PHYSIOLOGICAL CHARACTERISTICS RELATED TO CARBON SEQUESTRATION OF TREE SPECIES IN HIGHLAND FOREST ECOSYSTEM OF MOUNT HALIMUN-SALAK NATIONAL PARK
Nuril Hidayati1,2 , M. Mansur and Titi Juhaeti Received : 8 March 2010, Accepted : 15 February 2012
ABSTRACT
Biological diversity can have significant contribution to reduce the build-up of greenhouse gases in the atmosphere. The trees in a forest stand form an essential part in the functioning of the terrestrial biosphere, especially in the carbon cycle. Yet tree photosynthesis is far less studied than crop photosynthesis for several reasons: the large number of species; difficulty in measuring photosynthesis of entire trees or of forest stands. This research aims to assess the contribution of biological diversity in carbon sequestration by analyzing the physiological characteristics (photosynthesis, transpiration, stomatal conductance, leaf chlorophyll content) of species native to tropical highland forest ecosystem ofMount Halimun-Salak National Park. The results showed that there was a wide range of variation of
CO2 assimilation rate between tree species. The overall CO2 absorption rate ranged 1.1913 - 31.3875 µmolm-2 s -1 , the highest rate was reached byLithocarpus sp . (pasang parengpeng) (31.3875 µmolm-2 s -1 ), followed byLitsea noronhae (huru lumlum) (21.5750 µmolm-2 s -1 ), Saurauia nudiflora (kilebo) (11.8175 µmolm-2 s -1 ),Vernonia arborea ( hamirung) (6.7125 µmolm-2 s -1 ) and Litsea sp . (huru bodas) (6.2725 µmolm- 2-1 s ). The rate of CO2 assimilation was affected by incident radiation and thus the photon flux (Q leaf).
Correlation between CO2 assimilation and Q leaf under certain environmental condition was considerably high. Incident radiation and Q leaf also affected stomatal conductance and thus rate of transpiration.
Keywords: Biological diversity, photosynthesis, carbon sequestration, greenhouse gases
I. INTRODUCTION greenhouse gases in the atmosphere. Each year about 60 gigatonnes (GT) of carbon (C) are taken Forests represent 21% of the continental area up and released by land-based ecosystems and which are 76% of terrestrial biomass and 37% of about 90 GT are taken up and released by ocean its bioproductivity (Ceulmans and Sauger, 1991). systems. These natural fluxes are large compared A biologically diverse tropical forest holds 50 to the approximately 6.3 GT being emitted from times more carbon per unit of area than a fossil fuels and industrial processes, and about 1.6 monoculture plantation. Thus, the trees in a forest GT per year from deforestation (CBD,2008). stand form an essential part in the functioning of A reliable method for restoration of forest and the terrestrial biosphere, especially in the carbon reforestation is using native tree species. The cycle. Yet tree photosynthesis is far less studied trees which are suitable for CDM (Clean than crop photosynthesis for several reasons: the Development Mechanism) purpose should have large number of species; difficulty in measuring the three following characteristics; (1) seedling photosynthesis of entire trees or of forest stands. that can adapt easily to open sites, after Biological diversity can make a significant transplanting from shade (nurseries) to the sunlit contribution to reducing the build-up of conditions; (2) fast-growing species that is able to compete with weeds and fern (Ashton, 1998); (3) 1Biology Research Center - Indonesia Institute of Science 2 tree species that have high CO assimilation Corresponding author:[email protected] 2
49 Journal of Forestry Research Vol. 9 No. 2, 2012: 49-61 capacity and long live. However, these highland forest ecosystem examined. This physiological characteristics differ widely among research aimed to provide informations on tree tree species. In order to attain successful characteristics related to high carbon reforestation, it is necessary to carefully select sequestration by analyzing their physiological appropriate trees for transplanting based on these characteristics (CO22 absorption, CO assimilation, characteristics. For evaluation of the appropriate stomatal conductance, chlofophyll content), and trees, ecological (dominance, biomass, carbon ecological characteristics (dominance, biomass content) and physiological (photosynthesis, production and carbon sequestration ). transpiration) characteristics are suitable indicators (Takahashiet al ., 2005, 2006; Ashton, 1998). II. MATERIAL AND METHODS
Variance in CO2 assimilation rate is large among trees grown under sunlit conditions not A. Study Site only across the continental transect as well as This study was conducted in two locations across tropical climate regions (Matsumotoet al ., representing humid highland forest in Mount 2003). Fast-growing trees often have relatively Halimun-Salak National Park : 1) Cikaniki and 2) higher CO assimilation rate in tropical climate 2 Citalahab. Two plots were established in two zone suggesting that CO assimilation rate can be 2 different sites. The first plot of 100 m x 100 m was an indicator for evaluating fast-growing located in Cikaniki station of 06o 44'57” S and characteristics (Presset al ., 1996, Matsumoto et al ., o 106 32'08” E, 1,100 m above sea level and the 2003). second one of 50 m x 50 m was located in In this study ecological and physiological oo characteristics of tree species native to humid Citalahab of 06 44' 32.2” S and 116 31' 44.0” E, 1,076 m above sea level (Figure 1).
