoneda, Y. Matsumoto, M.A. Azani, and M.M. Nik. 2008. “Leaf photosynthetic and growth responses on four tropical trees species to different light conditions in degraded tropical secondary forest, Peninsular Malaysia.” Japan Agricultural Research Quarterly 42: 299-306. Osunkoya, O.O., J.E. Ash, A.W. Graham, and M.S. Hopkins. 1993. “Growth of tree seedlings in tropical rain forests of north Queensland, Australia.” Journal of Tropical Ecology 9: 1-18. King, D.A. 1991. “Correlations between biomass allocation, relative growth rate and light environment in tropical forest saplings.” Functional Ecology 5: 485-492. Kitajima, K. 1994. “Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees.” Oecology 98: 419-428. Kozlowski, T.T. and S.G. Pallardy. 1996. Physiology of Woody Plants. 2nd Edition. Academic Press, London, New York, 411 pp. Krishnapillay, B. 2002. A Manual for Forest Plantation Establishment in Malaysia. Malayan Forest Records No. 45. FRIM, Kuala Lumpur, Malaysia. 286 pp. Lambers, H., F.S. Chapin, and T.L. Pons. 1998. Plant Physiological Ecology. Springer-Verlag, New York, 513 pp. Lao Tree Seed Project. 2003. 50 Species Monographs for Plantation in Lao PDR. NAFRI/LTSP/DANIDA. Lee, Y.K. 2006. Ecosystem Structure and Functions in Relation to Restoration of Degraded Tropical Forest in the La Mesa Watershed and Mt. Makiling, Philippines. Ph.D. Dissertation. Seoul National University, Republic of Korea. 165 pp. Lichtenthaler, H. and G. Akoyunoglou. 1981. Photosynthesis IV. Philadelphia Balaban Internet Science Service, p. 273-285. Long, N. 2011.The Importance of Light to a Plant.Access: http://www. ehow.com/facts_7619414_importance-light-plant.html. Poorter, L. 2001. “Light-dependent changes in biomass allocation and their importance for growth of rain forest tree species.” Functional Ecology 15: 113-123. Poorter, L. 1999. “Growth responses of 15 rain-forest tree species to light gradient: the relative importance of morphological and physiological Traits.” Functional Ecology 13: 396-410. Popma, J. and F. Bongers. 1988. “The effect of canopy gaps on growth and morphology of seedlings of rain forest species.” Oecology 75: 625-632. Popma, J. and F. Bongers. 1991. “Acclimation of seedlings of three Mexican tropical rain forest tree species to change in light availability.” Journal of Tropical Ecology 7: 85-97. Rincon, E. and P. Huante. 1993. “Growth responses of tropical deciduous tree seedlings to contrasting light conditions.” Trees 7: 202-207. Sack, L. and P.J. Grubb. 2002. “The combined impacts of deep shade and drought on the growth and biomass allocation of shade-tolerant woody seedlings.” Oecologia 131: 175-185. Sims, D.A. and J.A. Gamon. 2002. “Relationships between leaf pigment content and spectral reflectance across a wide range of species, leaf structures and developmental stages.” Remote Sensing of Environment 81, 337–354. Whitmore, T.C. 1998. An Introduction to Tropical Rain Forests. 2nd Edition. Oxford University Press. Oxford, UK. 282 pp. Wright, S.J. 2005. “Tropical forests in a changing environment.” Trends in Ecology and Evolution 20: 553-560. Mesquita, R.C.G. 2000. “Management of advanced regeneration in secondary forests of the Brazilian Amazon.” Forest Ecology and Management 130: 131-140. ACKNOWLEDGMENT Minotta, G. and S. Pinzauti. 1996. “Effects of light and soil fertility on growth, leaf chlorophyll content and nutrient use efficiency of beech (Fagus sylvatica L.) seedlings.” Forest Ecology and Management 86: 61-71. This study was carried out with the support of "ASEANKorea Environmental Cooperation Project" provided by Korea Ministry of Foreign Affairs and Trade. The authors sincerely thank the staff of Forestry Research Centre of Lao PDR, for their assistance throughout the experiment. Montgomery, R.A. and R.L. Chazdon. 2002. “Light gradient
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