云南普洱季风常绿阔叶林优势物种不同生长阶段叶片碳、氮、磷化学计量特征

doi:10.17521/cjpe.2015.0006

植物生态学报 ›› 2015, Vol. 39 ›› Issue (1): 52-62.DOI: 10.17521/cjpe.2015.0006

所属专题：生态化学计量

云南普洱季风常绿阔叶林优势物种不同生长阶段叶片碳、氮、磷化学计量特征

刘万德^1,², 苏建荣^1,^2,^*(), 李帅锋^1,², 郎学东^1,², 张志钧^1,², 黄小波^1,²

¹中国林业科学研究院资源昆虫研究所, 昆明 650224
²国家林业局云南普洱森林生态系统国家定位观测研究站, 昆明 650224

收稿日期:2014-08-25 接受日期:2014-11-06 出版日期:2015-01-10 发布日期:2015-01-22
通讯作者: 苏建荣
作者简介:
# 共同第一作者
基金资助:
基金项目国家自然科学基金(31200461和3137- 0592)和林业公益性行业科研专项经费(201404211)资助

Leaf carbon, nitrogen and phosphorus stoichiometry at different growth stages in dominant tree species of a monsoon broad-leaved evergreen forest in Pu’er, Yunnan Province, China

LIU Wan-De^1,², SU Jian-Rong^1,^2,^*(), LI Shuai-Feng^1,², LANG Xue-Dong^1,², ZHANG Zhi-Jun^1,², HUANG Xiao-Bo^1,²

¹Research Institute of Resources Insect, Chinese Academy of Forestry, Kunming 650224, China
²Pu’er Forest Ecosystem National Research Station, State Forestry Administration, Kunming 650224, China

Received:2014-08-25 Accepted:2014-11-06 Online:2015-01-10 Published:2015-01-22
Contact: Jian-Rong SU
About author:
# Co-first authors

摘要/Abstract

摘要：

为探索植物叶片氮(N)、磷(P)、碳(C)生态化学计量特征随植物生长发育的变化规律, 在普洱季风常绿阔叶林中, 选取6种优势植物种(红锥(Castanopsis hystrix)、短刺锥(Castanopsis echidnocarpa)、泥柯(Lithocarpus fenestratus)、截果柯(Lithocarpus truncatus)、西南木荷(Schima wallichii)、茶梨(Anneslea fragrans))采集叶片, 分析其N、P、C含量及化学计量比随植物生长发育的变化。结果显示: 6种植物在不同生长阶段的N含量变化范围为7.90-17.72 mg·g^-1, P为0.34-1.39 mg·g^-1, C为458.48-516.87 mg·g^-1, C:N为28.04-65.70, N:P为11.41-63.50, C:P为355.23-1878.17, 且不同生长阶段6种植物及总体叶片N、P、C含量及其化学计量比变化趋势各异。在变异系数上, N:P比整体变异最大, 为36.46% (变化范围19.19%-91.65%), 其次为C:P, 为34.80% (变化范围15.99%-91.60%), C的整体变异最小, 为3.12% (变化范围1.61%-5.89%)。变异来源分析结果显示, N含量、C含量、C:N、N:P及C:P均主要受植物生长阶段的影响, 而P含量主要受物种与生长阶段的交互作用影响。

关键词: 化学计量学, 优势物种, 季风常绿阔叶林, 交互作用, 变异系数

Abstract: Aims

Our objective was to explore the effect of growth stage on leaf C:N:P stoichiometry in dominant species of a monsoon broad-leaved evergreen forest in Pu’er, Yunnan Province.

Methods

We collected leaves from six dominant species (i.e. Castanopsis hystrix, C. echidnocarpa, Lithocarpus fenestratus, L. truncatus, Schima wallichii and Anneslea fragrans) at different growth stages from seedlings to mature trees. The leaf N, P and C contents were measured. The effects of species, growth stage and their interaction on leaf C, N and P contents and stoichiometry were analyzed by Repeated-Measure ANOVA in SPSS 19.0.