Figure 1. Cikaniki and Citalahab study sites in Mount Halimun-Salak National Park
50 Physiological Characteristics Related to Carbon Sequestration of Tree Species tn Highland Forest Ecosystem ..... (Nuril Hidayati)
B. Ecological Analysis measurement of carbon dioxide (CO2 ) uptake is a Analysis of vegetation was conducted by direct method of measuring carbon exchange, making plots of representative sizes. Each plot was with important advantages: it is instantaneous and divided into smaller plots 10 m x 10 m. From these non destructive and it allows measurement of the smaller plots assessment and identification were total carbon gain by a plant separation of made for name of the tree species, number of photosynthetic gain from respiratory loss. This measurement of CO exchange involved enclosure species, stem diameter (>10 cm), height, 2 coordinate of the trees (x, y). For the unknown methods which is enclosure of a leaf in a species, samples were collected for identification. transparent chamber. The collected data was analyzed using Cox (1967) The rate of CO2 assimilation by the leaf and Greigh-Smith(1964) method for obtaining the enclosed is determined by measuring the change in values of basal area, relative frequency, relative the CO2 concentration of the air flowing across the density, relative dominance and value of chamber. In this close system air is pumped from importance by the following calculation: the chamber enclosing a leaf into an IRGA (Infra Red Gas Analyzer) which continuously records the BA = (0.5xD)2x3.14 CO2 concentration of the system. The air is then -2 -1 Where BA (m h ) is basal area; D is diameter and recycled back to the chamber. No air leaves or value of 3.14 is a constant. enters the system. If the leaf enclosed in the Plant biomass and carbon content were estimated chamber as photo-synthesizing, the CO2 by using the following calculation : concentration in the system will decline, and b continue to decline until the CO2 compensation W=aD point of photosynthesis is reached. The rate of
W= biomass; a and b = W= biomass; a and b = CO2 assimilation is equal to the change in the constant for estimating biomass of tree amount of CO2 in the system per unit time. community (a =0.19 and b=2.37); D = stem Rate of CO2 -assimilation was measured under diameter (Brown, 1997). certain range of CO2 concentrations, photon C = 0.5 W flux, and leaf temperature (Table 1). For C = carbon content; W = biomass. measurement of physiological characteristic, a fully expanded (young) and older leaves were C. Physiological Analysis chosen per sampling. Three different plants individuals of each species were measured. The measurement of physiological and Simultaneous measurements of microclimate, photosynthetic characteristics were carried out in photosynthesis, chlorophyll content and April and June 2010. The height of tree sampled transpiration rate were conducted. The measuring was 50 cm – 150 cm. Simultaneous measurements time for each species was between 09.00 - 12.00 am of CO2 assimilation, stomatal conductance and under completely clear sky. transpiration were conducted by using portable The measurement of microclimate was LCi ADC Bioscientific Ltd. Photosynthesis System. The conducted for each plant species. Air temperature
Table 1. Range of CO22 concentration in stomata, CO reference, foton flux, vapour pressure and temperature (leaf and chamber) during the measurement. Halimun National Park Parameters Cikaniki Citalahab CO2 reference (cref: vpm) 370 – 750 371 - 433 Analytical CO2 (can: vpm) 360 – 703 365 - 438 CO2 in stomata (ci: vpm) 260 – 500 345 - 412 Foton flux (Qleaf: µmolm-2s-) 8 – 250 10.25 - 957 Chamber temperature (Tch: oC) 24 – 29.5 21.6 – 30.8 Leaf temperature(Tie: oC) 29 – 30 20 - 29 Vapour pressure (P: mbar) 900 900
51 Journal of Forestry Research Vol. 9 No. 2, 2012: 49-61 and relative humidity were measured using Digital The rate of transpiration ranged between Thermohygrometer AS ONE TH-321, soil pH and 0.4175 - 1.8975 molm-2 s -1 . The highest was moisture content were measured usingSoil Tester , reached byEugenia sp.1 (kibeusi) which was and light intensity was measured by using Lux meter 1.8975, followed by Castanopsis acuminatissima Luxor. Leaf chlorophyll content was measured (kianak) 1.8950 molm-2 s -1 ,Quercus lineata (pasang using spectrophotometer and chlorophyll meter batarua) 1.7875 molm-2 s -1 ,Schima wallichii (puspa) SPAD-502; Minolta Co.Ltd., Osaka, Japan. 