Carbon storage and its allocation in Betula platyphylla forests of different ages in cold temperate zone of China

doi:10.17521/cjpe.2019.0127

Abstract

Abstract:

Aims The objective of this study was to estimate the carbon storage and its allocation in the Betula platyphylla forests of three different ages (25, 40, 61-year-old) in the cold temperate zone, NE China. Methods Through analyzing the field data, we estimated the carbon storage and sequestration rates of tree, understory layer (shrub layer, herb layer, litter layer) and soil layer (0-100 cm) of the 25, 40 and 61-year-old Betula platyphylla ecosystems in the north section of the Da Hinggan Ling Mountains. Important findings The results showed that the carbon content of each organ in the tree layer of the forests ranged from 440.7 to 506.7 g·kg^-1, that decreased as the forest age increases. Carbon content in the shrub and herb layers decreased first and then increased as the forest aged, while that in the litter layer decreased with the increase of forest age. The carbon content in soil layer (0-100 cm) increased significantly with the forest age (p < 0.05), and decreased as soil drought intensified. The carbon storage of B. platyphylla ecosystem at all levels increased significantly with the increase of forest age. The carbon storage of tree layer in the forests of 25, 40 and 61-year-old were 11.9, 19.1 and 34.2 t·hm^-2, respectively. The carbon storage of the organs follow the order of: trunk > root > branch > leaf, and the allocation ratio of trunk carbon increased as the forest aged. The carbon storage in the Betula platyphylla forest ecosystems of 25, 40 and 61-year-old were 77.4, 180.9 and 271.4 t·hm ^-2, respectively. Soil layer, the main carbon pool of the ecosystems, accounted for 81.6%, 87.7% and 85.9% of the total carbon storage. The annual net productivity (2.0-4.4 t·hm ^-2·a ^-1) and annual net carbon sequestration (1.0-2.1 t·hm ^-2·a ^-1) of the forests increased with the age increase of the forest, and the old-growth B. platyphylla forests hold a strong carbon sequestration capacity.

Key words: Betula platyphylla forests, carbon storage, carbon sequestration, carbon content, forest age

WEI Hong, MAN Xiu-Ling. Carbon storage and its allocation in Betula platyphylla forests of different ages in cold temperate zone of China[J]. Chin J Plant Ecol, 2019, 43(10): 843-852.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2019.0127