<i>Important findings</i>

The ranges of leaf N, P and C contents and mass ratios of C:N, N:P and C:P in the six species across different growth stages were 7.90-17.72 mg·g^-1, 0.34-1.39 mg·g^-1, 458.48-516.87 mg·g^-1, 28.04- 65.70, 11.41-63.50 and 355.23-1878.17, respectively. The coefficient of variation (CV) for N:P mass ratio ranged between 19.19%-91.65%, with an average of 36.46%; CV for C:P varied between 15.99%-91.60%, with an average of 34.80%; whilst CV for C varied between 1.61%-5.89%, with an average of 3.12%. Factorial analysis of the variation for each variable, with growth stage and species as independent factors, showed that leaf N and C contents and the mass ratios of leaf C:N, N:P and C:P were mainly determined by growth stage, while leaf P content was mainly determined by an interaction between growth stage and species.

Key words: stoichiometry, dominant species, monsoon broad-leaved evergreen forest, interaction, coefficient of variation

刘万德, 苏建荣, 李帅锋, 郎学东, 张志钧, 黄小波. 云南普洱季风常绿阔叶林优势物种不同生长阶段叶片碳、氮、磷化学计量特征. 植物生态学报, 2015, 39(1): 52-62. DOI: 10.17521/cjpe.2015.0006

LIU Wan-De,SU Jian-Rong,LI Shuai-Feng,LANG Xue-Dong,ZHANG Zhi-Jun,HUANG Xiao-Bo. Leaf carbon, nitrogen and phosphorus stoichiometry at different growth stages in dominant tree species of a monsoon broad-leaved evergreen forest in Pu’er, Yunnan Province, China. Chinese Journal of Plant Ecology, 2015, 39(1): 52-62. DOI: 10.17521/cjpe.2015.0006

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https://www.plant-ecology.com/CN/Y2015/V39/I1/52

图/表 6

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物种 Species	科 Family	属 Genus
红锥 Castanopsis hystrix	壳斗科 Fagaceae	锥属 Castanopsis
短刺锥 C. echidnocarpa	壳斗科 Fagaceae	锥属 Castanopsis
泥柯 Lithocarpus fenestratus	壳斗科 Fagaceae	柯属 Lithocarpus
截果柯 L. truncatus	壳斗科 Fagaceae	柯属 Lithocarpus
西南木荷 Schima wallichii	山茶科 Theaceae	木荷属 Schima
茶梨 Anneslea fragrans	山茶科 Theaceae	茶梨属 Anneslea

物种 Species	科 Family	属 Genus
红锥 Castanopsis hystrix	壳斗科 Fagaceae	锥属 Castanopsis
短刺锥 C. echidnocarpa	壳斗科 Fagaceae	锥属 Castanopsis
泥柯 Lithocarpus fenestratus	壳斗科 Fagaceae	柯属 Lithocarpus
截果柯 L. truncatus	壳斗科 Fagaceae	柯属 Lithocarpus
西南木荷 Schima wallichii	山茶科 Theaceae	木荷属 Schima
茶梨 Anneslea fragrans	山茶科 Theaceae	茶梨属 Anneslea

参数 Para- meter	物种 Species	平均值 Mean	变异系数 Coefficient of variation (%)	极差 Range
N	茶梨 Anneslea fragrans	8.78	15.72	5.40
	红锥 Castanopsis hystrix	12.73	11.23	4.54
	短刺锥 C. echidnocarpa	14.20	13.66	8.65
	西南木荷 Schima wallichii	12.75	27.61	14.17
	泥柯 Lithocarpus fenestratus	17.01	7.47	5.72
	截果柯 L. truncatus	14.55	15.60	7.19
	合计 Total	13.24	24.24	17.85
P	茶梨 Anneslea fragrans	0.46	17.39	0.35
	红锥 Castanopsis hystrix	0.40	17.50	0.22
	短刺锥 C. echidnocarpa	0.71	66.20	1.41
	西南木荷 Schima wallichii	0.41	19.51	0.28
	泥柯 Lithocarpus fenestratus	1.02	63.73	1.71
	截果柯 L. truncatus	0.53	18.87	0.33
	合计 Total	0.59	40.68	1.71
C	茶梨 Anneslea fragrans	508.62	3.00	53.34
	红锥 Castanopsis hystrix	464.75	1.61	20.73
	短刺锥 C. echidnocarpa	480.35	1.97	41.36
	西南木荷 Schima wallichii	471.38	5.89	107.37
	泥柯 Lithocarpus fenestratus	502.91	3.75	57.73
	截果柯 L. truncatus	495.10	2.49	47.95
	合计 Total	487.18	3.12	107.37
C:N	茶梨 Anneslea fragrans	59.49	18.34	43.33
	红锥 Castanopsis hystrix	37.05	13.77	16.21
	短刺锥 C. echidnocarpa	34.36	12.17	17.70
	西南木荷 Schima wallichii	38.84	20.29	32.10
	泥柯 Lithocarpus fenestratus	29.68	6.84	8.99
	截果柯 L. truncatus	34.83	16.48	19.27
	合计 Total	39.30	15.19	66.81
N:P	茶梨 Anneslea fragrans	19.80	24.80	20.96
	红锥 Castanopsis hystrix	32.71	19.20	19.23
	短刺锥 C. echidnocarpa	25.56	38.93	33.09
	西南木荷 Schima wallichii	31.58	23.21	24.57
	泥柯 Lithocarpus fenestratus	31.60	91.65	71.06
	截果柯 L. truncatus	28.46	20.98	21.45
	合计 Total	28.28	37.38	71.77
C:P	茶梨 Anneslea fragrans	1 153.75	22.23	1100.84
	红锥 Castanopsis hystrix	1 195.89	15.99	622.73
	短刺锥 C. echidnocarpa	855.62	35.06	942.17
	西南木荷 Schima wallichii	1 195.06	22.49	911.85
	泥柯 Lithocarpus fenestratus	934.20	91.60	2075.46
	截果柯 L. truncatus	977.11	21.45	729.09
	合计 Total	1 051.94	32.98	2075.46