1.6225 molm-2 s -1 andLitsea noronhae (huru lumlum) which was 1.5300 molm-2 s -1 (Table 2). Chlorophyll content of the leaf varied between III. RESULTS AND DISCUSSION 0.6015 - 3.8370 mg chlorophyll. The highest leaf chlorophyll content was on Symplocos fasciculata A. Results (jirak) which was 3.8370, followed byLitsea sp.2 1. Cikaniki Site (huru hejo) 3.2862,Litsea brachystachia (huru hiris) Tree distribution in the higher site was 3.1232,Litsea sp.3 (huru buah) 2.7065 and Litsea dominated byAltingia excelsa (rasamala) which sp.1(huru bodas) 2.6268 (Table 2). contributed to 32% of the total basal area (BA), Variation in microclimate conditions during followed bySchima wallichii (puspa) (10%) and the measurements resulted in variation of the Quersus lineata (6%). measurements. The rate of CO2 assimilation was Microclimate and soil conditions during the affected by incident radiation intensity and thus measurements were presented in Appendix Table the quantum leaf (Q leaf) (Figure 2). Incident 1. Soil pH ranged from 5.58 - 6.60, soil moisture radiation and Q leaf also affected stomatal ranged from 55.5 - 90.3%, relative humidity was conductance and thus rate of transpiration. 62.5 - 29.4%, air temperature ranged from 23.5 - Correlation between stomatal conductance and 29.4o C and light intensity was 597.5 - 22893.5 lux. transpiration rate was R= 0.5741 (Figure 3). Under such microclimate and soil conditions CO2 assimilation of young leaves was not the overall CO2 assimilation range from 1.1913 - significantly different with that of older leaves, 31.3875 µmolm-2 s -1 . The highest rate was reached although stomatal resistance of young leaves was byLithocarpus sp.1 (pasang parengpeng) which was commonly higher than that of old leaves 31.3875 µmolm-2 s -1 , followed by Litsea noronhae (Appendix 2). Under such environmental -2 -1 (huru lumlum) 21.5750 µmolm s , Saurauia conditions CO2 assimilation was affected more by nudiflora(kilebo) 11.8175 µmolm-2 s -1 , Vernonia external factors (i.e. solar radiation) than by the arborea (hamirung) 6.7125 µmolm-2 s -1 and Litsea leaf stomatal character although some theory -2 -1 stated that stomatal conductance correlates with sp.1(huru bodas) which was 6.2725 µmolm s photosynthetic capacity (Wonget al ., 1979). Under (Table 2). certain range of Q leaf, there was a linear
Figure 2. Correlation between photon flux (QLeaf) and photosynthesis (Ps) In Cikaniki plot (names of species are presented in Table 2)
52 Physiological Characteristics Related to Carbon Sequestration of Tree Species tn Highland Forest Ecosystem ..... (Nuril Hidayati)
yl
1.96 1.09 1.70 2.53 1.81 1.04 2.03 1.21 2.71 1.47 2.45 1.80 2.63 3.84 2.04 3.29 1.25 0.60 2.10 2.30 1.32
1.92
3.12
2.61 2.94 1.86 1.93
Chloroph
content (mg)
Park
)
-1
s
-2
0.733 0.328 0.725 0.630 0.265 0.535 0.410 0.080 0.393 0.065 0.173 0.500 0.133 0.053 0.585 0.293 0.223 0.188 0.365 0.545 0.103
0.430
0.208
0.273 0.218 0.250 0.180
Stomatal
(molm
conductance
)
-1
s
-2
1.260 1.133 1.263 1.530 0.858 1.140 1.045 0.893 0.980 0.835 1.355 1.895 0.790 0.418 1.898 0.950 1.258 1.210 0.650 1.623 0.858
1.788
1.258
1.208 1.473 1.013 1.058
(molm
Transpiration
plot - Halimun-Salak National
)
-1
s
-2
2
CO
4.570 5.778 1.703 3.455 6.713
2.928 1.198 1.913 2.080 3.290 6.273 2.918 1.835 4.110 5.628 1.503 5.138 5.668 1.868
4.623
2.650
9.333
6.335 4.153
21.158 11.818
31.388
(µmolm
Assimilation
)
-1
s
-2
4.50 5.75
8.25 7.50
11.75 65.75
99.25 16.75 22.50 54.75 20.75 10.28 40.75 14.75 17.50 30.00 52.50
32.50
31.00
61.75 44.25 70.25 29.25
249.75 107.25
263.00
219.25
Q leaf
(µmolm
2
2
(vpm)
513.75 408.50 584.00 381.00 387.50 503.25 386.00 440.75 361.50 384.50 435.00 367.00 396.25 377.75 417.00 402.25 396.25 384.50 384.75 392.25 371.25 431.50
440.00
387.50 537.00 703.50 410.00
Analytical CO
tics related to CO absorption of tree Species in Cikaniki
peng)
ua)
(kalimorot)
characteris
(kianak)
(salam)
(jirak)
(saninten)
(marabodas)
(huru hiris)
hamirung)
(kilebo)
(hamirung)
(kiajag)
(pasang reuuey)
(pasang pareng
(puspa)
(rasamala)
ea (
(huru lumlum)
(pasang batar
(hawoyang)
(ceri)
ea
(kopo)
(kinolka)
gentea
(kisireum)
(kibonteng)
anthum
(kibeusi)
ea
sp.2
sp.1
ystachia
ea
(huru hejo)
(huru bodas)
(huru buah)
celsa
onhae
sp.1 sp.2
sp.3 sp.2 sp.1
Vernonea arbor
Prunus arbor Knema ciner Vernonia arbor Garcinia dioica Macaranga tanarius Litsea nor Symplocos fasciculata Saurauia nudiflora Eugenia opaca Castanopsis acuminatissima Eugenia Litsea Ardisia zollingeri Species (Local name) Litsea Eugenia Litsea Altingia ex Schima wallichii Quercus lineata Platea excelsa Lithocarpus
Litsea brach
Castanopsis ar
Lithocarpus Syzygium poly Lithocarpus pseudomoluccanus
1. 2.