https://www.plant-ecology.com/EN/Y2019/V43/I10/843

Figures/Tables 7

References 41

[1]	. Bradford JB, Kastendick DN (2010). Age-related patterns of forest complexity and carbon storage in pine and aspen- birch ecosystems of northern Minnesota, USA. Canadian Journal of Forest Research, 40, 401-409. DOI URL
[2]	. Chen HYH, Luo Y (2015). Net aboveground biomass declines of four major forest types with forest ageing and climate change in western Canada’s boreal forests. Global Change Biology, 21, 3675-3684. DOI URL PMID
[3]	. Ding GJ, Wang PC (2002). Study on change laws of biomass and productivity of Masson pine forest plantation II. Biomass and productivity of stand at different ages. Forest Research, 15(l), 54-60.
	[ 丁贵杰, 王鹏程 (2002). 马尾松人工林生物量及生产力变化规律研究II. 不同林龄生物量及生产力. 林业科学研究, 15(l), 54-60.]
[4]	. Fang JY, Chen AP, Peng CH, Zhao SQ, Ci LJ (2001). Changes in forest biomass carbon storage in China between 1949 and 1998. Science, 292, 2320-2322. DOI URL PMID
[5]	. Fang JY, Piao SL, Zhao SQ (2001). The carbon sink: The role of the middle and high latitudes terrestrial ecosystems in the Northern Hemisphere. Acta Phytoecologica Sinica, 25, 594-602.
	[ 方精云, 朴世龙, 赵淑清 (2001). CO₂失汇与北半球中高纬度陆地生态系统的碳汇. 植物生态学报, 25, 594-602.]
[6]	. Girardin MP, Bouriaud O, Hogg EH, Kurz W, Zimmermann NE, Metsaranta JM, de Jong R, Frank DC, Esper J, Büntgen U, Guo XJ, Bhatti J (2016). No growth stimulation of Canada’s boreal forest under half-century of combined warming and CO₂ fertilization. Proceedings of the National Academy of Sciences of the United States of America, 113, 8406-8414.
[7]	. Han YY, Huang W, Sun T, Lu B, Mao ZJ (2015). Soil organic carbon stocks and fluxes in different age stands of secondary Betula platyphylla in Xiaoxingʼan Mountain, China. Acta Ecologica Sinica, 35, 1460-1469.
	[ 韩营营, 黄唯, 孙涛, 陆彬, 毛子军 (2015). 不同林龄白桦天然次生林土壤碳通量和有机碳储量. 生态学报, 35, 1460-1469.]
[8]	. Hu HQ, Luo BZ, Wei SJ, Wei SW, Wen ZM, Sun L, Luo SS, Wang LM, Ma HB (2015). Estimating biological carbon storage of five typical forest types in the Daxingʼan Ling Mountains, Heilongjiang, China. Acta Ecologica Sinica, 35, 5745-5760.
	[ 胡海清, 罗碧珍, 魏书精, 魏书威, 文正敏, 孙龙, 罗斯生, 王立明, 马洪斌 (2015). 大兴安岭5种典型林型森林生物碳储量. 生态学报, 35, 5745-5760.]
[9]	. Huang GS, Ma W, Wang XJ, Xia CZ, Dang YF (2014). Carbon storage measurement of larch forest in northeastern China. Scientia Silvae Sinicae, 50(6), 167-174. DOI URL
	[ 黄国胜, 马炜, 王雪军, 夏朝宗, 党永锋 (2014). 东北地区落叶松林碳储量估算. 林业科学, 50(6), 167-174.] DOI URL
[10]	. Lan SB (2018). Comparison, selection and its progeny test of natural population of Betula platyphylla in northern China. Journal of Beihua University (Natural Science), 19, 582-587.
	[ 兰士波 (2018). 中国北方白桦自然群体比较选择及子代测定. 北华大学学报(自然科学版), 19, 582-587.]
[11]	. Li JQ, Ju H, Zhang QL (2010). Study on the Betula platyphylia natural forest biomass model. Journal of Inner Mongolia Agricultural University (Natural Science Edition), 31(1), 76-82.
	[ 李建强, 菊花, 张秋良 (2010). 白桦天然林生物量模型的研究. 内蒙古农业大学学报(自然科学版), 31(1), 76-82.]
[12]	. Li NN, Mu CC, Zheng T, Zhang Y, Cheng JY, Cao WL (2015). Effect of site types on carbon storage of natural white birch forest ecosystem in Changbai Mountains, Northeast China. Forest Research, 28, 618-626.
	[ 李娜娜, 牟长城, 郑瞳, 张毅, 程家友, 曹万亮 (2015). 立地类型对长白山天然白桦林生态系统碳储量的影响. 林业科学研究, 28, 618-626.]
[13]	. Li WY, Man XL, Zhang YW (2009). Soil properties and water conservation function of Betula platyphylla secondary forest with different stand ages. Science of Soil and Water Conservation, 7(5), 63-69.
	[ 李文影, 满秀玲, 张阳武 (2009). 不同林龄白桦次生林土壤特性及其水源涵养功能. 中国水土保持科学, 7(5), 63-69.]