参数 Para- meter	物种 Species	平均值 Mean	变异系数 Coefficient of variation (%)	极差 Range
N	茶梨 Anneslea fragrans	8.78	15.72	5.40
	红锥 Castanopsis hystrix	12.73	11.23	4.54
	短刺锥 C. echidnocarpa	14.20	13.66	8.65
	西南木荷 Schima wallichii	12.75	27.61	14.17
	泥柯 Lithocarpus fenestratus	17.01	7.47	5.72
	截果柯 L. truncatus	14.55	15.60	7.19
	合计 Total	13.24	24.24	17.85
P	茶梨 Anneslea fragrans	0.46	17.39	0.35
	红锥 Castanopsis hystrix	0.40	17.50	0.22
	短刺锥 C. echidnocarpa	0.71	66.20	1.41
	西南木荷 Schima wallichii	0.41	19.51	0.28
	泥柯 Lithocarpus fenestratus	1.02	63.73	1.71
	截果柯 L. truncatus	0.53	18.87	0.33
	合计 Total	0.59	40.68	1.71
C	茶梨 Anneslea fragrans	508.62	3.00	53.34
	红锥 Castanopsis hystrix	464.75	1.61	20.73
	短刺锥 C. echidnocarpa	480.35	1.97	41.36
	西南木荷 Schima wallichii	471.38	5.89	107.37
	泥柯 Lithocarpus fenestratus	502.91	3.75	57.73
	截果柯 L. truncatus	495.10	2.49	47.95
	合计 Total	487.18	3.12	107.37
C:N	茶梨 Anneslea fragrans	59.49	18.34	43.33
	红锥 Castanopsis hystrix	37.05	13.77	16.21
	短刺锥 C. echidnocarpa	34.36	12.17	17.70
	西南木荷 Schima wallichii	38.84	20.29	32.10
	泥柯 Lithocarpus fenestratus	29.68	6.84	8.99
	截果柯 L. truncatus	34.83	16.48	19.27
	合计 Total	39.30	15.19	66.81
N:P	茶梨 Anneslea fragrans	19.80	24.80	20.96
	红锥 Castanopsis hystrix	32.71	19.20	19.23
	短刺锥 C. echidnocarpa	25.56	38.93	33.09
	西南木荷 Schima wallichii	31.58	23.21	24.57
	泥柯 Lithocarpus fenestratus	31.60	91.65	71.06
	截果柯 L. truncatus	28.46	20.98	21.45
	合计 Total	28.28	37.38	71.77
C:P	茶梨 Anneslea fragrans	1 153.75	22.23	1100.84
	红锥 Castanopsis hystrix	1 195.89	15.99	622.73
	短刺锥 C. echidnocarpa	855.62	35.06	942.17
	西南木荷 Schima wallichii	1 195.06	22.49	911.85
	泥柯 Lithocarpus fenestratus	934.20	91.60	2075.46
	截果柯 L. truncatus	977.11	21.45	729.09
	合计 Total	1 051.94	32.98	2075.46