4. 3. 5.
6.
7.
8.
9.
16. 18. 19. 15. 17. 20. 14. 23. 13. 22. 21. 24.
25. 26. 27.
10. 11. 12.
No.
Table 2. Variation in physiological
53 Journal of Forestry Research Vol. 9 No. 2, 2012: 49-61
Transpiration Rate (mmol m-2 s -1 ) Figure 3. Correlation between stomatal conductance and transpiration in Cikaniki Plot (R= 0.5741)
relationship between CO2 assimilation Q leaf. Microclimate conditions during the This findings agree with the report by Long and measurement were relatively low temperature hallgren (1993). There was positive correlation and low light intensity (Appendix 3). This between CO2 assimilation and leaf chlorophyll microclimate conditions resulted in low photon content although the value was not significant. flux (Q leaf) and relatively low in CO2 assimilation
However this relationship need to be studied rate. The highest CO2 assimilation rate in this area more thoroughly under controlled environment. was 8.77 µmolm-2 s -1 (Altingia excelsa ), followed by Tricalysia singularis which was 7.10 µmolm-2 s -1 and 2. Citalahab Site Litsea resinosa which was 6.22 µmolm-2 s -1 (Table 3 In Citalahab plot, there were 337 individual and Figure 4). Variation in transpiration rate and trees (diameter > 10 cm) which consists of 71 stomatal conductance were relatively small under species from 32 genus and 50 families. The most this environmental conditions. Under certain commonly family found were Lauraceae, conditions, there was positive correlation between Fagaceae, Myrtaceae, Rubiaceae, Meliaceae and Q leaf and CO2 assimilation and between stomatal Euphorbiaceae. conductance and transpiration rate although the There were 20 trees species that have the values were relatively low due to the uncontrolled highest “value of importance” including Altingia environmental factors. excelsa, Blumeodendron elateriospermum, Ardisia Dry matter and carbon content reached zollingeri, Gordonia excelsa, Tricalysia singularis, 152,247.91 (152.3 ton ha-1 ) obtained from basal Castanopsis acuminatissima, Knema cinerea, Laportea area of 28.89 m-2 ha -1 and the number of individu stimulans, Vernonia arboreaand Dysoxylum excelsum . 337 trees ha-1 . Five tree species with the highest Distribution of diameter classification was biomass were recorded i.e.:Altingia excelsa (938.25 dominated by class 10-20 cm, that reached ton ha-1 ),Blumeodendron elateriospermum (2882.31 51. 63% of the total individu. The most common ton ha-1 ),Castanopsis acuminatissima (1930.21 ton tree species within this class include Ardisia ha-1 ),Engelhardtia serrata (1608.47 ton ha -1 ) and zollingeri, Laportea stimulans, Gordonia excelsa and Vernonia arborea (1372.75 ton ha-1 ). The ecological Urophyllum glabrum. findings are discussed in different paper.
54 Physiological Characteristics Related to Carbon Sequestration of Tree Species tn Highland Forest Ecosystem ..... (Nuril Hidayati)
-2 -1
3.622 3.315 4.853 3.105 3.553 3.228 4.148 3.633 3.443 3.488 2.983 4.253 3.815 3.525 3.370 3.390 3.638 3.430 3.338 3.793 3.908 3.668 3.220 3.005 3.653
(molm s )
Transpiration
-2 -1
0.470 0.575 0.730 0.668 0.205 1.613 0.815 1.168 0.450 0.223 3.808 0.293 0.350 3.518 0.685 0.358 0.745 0.278 0.915 0.305 0.250 0.305 0.328 0.363
1.1875
Stomatal
(molm s )
conductance
un-Salak
s)
-2 -1
CO2
4.190 3.308 2.085 1.825 5.595 2.680 4.235 2.265 3.605 2.105 3.990 4.148 8.770 7.100 6.218 3.798 2.890 0.970 2.498 1.480 2.168 4.043 2.610 1.745 1.123
(µmolm
Assimilation
)
-1
s
-2
80.75 26.00 77.25 65.75 94.25 88.75 70.25 24.00 66.75 86.75 72.00 11.00 17.00 13.25
41.50 29.00 10.25
Q leaf
130.50 491.75 957.25 312.25 303.00 292.75
127.25 527.25
(µmolm
(vpm)
2
377.50 411.25 381.25 368.25 402.00 382.25 389.50 395.50 382.25 423.50 389.00 396.75 394.00 438.50 399.75 385.75 372.25 398.50 380.50 414.25 380.00 387.50 372.25 374.00 379.50
Analytical
CO
absorption of tree species in Citalahab - Halim
2
Local Name maja burununggul mara ganitri hamirung kianak kawoyang kimokla mumuncangan huru buah jirak kacapi rasamala dawolong huru minyak huru hiris puspa kibonteng pasang batarua bengang kokopian huru bodas saninten kiajag pasang parengpeng
tics related to CO
haracteris
mum
ysiological c
pa
ea
sp.