[14]	. Li XD, Yi MJ, Son Y, Park PS, Lee KH, Son YM, Kim RH, Jeong MJ (2011). Biomass and carbon storage in an age-sequence of Korean pine (Pinus koraiensis) plantation forests in central Korea. Journal of Plant Biology,54, 33-42. DOI URL
[15]	. Liu LX, Wang J, Yang XJ, Liu CZ, Wang XW (2018). Forest plant community and soil organic carbon density in Da Xingʼan Mountains. Ecology and Environmental Sciences, 27, 1610-1616.
	[ 刘林馨, 王健, 杨晓杰, 刘传照, 王秀文 (2018). 大兴安岭不同森林群落植被多样性对土壤有机碳密度的影响. 生态环境学报, 27, 1610-1616.]
[16]	. Liu SN, Zhou T, Wei LY, Shu Y (2012). The spatial distribution of forest carbon sinks and sources in China. Chinese Science Bulletin, 57, 943-950, 987.
	[ 刘双娜, 周涛, 魏林艳, 舒阳 (2012). 中国森林植被的碳汇/源空间分布格局. 科学通报, 57, 943-950, 987.]
[17]	. Liu X, Man XL, Tian YH (2015). Hydro-chemical and nutrient importing characteristics of precipitation in secondary Betula platyphylla forests in northern Great Xing’an Mountains, northeastern China. Journal of Beijing Forestry University, 37(8), 83-89.
	[ 刘茜, 满秀玲, 田野宏 (2015). 白桦次生林降雨水化学及养分输入特征. 北京林业大学学报, 37(8), 83-89.]
[18]	. Lu BQ, Zang SY, Sun L (2019). The effects of freezing- thawing process on soil active organic carbon and nitrogen mineralization in Daxingʼanling Mountain forests. Acta Scientiae Circumstantiae, 39, 1664-1672.
	[ 鲁博权, 臧淑英, 孙丽 (2019). 冻融作用对大兴安岭典型森林土壤活性有机碳和氮矿化的影响. 环境科学学报, 39, 1664-1672.]
[19]	. Lü CQ, Sun SC (2004). A review on the distribution patterns of carbon density in terrestrial ecosystems. Acta Phytoecologica Sinica, 28, 692-703.
	[ 吕超群, 孙书存 (2004). 陆地生态系统碳密度格局研究概述. 植物生态学报, 28, 692-703.]
[20]	. Ma FF, Zhang CM, Li YZ (2016). Density and distribution of organic carbon of Larix kaempferi plantation ecosystem in Subtropics. Journal of Central South University of Forestry & Technology, 36(1), 94-100.
	[ 马丰丰, 张灿明, 李有志 (2016). 亚热带日本落叶松人工林生态系统碳密度及其分配特征. 中南林业科技大学学报, 36(1), 94-100.]
[21]	. Ma L (2012). Xiaoxing’anling Birch Natural Forest Ecosystem Carbon Storage Research. Master degree dissertation, Northeast Forestry University, Harbin.
	[ 马林 (2012). 小兴安岭白桦天然林生态系统碳储量的研究. 硕士学位论文, 东北林业大学, 哈尔滨.]
[22]	. Ming AG, Jia HY, Tian ZW, Tao Y, Lu LH, Cai DX, Shi ZM, Wang WX (2014). Characteristics of carbon storage and its allocation in Erythrophleum fordii plantations with different ages. Chinese Journal of Applied Ecology, 25, 940-946.
	[ 明安刚, 贾宏炎, 田祖为, 陶怡, 卢立华, 蔡道雄, 史作民, 王卫霞 (2014). 不同林龄格木人工林碳储量及其分配特征. 应用生态学报, 25, 940-946.]
[23]	. Pan YD, Birdsey RA, Fang JY, Houghton R, Kauppi PE, Kurz WA, Phillips OL, Shvidenko A, Lewis SL, Canadell JG, Ciais P, Jackson RB, Pacala SW, David McGuire A, Piao SL, Rautiainen A, Sitch S, Hayes D (2011). A large and persistent carbon sink in the world’s forests. Science, 333, 988-993. DOI URL
[24]	. Pregitzer KS, Euskirchen ES (2004). Carbon cycling and storage in world forests: Biome patterns related to forest age. Global Change Biology, 10, 2052-2077. DOI URL
[25]	. Qi G, Wang QL, Wang XC, Qi L, Wang QW, Ye YJ, Dai LM (2011). Vegetation carbon storage in Larix gmelinii plantations in Great Xing’an Mountains. Chinese Journal of Applied Ecology, 22, 273-279.
	[ 齐光, 王庆礼, 王新闯, 齐麟, 王庆伟, 叶雨静, 代力民 (2011). 大兴安岭林区兴安落叶松人工林植被碳贮量. 应用生态学报, 22, 273-279.]
[26]	. Raich JW, Russell AE, Kitayama K, Parton WJ, Vitousek PM (2006). Temperature influences carbon accumulation in moist tropical forests. Ecology, 87, 76-87. DOI URL PMID
[27]	. Shu Y, Zhou M, Zhao PW, Zeng N, Shi L, Wang ZX, Wang D, Ge P, Zhang B (2016). Carbon storage and its distribution characteristics in the planted forest of Larix principis in southern Daxinganling. Ecology and Environmental Sciences, 25, 1604-1611.
	[ 舒洋, 周梅, 赵鹏武, 曾楠, 石亮, 王梓璇, 王鼎, 葛鹏, 张波 (2016). 大兴安岭南段华北落叶松人工林碳储量及分配特征研究. 生态环境学报, 25, 1604-1611.]