参数 Parameter	生长阶段 Growth stages	平均值 Mean	变异系数 Coefficient of variation (%)	极差 Range
N	幼苗幼树 Seedling	12.92	22.45	12.27
	小树 Sapling	12.96	29.08	16.89
	中树 Medium tree	13.94	22.83	12.29
	大树 Mature tree	12.87	16.72	6.96
P	幼苗幼树 Seedling	0.45	35.22	0.76
	小树 Sapling	0.53	44.66	1.01
	中树 Medium tree	0.66	72.98	1.61
	大树 Mature tree	1.01	65.71	1.53
C	幼苗幼树 Seedling	484.09	5.06	77.23
	小树 Sapling	487.59	4.15	74.98
	中树 Medium tree	488.38	5.11	107.37
	大树 Mature tree	489.16	2.88	42.31
C:N	幼苗幼树 Seedling	39.77	30.18	58.16
	小树 Sapling	40.97	32.30	53.72
	中树 Medium tree	37.24	28.78	41.14
	大树 Mature tree	39.19	21.22	26.09
N:P	幼苗幼树 Seedling	33.39	54.41	63.98
	小树 Sapling	26.22	30.26	31.49
	中树 Medium tree	26.44	38.23	34.00
	大树 Mature tree	19.86	64.21	26.90
C:P	幼苗幼树 Seedling	1 225.22	38.58	1 808.29
	小树 Sapling	1 023.82	27.24	1 094.90
	中树 Medium tree	954.83	36.11	1 209.56
	大树 Mature tree	803.67	74.58	1 647.69

云南普洱季风常绿阔叶林优势物种不同生长阶段叶片碳、氮、磷化学计量特征

Leaf carbon, nitrogen and phosphorus stoichiometry at different growth stages in dominant tree species of a monsoon broad-leaved evergreen forest in Pu’er, Yunnan Province, China

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参数 Parameter	变异来源 Source of variation	离差平方和 Sum of squares of deviations (SS)	df	均方 Mean square (MS)	F	p
N (mg·g^-1)	物种 Species	515.68	5	103.14	11.38	0.00
	物种间误差 Error among species	217.49	24	9.06
	生长阶段 Growth stage	2 147.38	3	715.79	84.31	0.00
	生长阶段间误差 Error among growth stages	611.32	72	8.49
	物种×生长阶段 Species × Growth stage	757.75	15	50.52	5.95	0.00
P (mg·g^-1)	物种 Species	3.60	5	0.72	21.32	0.00
	物种间误差 Error among species	0.81	24	0.03
	生长阶段 Growth stage	2.40	3	0.80	16.89	0.00
	生长阶段间误差 Error among growth stages	3.41	72	0.05
	物种×生长阶段 Species × Growth stage	10.57	15	0.71	14.90	0.00
C (mg·g^-1)	物种 Species	266 602.48	5	53 320.50	7.13	0.00
	物种间误差 Error among species	179 485.43	24	7 478.56
	生长阶段 Growth stage	2 773 966.81	3	924 655.60	127.31	0.00
	生长阶段间误差 Error among growth stages	522 960.01	72	7 263.33
	物种×生长阶段 Species × Growth stage	712 916.28	15	47 527.75	6.54	0.00
C:N	物种 Species	10 483.35	5	2 096.67	32.81	0.00
	物种间误差 Error among species	1 533.77	24	63.91
	生长阶段 Growth stage	18 289.86	3	6 096.62	72.08	0.00
	生长阶段间误差 Error among growth stages	6 089.92	72	84.58
	物种×生长阶段 Species × Growth stage	5 421.81	15	361.45	4.27	0.00
N:P	物种 Species	1 155.88	5	231.18	2.24	0.08
	物种间误差 Error among species	2 472.71	24	103.03
	生长阶段 Growth stage	13 145.00	3	4 381.67	68.61	0.00
	生长阶段间误差 Error among growth stages	4 598.38	72	63.87
	物种×生长阶段 Species × Growth stage	10 189.61	15	679.31	10.64	0.00
C:P	物种 Species	1 654 291.72	5	330 858.35	2.87	0.04
	物种间误差 Error among species	2 766 919.49	24	115 288.31
	生长阶段 Growth stage	17 259 909.97	3	5 753 303.30	64.52	0.00
	生长阶段间误差 Error among growth stages	6 420 001.91	72	89 166.69
	物种×生长阶段 Species × Growth stage	8 581 371.25	15	572 091.42	6.42	0.00

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