gentea
oetjape
ea
on elateriosper
gans
celsa
ystachia
ea
celsa
a triloba
ollingeri
Magnolia ele Blumeodendr Castanopsis acuminatissima Species Elaeocarpus ganitrus Knema ciner Gordonia ex Prunus arbor Symplocos fasciculata Vernonia arbor Altingia ex Tricalysia singularis Litsea resinosa Litsea brach Schima wallichii Platea latifolia Sandoricum k Cinnamomum Quercus lineata Neesia altissima Urophyllum corymbosum Acer laurinum Castanopsis ar Ardisia z Quersus oidocar Macarang
1. 2. 3. 4. 5. 6. 7. 8. 9.
10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25.
No.
Table 3. Variation in ph
55 Journal of Forestry Research Vol. 9 No. 2, 2012: 49-61
Figure 4. Correlation between photon flux (QLeaf) and photosynthesis (Ps) In Citalahab Plot (name of species is presented in Table 3)
B. Discussion molm-2 s -1 in Citalahab. Findings were reported S. balangeran A. The significant differences between the that gs of fast growing of and mangiumwere 0.49 molm-2 s -1 (Takahashi et al ., 2005; response of physiological characteristics and -2 -1 environmental conditions were represented by Takahashiet al ., 2006) and 1.3 molm s tree species. The response of physiological (Matsumotoet al ., 2003). The high gs play a role characteristic showed typical acclimation to sunlit for high capacity of ventilation due to high transpiration rate in open site, being able to avoid condition such as increasing CO2 assimilation rate. The results of this study agree with some extremely increase of leaf temperature. findings that leaf photosynthesis in trees fairly This study suggested that for evaluation of the variable with maximum values under natural appropriate tree species, morphological and conditions ranged from 3 - 30 µmolm-2 s - . The photosynthetic characteristic of leaves are -2 -1 suitable indicators. In general, sun leaf has higher values varies between 2 - 25 µmolm s for light saturated CO assimilation rate and lower deciduous broad leafed trees, 2 - 10 µmolm-2 s - for 2 apparent quantum yield of CO assimilation rate coniferous trees, 3 - 6 µmolm-2 s - for certain broad 2 compared with shade leaf (Boardman, 1977; leafed species such asQuersus and Fagus , more -2 - Larcher, 1995; Presset al ., 1996). Shade leaf has than 25 µmolm s for poplar, oil palms and high light-use efficiency for CO assimilation eucalypt (Raghavendra, 1991). Photosynthesis of 2 under low light condition due to high Shorea-2 - Shorea was reported of 7 - 21 µmolm s , accumulation light harvesting system in balangeran 21.9 µmolm-2 s - in Central Kalimantan, -2 - photosynthesis, however, under open condition, Acacia mangiumof 24.2 µmolm s , 16 for Hopea shade leaf does not have high light-use efficiency odorata, 27.8 µmolm-2 s - for Ochroma lagopus and the reduction of CO2 assimilation rate often (Chazdonet al ., 1996; Press et al ., 1996; occurs due to light oxidation by excess light Matsumotoet al. , 2003). Photosynthesis of energy called photo-inhibition (Clearwateret al ., tropical woody plants for the first stage of -2 - 1999). Furthermore, Presset al . (1996) succession ranged 10 - 20 µmolm s , scarcely 25 demonstrated that the degree of photosynthetic -2 - µmolm s (Larcher, 1995). plasticity in response to changes of light regimes The plasticity of stomatal conductance (gs) was high in the most-light demanding species, under relatively shade and lower temperature was therefore it is recommendable to select trees -2 -1 -2 -1 low, between 0.030 molm s to 0.263 molm s in which have higher CO2 assimilation rate of sunlit -2 -1 Cikaniki and between 0.043 molm s to 0.223 leaf and higher degree of plasticity.