[28]	. van Huysen TL, Perakis SS, Harmon ME (2016). Decomposition drives convergence of forest litter nutrient stoichiometry following phosphorus addition. Plant and Soil, 406, 1-14. DOI URL
[29]	. Wang Z, Liu GB, Xu MX, Zhang J, Wang Y, Tang L (2012). Temporal and spatial variations in soil organic carbon sequestration following revegetation in the hilly Loess Plateau, China. Catena, 99, 26-33. DOI URL
[30]	. Wu XR, Zhang JB, Tian Y, Li P (2014). Provincial agricultural carbon emissions in China: Calculation, performance change and influencing factors. Resources Science, 36, 129-138.
	[ 吴贤荣, 张俊飚, 田云, 李鹏 (2014). 中国省域农业碳排放: 测算、效率变动及影响因素研究——基于DEA-Malmquist指数分解方法与Tobit模型运用. 资源科学, 36, 129-138.]
[31]	. Xiang YX, Chen SK, Pan P, Ouyang XZ, Ning JK, Li Q (2019). Stoichiometric traits of C, N and P of leaf-litter-‌soil system of Pinus massoniana forest. Journal of Forest and Environment, 39(2), 120-126.
	[ 向云西, 陈胜魁, 潘萍, 欧阳勋志, 宁金魁, 李琦 (2019). 马尾松叶片-凋落物-土壤的碳氮磷化学计量特征. 森林与环境学报, 39(2), 120-126.]
[32]	. Xie XL, Sun B, Zhou HZ, Li ZP (2004). Soil carbon stocks and their influencing factors under native vegetations in China. Acta Pedologica Sinica, 41, 687-699.
	[ 解宪丽, 孙波, 周慧珍, 李忠佩 (2004). 不同植被下中国土壤有机碳的储量与影响因子. 土壤学报, 41, 687-699.]
[33]	. Yang LL, Wang YH, Wen SZ, Liu YH, Du M, Hao J, Li ZH (2015). Carbon and nitrogen storage and distribution in four forest ecosystems in Liupan Mountains, northwestern China. Acta Ecologica Sinica, 35, 5215-5227.
	[ 杨丽丽, 王彦辉, 文仕知, 刘延惠, 杜敏, 郝佳, 李振华 (2015). 六盘山四种森林生态系统的碳氮储量、组成及分布特征. 生态学报, 35, 5215-5227.]
[34]	. Zhang Y (2015). Effect of Site Types on Carbon Storage of Natural White Birch Forest Ecosystem in Xiaoxing’an Mountains of China. Master degree dissertation, Northeast Forestry University, Harbin.
	[ 张毅 (2015). 立地类型对小兴安岭天然白桦次生林生态系统碳储量的影响. 硕士学位论文, 东北林业大学, 哈尔滨.]
[35]	. Zhang Y, Mu CC, Zheng T, Li NN (2015). Ecosystem carbon storage of natural secondary birch forests in Xiaoxing’an Mountains of China. Journal of Beijing Forestry University, 37, 38-47.
	[ 张毅, 牟长城, 郑曈, 李娜娜 (2015). 小兴安岭天然白桦林生态系统碳储量. 北京林业大学学报, 37, 38-47.]
[36]	. Zhang ZJ, Zhang XQ, Wang YH, Luo YJ, Li ZY, Cao L (2009). Carbon storage and distribution of Pinus massoniana forest ecosystem in Tieshanping of Chongqing. Scientia Silvae Sinicae, 45(5), 49-53. DOI URL
	[ 张治军, 张小全, 王彦辉, 罗云建, 李志勇, 曹磊 (2009). 重庆铁山坪马尾松林生态系统碳贮量及其分配特征. 林业科学, 45(5), 49-53.] DOI URL
[37]	. Zheng T, Mu CC, Zhang Y, Li NN (2016). Effects of site condition on ecosystem carbon storage in a natural Betula platyphylla forest in the Zhangguangcai Mountains, China. Acta Ecologica Sinica, 36, 6284-6294.
	[ 郑瞳, 牟长城, 张毅, 李娜娜 (2016). 立地类型对张广才岭天然白桦林生态系统碳储量的影响. 生态学报, 36, 6284-6294.]
[38]	. Zhou XL, Cai Q, Xiong XY, Fang WJ, Zhu JX, Zhu JL, Fang JY, Ji CJ (2018). Ecosystem carbon stock and within-‌system distribution in successional Fagus lucida forests in Mt. Yueliang, Guizhou, China. Chinese Journal of Plant Ecology, 42, 703-712. DOI URL
	[ 周序力, 蔡琼, 熊心雨, 方文静, 朱剑霄, 朱江玲, 方精云, 吉成均 (2018). 贵州月亮山不同演替阶段亮叶水青冈林碳储量及其分配格局. 植物生态学报, 42, 703-712.] DOI URL
[39]	. Zhou Y, Hartemink AE, Shi Z, Liang ZZ, Lu YL (2019). Land use and climate change effects on soil organic carbon in North and Northeast China. Science of the Total Environment, 647, 1230-1238. DOI URL PMID
[40]	. Zhu JX, Hu HF, Tao SL, Chi XL, Li P, Jiang L, Ji CJ, Zhu JL, Tang ZY, Pan YD, Birdsey RA, He XH, Fang JY (2017 a). Carbon stocks and changes of dead organic matter in China’s forests. Nature Communications, 8, 151-160. DOI URL PMID
[41]	. Zhu JX, Zhou XL, Fang WJ, Xiong XY, Zhu B, Ji CJ, Fang JY (2017b) . Plant debris and its contribution to ecosystem carbon storage in successional Larix gmelinii forests in northeastern China. Forests, 8, 191-200. DOI URL