56 Physiological Characteristics Related to Carbon Sequestration of Tree Species tn Highland Forest Ecosystem ..... (Nuril Hidayati)
Abiotic factors such as light, temperature, CO2 tree species. The overall CO2 absorption concentration, vapour pressure deficit and rate ranged 1.1913 - 31.3875 µmolm-2 s -1 , the nutrient status have a major effect on net highest rate was reached byLithocarpus sp.1 photosynthesis, and thus on growth and (pasang parengpeng) (31.3875 µmolm-2 s -1 ), productivity. All environmental conditions that followed byLitsea noronhae (huru lumlum) tend to reduce photosynthetic rate ( e.g. low light, (21.5750 µmolm-2 s -1 ),Saurauia nudiflora (kilebo) low temperature, low nutrient availability) reduce (11.8175 µmolm-2 s -1 ), Vernonia arborea the photosynthetic carbon gain (Ceulmens and (hamirung) (6.7125 µmolm-2 s -1 ) and Litsea sp.1 Sauger, 1991). (huru bodas) (6.2725 µmolm-2 s -1 ). Photosynthetic capacity varies not only with 2. Different microclimate conditions during the environment but also with age and position of the measurements resulted in variance CO2 leaves in the canopy. Stomatal conductance (and assimilation rate. The rate of CO assimilation net photosynthesis) inQuersus reached a 2 was affected by photon flux (Q leaf). Incident maximum several weeks after maximum leaf size. radiation and Q leaf also affected stomatal Leaves of 10-year old oil palm remainded conductance and thus rate of transpiration. photosynthetically active for 21 months. This has Correlation between stomatal conductance important implications for the whole-tree and transpiration under certain environmental photosynthetic CO uptake (Ceulmens and 2 condition was considerably high. Sauger, 1991). 3. Some remark need to be measurement of CO Some important remarks should be made 2 assimilation. about the correct interpretation of the values of photosynthetic rate. First, growth conditions as well as the experimental methods have important ACKNOWLEDGEMENT implication on CO2 exchange rate. Plant raised under natural conditions and/or measured in situ We thank to Mount Halimun-Salak National tend to have higher CO2 exchange rate than do Park for study site, cooperation and assistance plants grown under controlled environment such during data collection. Research Competitive as greenhouse condition. Therefore specification Programme The Indonesian Institute of Sciences on tree size, measurements conditions and for financial supports. methods used are mentioned in this paper. In many cases net photosynthesis has been found to be poorly correlated with growth rate for REFERENCES some reasons, i.e difference in leaf area, pattern of carbon partitioning and variation in wood and Ashton, M.S. 1998. Seedling Ecology of Mixed- root respiration rate. The harvestable product of Dipterocarp Forest.In : Appanah, S. and a tree (the stem) depends not only on the J.M. Thurnbull (Eds.), Review of photosynthetic carbon uptake by the foliage but Dipterocarps, Taxonomy, Ecology and also on respiration of the various organs and Silviculture. CIFOR, Bogor, pp. 89-98. carbon investment into renewable organs (leaves, Boardman, N.K. 1997. Comparative photo- fine roots) and non harvestable organs (branches synthesis of sun and shade plants. Annu. and large roots). Consequently, there is no Rev.Plant Physiol. 28: 57-107. obvious relationship between photosynthesis and Brown, S. 1997. Estimating biomass and biomass biomass production. However, a fast growing tree change of tropical forest, a primer. FAO needs a high photosynthesis, but the reverse is not Forestry Paper 134. FAO,Rome. necessarily true (Raghavendra, 1991). Chazdon R.L., R.W. Pearey, D.W. Lee and N. Fetcher. 1996. Photoshyntetic responses of IV.CONCLUSION tropical forest plants to contransting light environments.In : Mulkey S.S., R.L. 1. The results showed that there was a wide range Chazdon and A.P. Smith (Eds.) Tropical of variation of CO2 assimilation rate between
57 Journal of Forestry Research Vol. 9 No. 2, 2012: 49-61
Forest Plant Ecophysiology. Chapman and tropical tree species in Pasoh forest reserve. Hall, Newyork. pp. 5-55 In: Okuda, T., Manokaran, N., Matsumoto, Convention of Biological Diversity (CBD). Y., Niiyama, K., Thomas, S.C., Ashton, PS. 2008. Biodiversity: A Missing Link for (Eds.). Ecological of Lowland Rain Forest Mitigating Climate Change. World in Southeast Asia. Springer-Verlag, Tokyo, Environment Day Celebrated in Montreal pp. 241-250. (Press Release). Morenco, R.A. and G. Viera. 2005. Specific leaf Ceulmens R.J. and B. Sauger. 1991. Photo- area and photosynthetic parameters of tree synthesis.In : Raghavendra. A.S. 1991. species in the forest undestory as a function Physiology of Trees. Wiley & Sons Publ. of microsite light environment in Central New York 262p. Amazonia.J. Tropical Forest Science 17 : 265- 278. Clearwater, M.J., R. Susilawati, R. Effendi and P.R.van Gardingen. 1999. Rapid Press, M.C., N.D. Brown, M.G. Baker and S.W. photosynthetic acclimation of Shorea Zipperlen. 1996. Photosynthetic responses johorensis seedlings after logging to light in tropical rain forest tree seedlings. disturbance in Central Kalimantan. In: Swaine, M.D., (Ed.). The Ecology of Oceanologia 121: 478-488. Tropical Forest Tree Seedlings. UNESCO, Paris, pp. 41-58. Cox, G.W. 1967. Laboratory Manual of General Ecology. M.C. Crown, Iowa. Takahashi, K., M. Osaki, M. Shibuya, Y.Tamai, H. Saito, L.H. Swido, S.J.Tuah, A.R. Susanto, C. Greigh-Smith, P. 1964. Quantitative Plant Pidjath and P. Erosa. 2005. Growth Ecology. Second Edition. Butterworth, phenology and photosinthetic traits of tree London. species native to peat-swamp forest. Lacher, W.1995. Physiological Plant Ecology (3rd ). Annual Report: Environmental Springer, Berlin. Conservation and Land Use Management of Wetland Ecosystem in Southeast Asia. Long, S.P.and J.E. Hallgren. Measurement of CO2 assimilation by plants in the field and the p. 68-70. laboratory.In : Hall, D.O., M.O. Scurlock, Takahashi, K., M. Shibuya, Y. Tamai, H. Saito, H.R. Bolhar-Nordenkampf, R.C. Leegood L.H. Swido, S.J. Tuah, A.R. Susanto and P. and S.P. Long (Eds.). Photosynthesis and Erosa. 2006. Morphological and Production in a Changing Environment : photosynthetic characteristics of Shorea A Field and Laboratory Manual. Chapman selanicaand S. balangeran sapling planted at & Hall. 464 p. open and understory conditions on peat soil Matsumoto, Y., Y. Maruyama, A. Uemura, H. in Central Kalimantan. Annual Report: Shigenaga, S. Okuda, H. Harayama, H. Environmental Conservation and Land Use Kawarasaki, L.H. Ang and S.K. Yap. 2003. Management of Wetland Ecosystem in Gas exchange and turgon maintenance of Southeast Asia. p.62-68.