林龄 Stand age (a)	坡度 Slope (°)	密度 Stand density (No.·hm^-2)	郁闭度 Canopy density	平均树高 Mean tree height (m)	平均胸径 Mean DBH (cm)	林下主要植物 Main understory plants
25 ± 6	3 ± 1	1 825 ± 225	0.7 ± 0.1	8.6 ± 1.6	7.1 ± 1.8	1, 2, 3, 5, 6, 8
40 ± 4	2 ± 1	2 075 ± 300	0.9 ± 0.1	11.1 ± 3.0	9.5 ± 1.8	1, 2, 4, 5, 7, 9
61 ± 7	6 ± 2	1 820 ± 250	0.8 ± 0.1	15.2 ± 3.5	12.4 ± 2.2	1, 2, 4, 5, 6, 7, 8

林龄 Stand age (a)	坡度 Slope (°)	密度 Stand density (No.·hm^-2)	郁闭度 Canopy density	平均树高 Mean tree height (m)	平均胸径 Mean DBH (cm)	林下主要植物 Main understory plants
25 ± 6	3 ± 1	1 825 ± 225	0.7 ± 0.1	8.6 ± 1.6	7.1 ± 1.8	1, 2, 3, 5, 6, 8
40 ± 4	2 ± 1	2 075 ± 300	0.9 ± 0.1	11.1 ± 3.0	9.5 ± 1.8	1, 2, 4, 5, 7, 9
61 ± 7	6 ± 2	1 820 ± 250	0.8 ± 0.1	15.2 ± 3.5	12.4 ± 2.2	1, 2, 4, 5, 6, 7, 8

器官 Organ	回归方程 Regression equation	R²
树干 Stem	W = 0.046D^1.55H^1.163	0.969
树枝 Branch	W = 5.921D^1.925H^-1.54	0.793
树叶 Leaf	W = 0.012D^1.662H^0.49	0.711
树根 Root	W = 0.016D^2.171H^0.281	0.933

器官 Organ	回归方程 Regression equation	R²
树干 Stem	W = 0.046D^1.55H^1.163	0.969
树枝 Branch	W = 5.921D^1.925H^-1.54	0.793
树叶 Leaf	W = 0.012D^1.662H^0.49	0.711
树根 Root	W = 0.016D^2.171H^0.281	0.933