58 Physiological Characteristics Related to Carbon Sequestration of Tree Species tn Highland Forest Ecosystem ..... (Nuril Hidayati)
930.0 807.5 830.0 585.0
952.3
597.5
(Lux)
1307.5 1612.5 3530.0 1960.0 2410.0 1407.5 2227.5 2560.0 2427.5 2870.0 3492.5 5162.5 4780.0 1315.0 1355.0 2780.0 2327.5 3587.5 3455.0
22892.5
18845.0
Light Intencity
C)
o
Air
(
23.5 24.8 26.9 26.2 27.2 25.8 25.1 25.0 25.4 25.3 25.3 27.5 26.0 28.2 26.3 29.4 26.6 24.9 26.4 26.8 26.3 26.8 26.0 25.4 26.6 26.4 27.6
Temperature
88.5 86.5 69.3 78.3 73.2 80.3 77.0 82.8 76.5 81.5 74.8 70.2 78.0 67.3 70.3 70.8 69.3 72.9 68.3 71.5 69.8 73.8 71.4 77.5 70.5 78.5 62.5
Relative
Humidity (%)
(%)
77.5 78.0 73.0 82.0 79.5 77.0 79.8 73.8 83.3 85.3 67.0 67.0 72.5 77.5 65.0 67.5 53.8 71.3 78.3 55.5 72.5 62.8 72.5 68.8 90.3 81.8 84.3
Soil Moisture
6.6 6.4 5.8 6.3 6.4 6.1 6.4 6.0 6.5 5.8 6.1 6.3 6.0 5.8 6.0 6.1 6.4 5.9 6.1 6.2 6.2 6.1 6.0 6.2 5.6 6.2
5.83
282.0 236.5 443.0 264.0 283.0 396.0 241.5 261.5 363.5 404.5 347.0 492.0 214.0 280.5 192.5 458.5 313.0 416.5 288.5 424.0 270.0 354.5 114.0 313.0 265.0 318.0 212.5
Height (cm) Soil pH
1.9 1.3 2.0 2.6 2.9 1.8 1.6 2.4 3.1 2.9 6.3 1.7 2.4 1.3 3.9 2.0 4.5 3.8 4.8 2.0 1.4 1.9 1.9 1.8 1.9 3.4
(cm)
3. 8
Diameter
ize measured in Cikaniki plot
Microclimate and tree s
Local Name
1.2. kianak kibeusi 3. huru bodas 4. huru hejo 5. rasamala 6. Ppsang batarua 7. huru hiris 8. saninten 9. pasang parengpeng
15. hamirung 16. hawoyang 17. ceri 18. kinokla 19. kirung 20. marabodas 10. salam 21. kiajag 11. kalimorot 22. kopo 12. huru buah 23. jirak 13. kilebo 24. kisireum 14. huru lumlum 25. puspa 26. kibonteng 27. pasang reuney
No.
Appendix 1.
59 Journal of Forestry Research Vol. 9 No. 2, 2012: 49-61
0.25 0.33 0.74 2.05 0.83 0.39 0.46 0.05
0.25 0.16 0.31 0.13 0.21 0.59 0.47 0.32 0.07
0.66 0.08
0.49 0.08
0.16 0.18 0.18 1.83 0.16
0.59
Old
-2 -1
ms)
(mol
0.19 0.49 0.52 0.97 0.27 1.08 0.72 0.16
0.29 0.11 0.35 0.22 0.17 0.48 0.12 1.14 0.07
0.07
0.37 0.03
0.26 0.37 0.26 2.22 0.21
0.20
Young
Stomatal Conductance
0.080
-2 -1
ms)
1.45 1.18 1.46 1.83 1.97 1.37 1.86 0.83
0.83 0.78 1.23 1.12 1.36 1.04 1.09 1.06 0.82
0.81 0.92
1.90 0.47
1.30 0.96 1.31 0.91 1.05
0.81
Old
1.07 0.91 1.61 1.96 1.28 1.16 1.94 0.89
0.89 0.80 1.04 1.59 1.07 1.25 0.82 1.47 0.85
0.49 0.87
1.68 0.37
1.22 1.47 1.64 1.12 1.07
1.16
Young
Transpiration (mol
Old
1.05
1.19
4.35 3.09 1.28 1.49
2.35 4.34 1.54
4.32 5.62 1.66 2.68 0.92
4.15 1.79
3.49 3.45
3.09
6.21 2.48
6.07
6.46
12.27
4.470
27.66
33.37
-2 -1
ms)
Assimilation
4.79 3.49 2.39 2.25
1.06 8.21 1.47
5.06 3.91 1.29
6.13 2.38
5.76 2.39
2.21 5.19
6.39
8.21 2.10 3.38
10.15
14.66
21.00
29.41
2
12.00 10.75
22.59
Young
(µmol
CO
ves in Cikaniki plot
8.50
4.50 8.70
4.50
9.50
9.50
Old
-2 -1
24.00 88.00 17.00 47.50
12.50 42.50 55.00
71.00 35.50 68.00 69.00 54.50 12.00
37.00 21.50
29.00 16.00
47.50 88.00 84.50 37.00
ms)
4.50
7.00
5.50
36.00
28.00 11.85 57.50
11.00 67.00
46.00
25.50 77.00 21.50
28.00 20.00
33.00 13.50
76.00
56.00 21.50
Q Leaf (µmol
Young
126.50
183.50
228.00
257.50 129.50
161.00
128.00
tics of young and old lea
haracteris
Locak Name
aninten
alam
asamala
irak
puspa huru lumlum hamirung kianak kibeusi kibonteng
hawoyang huru bodas pasang reuney
ceri huru hejo kinokla kiajag marabodas kirung kopo
pasang batarua
huru hiris kisireum
pasang parengpeng
kalimorot huru buah
kileho
. Physiological c
2
1. 2. 3. 4. 5. r 6. 7. 8. s 9.
10. s 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. j 24. 25. 26. 27.
No.
Appendix
60 Physiological Characteristics Related to Carbon Sequestration of Tree Species tn Highland Forest Ecosystem ..... (Nuril Hidayati)
950.5
513.5
(Lux)
3800.0 1545.5 1580.0 4165.0 4970.0 1419.5 4200.0
3521.7 5745.0 5788.7 4060.0 1238.0
6102.5
35900.0 15245.0 25400.0 17950.0 11369.3 32522.5 15750.0 42600.0 26660.0
12037.5
Light Intencity
C)
o
Air
25.0 27.0 26.0 26.0 29.0 28.0 25.0 28.0 28.0 26.5 23.0 22.0 25.5 20.0 25.0 27.5 26.0 30.0 26.5 22.5 27.0 30.0 24.0 27.5 26.0
(
Temperature
50.0 62.0 57.0 50.0 50.0 58.0 51.0 58.0 60.0 51.5 68.0 65.0 48.5 72.0 70.0 57.5 53.0 40.0 60.0 59.0 49.0 39.0 60.5 55.0 61.5
Relative
Humidity (%)
(%)
30.0 40.0 41.0 40.0 60.0 10.0 10.0 48.0 10.0 20.0 80.0 50.0 20.0 50.0 50.0 30.0 40.0 75.0 60.0 52.5 10.0 60.0 90.0 80.0 35.0
Soil Moisture
6.0 7.0 6.0 6.1 5.8 4.5 4.5 7.0 5.8 6.8 6.6 4.0 5.5 6.5 5.5 5.8 6.5 5.5 4.0 5.0 5.5 5.0 3.7 4.0 5.5
Soil pH
262.0 254.0 144.0 382.5 376.5 504.0 364.0 197.5 127.0 231.5 136.0 204.0 132.0 241.5 133.0 157.0 369.0 147.0 121.5 623.5 416.0 148.5 146.5 185.0 415.5
2.1 1.2 1.7 3.9 3.3 2.7 2.9 1.5 0.7 3.1 1.1 2.8 1.4 1.7 2.6 2.2 2.4 1.1 0.8 4.3 2.4 1.1 1.8 1.4 4.4
Diameter (cm) Height (cm)
ize measured in Citalahab plot
peng
Local Name
. Microclimate and tree s
3
9. pasang batarua 8. puspa 5. hamirung 7. rasamala 4. kimolka 6. mumuncangan 3. ganitri 1.2. maja burununggul
12. saninten 14. pasang pareng 23. dawolong 11. huru bodas 13. kiajag 22. huru buah 24. kibonteng 10. kokopian 25. bengang 19. huru minyak 21. huru hiris 18. jirak 20. kacapi 17. kawoyang 16. kianak 15. mara
No.
Appendix
61