组分 Component		25 a			40 a			61 a
组分 Component		生物量 Biomass (t·hm^-2)	碳含量 Carbon Content (g·kg^-1)	碳储量 Carbon Storage (t·hm^-2)	生物量 Biomass (t·hm^-2)	碳含量 Carbon content (g·kg^-1)	碳储量 Carbon storage (t·hm^-2)	生物量 Biomass (t·hm^-2)	碳含量 Carbon content (g·kg^-1)	碳储量 Carbon storage (t·hm^-2)
乔木层 Arbor layer	树干 Stem	11.7 ± 0.6^Ac	490.3 ± 8.8 ^Ba	5.8 ± 0.5^Ac	24.8 ± 1.2^Ab	453.3 ± 8.7^Bb	11.2 ± 0.6^Ab	54.0 ± 2.7^Aa	440.7 ± 4.7^Db	23.8 ± 0.7^Aa
	树枝 Branch	9.4 ± 0.5^Bb	485.9 ± 8.4^Ca	4.6 ± 0.4^Bb	11.1 ± 0.5^Ba	473.7 ± 3.6^Aa	5.3 ± 0.3^Ba	11.4 ± 0.1^Ba	453.6 ± 6.8^Cb	5.2 ± 0.2^Ba
	树叶 Leaf	0.9 ± 0.1^Fc	506.7 ± 7.3^Ba	0.5 ± 0.0^Gc	1.7 ± 0.1^Db	484.2 ± 16.0^Ab	0.8 ± 0.0^Eb	3.0 ± 0.2^Da	467.9 ± 9.7^Cb	1.4 ± 0.0^Ea
	树根 Root	2.1 ± 0.1^Cc	505.9 ± 11.0^Ba	1.0 ± 0.1^Cc	3.9 ± 0.2^Cb	471.6 ± 12.5^Ab	1.8 ± 0.1^Cb	8.1 ± 0.5^Ca	465.6 ± 12.6^Cb	3.8 ± 0.1^Ca
	总计 Sum	24.1 ± 1.1^c		11.9 ± 1.0^c	41.5 ± 2.0^b		19.1 ± 1.0^b	76.5 ± 3.5^a		34.2 ± 1.0^a
灌木层 Shrub layer	地上部分 Aboveground	1.5 ± 0.1^Da	505.3 ± 1.7^Bb	0.8 ± 0.1^Db	1.0 ± 0.1^Eb	494.3 ± 14.9^Ab	0.5 ± 0.1^Fc	1.5 ± 0.1^Ea	575.3 ± 26.6^Aa	0.9 ± 0.2^Fa
	地下部分 Underground	1.2 ± 0.1^Ea	483.7 ± 1.9^Cb	0.6 ± 0.1^Fb	0.9 ± 0.0^Fb	465.7 ± 1.3^Bc	0.4 ± 0.1^Gc	1.3 ± 0.1^Fa	551.7 ± 7.5^Aa	0.7 ± 0.1^Ga
	总计 Sum	2.8 ± 0.2^a		1.4 ± 0.2^b	1.8 ± 0.1^b		0.9 ± 0.2^c	2.8 ± 0.2^a		1.6 ± 0.3^a
草本层 Herb layer	地上部分 Aboveground	0.2 ± 0.0^Gb	426.1 ± 2.9^Db	0.1 ± 0.0^Ib	0.7 ± 0.1^Ga	412.8 ± 2.7^Cc	0.3 ± 0.1^Ja	0.7 ± 0.0^Ga	462.4 ± 7.1^ABc	0.1 ± 0.0^Ib
	地下部分 Underground	0.1 ± 0.0^Hb	424.8 ± 1.5^Db	0.0 ± 0.0^Jc	0.5 ± 0.0^Ha	399.8 ± 9.1^Cc	0.2 ± 0.0^Ja	0.5 ± 0.0^Ha	471.5 ± 8.3^Ba	0.0 ± 0.0^Jb
	总计 Sum	0.2 ± 0.0^b		0.1 ± 0.0^b	1.1 ± 0.1^a		0.5 ± 0.1^a	0.2 ± 0.0^b		0.1 ± 0.0^b
凋落物层 Litter layer	未分解层 Undecomposed layer		544.0 ± 63.0^Aa	0.1 ± 0.0^Hc		478.1 ± 21.8^Ab	0.3 ± 0.1^Hb		433.0 ± 17.8^Dc	0.6 ± 0.1^Ha
	半分解层 partially- decomposed layer		486.3 ± 11.7^Ca	0.7 ± 0.1^Eb		488.1 ± 13.3^Aa	1.5 ± 0.2^Da		419.9 ± 2.4^Eb	1.8 ± 0.2^Da
	总计 Sum		515.2	0.8 ± 0.1^c		483.1	1.8 ± 0.3^b		426.4	2.3 ± 0.3^a