生态系统演替过程中土壤与微生物碳氮磷化学计量关系的变化

doi:10.17521/cjpe.2016.0218

摘要/Abstract

摘要：

土壤碳(C)、氮(N)、磷(P)化学计量特征会显著影响微生物的生长、群落结构、生物量C:N:P化学计量及其代谢活动。然而生态系统演替过程中土壤-微生物C:N:P化学计量的时间格局及其协调关系还不明确。为此, 该研究收集了2016年5月以前发表的文献中19个生态系统演替序列(包括13个森林、6个草地生态系统)的土壤-微生物生物量C:N:P研究结果, 整合分析了其中土壤-微生物生态化学计量的时间动态, 结果表明: (1)生态系统演替过程中土壤C:N没有一致的时间格局, 而土壤C:P和N:P均随演替进程显著增加, 其中土壤C:N:P与演替时间之间线性关系的斜率与相应演替序列的初始土壤有机C含量呈负相关关系。(2)演替进程中土壤-微生物生物量C:N:P没有一致的时间格局。(3)微生物生物量C占土壤有机C百分比(qMBC)、微生物生物量N占土壤全N百分比、微生物生物量P占土壤全P百分比均随着演替进程而显著增加, 即单位资源所能支持的微生物生物量随着演替进程而增加, 这与宏观生态系统演替理论相符。(4) qMBC随着土壤C:N、C:P和N:P以及C:N、C:P和N:P化学计量不平衡性(即土壤C:N、C:P和N:P分别除以微生物生物量C:N、C:P和N:P)的增加而减小; 其中, C:N、C:P和N:P化学计量不平衡性解释了qMBC变异性的37%-57%, 是演替时间解释率的7-17倍, 表明土壤-微生物生态化学计量关系对qMBC演替动态有重要影响。该研究强调了生态化学计量学理论和生态系统演替理论在土壤微生物时间动态研究中的重要作用, 表明适当地融合生态学宏观理论于土壤微生物研究可以加深对土壤-微生物生态过程的认识。

关键词: 碳, 氮, 磷, 生态化学计量, 土壤微生物, 演替

Abstract: AimsThe carbon (C), nitrogen (N) and phosphorus (P) stoichiometry (C:N:P) of soil profoundly influences the growth, community structure, biomass C:N:P stoichiometry, and metabolism in microbes. However, the relationships between soil and microbes in the C:N:P stoichiometry and their temporal dynamics during ecosystem succession are poorly understood. The aim of this study was to determine the temporal patterns of soil and microbial C:N:P stoichiometry and their relationships during ecosystem succession.MethodsAn extensive literature search was conducted and data were compiled for 19 age sequences of successional ecosystems, including 13 forest ecosystems and 6 grassland ecosystems, from 18 studies published up to May 2016. Meta-analyses were performed to examine the sequential changes in 18 variables that were associated with soil and microbial C, N and P contents and the stoichiometry. Important findings (1) There was no consistent temporal pattern in soil C:N along the successional stages, whereas the soil C:P and N:P increased with succession; the slopes of the linear relationships between soil C:N:P stoichiometry and successional age were negatively correlated with the initial content of the soil organic C within given chronosequence. (2) There was no consistent temporal pattern in microbial C:N:P stoichiometry along the successional stages. (3) The fraction of microbial biomass C in soil organic C (qMBC), the fraction of microbial biomass N in soil total N, and the fraction of microbial biomass P in soil total P all increased significantly with succession, in consistency with the theory of succession that ecosystem biomass per unit resource increases with succession. (4) The qMBC decreased with increases in the values of soil C:N, C:P, or N:P, as well as the stoichiometric imbalances in C:N, C:P, and N:P between soil and microbes (i.e., ratios of soil C:N, C:P, and N:P to microbial biomass C:N, C:P, and N:P, respectively). The C:N, C:P, and N:P stoichiometric imbalances explained 37%-57% variations in the qMBC, about 7-17 times more than that explainable by the successional age, illustrating the importance of soil-microbial C:N:P stoichiometry in shaping the successional dynamics in qMBC. In summary, our study highlights the importance of the theories of ecosystem succession and stoichiometry in soil microbial studies, and suggests that appropriately applying macro-ecological theories in microbial studies may improve our understanding on microbial ecological processes.

Key words: carbon, nitrogen, phosphorus, ecological stoichiometry, soil microbes, succession

周正虎, 王传宽. 生态系统演替过程中土壤与微生物碳氮磷化学计量关系的变化. 植物生态学报, 2016, 40(12): 1257-1266. DOI: 10.17521/cjpe.2016.0218

Zheng-Hu ZHOU, Chuan-Kuan WANG. Changes of the relationships between soil and microbes in carbon, nitrogen and phosphorus stoichiometry during ecosystem succession. Chinese Journal of Plant Ecology, 2016, 40(12): 1257-1266. DOI: 10.17521/cjpe.2016.0218

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2016.0218

https://www.plant-ecology.com/CN/Y2016/V40/I12/1257

图/表 6

表1 满足本研究筛选要求的数据介绍

Table 1 Summary of the data sets compiled in this study

参考文献 Reference	生态系统类型 Ecosystem type	年龄阶段数 No. of age stages	演替年龄跨度 Span of successional age (a)	C_soil	N_soil	P_soil	C_mic	N_mic	P_mic
An et al., 2009	草地 Grassland	8	78	√	√	NA	√	√	NA
Banning et al., 2008	森林 Forest	7	27	√	√	NA	√	√	NA
	森林 Forest	7	27	√	√	NA	√	√	NA
Cao et al., 2008	森林 Forest	4	24	√	√	NA	√	√	NA
Hu et al., 2016	森林 Forest	5	13	√	√	NA	√	√	NA
Jia et al., 2005	森林 Forest	5	36	√	√	NA	√	√	NA
Jia, 2006	草地 Grassland	9	27	√	√	√	√	√	√
Jiang et al., 2007	草地 Grassland	6	24	√	√	NA	√	√	NA
Jiang et al., 2009	草地 Grassland	6	139	√	√	NA	√	√	NA
Liu et al., 2012	森林 Forest	5	39	√	√	NA	√	NA	√
Liu et al., 2010	森林 Forest	7	101	√	√	NA	√	√	√
Liu et al., 2013	森林 Forest	5	56	√	√	√	√	√	NA
Singh et al., 2001	森林 Forest	5	58	√	√	√	√	√	√
Xiao et al., 2013	草地 Grassland	8	30	√	√	NA	√	√	NA
Xue et al., 2008	森林 Forest	6	31	√	√	√	√	√	√
Xue et al., 2007	森林 Forest	9	51	√	√	√	√	√	√
Xue et al., 2009	草地 Grassland	11	51	√	√	√	√	√	√
Yang et al., 2014	森林 Forest	4	49	√	√	√	√	√	√
Zhu et al., 2012	森林 Forest	4	51	√	√	NA	√	√	NA

图1 不同演替序列土壤和微生物C、N、P的浓度(A)、化学计量比(B)、化学计量不平衡性(C), 以及微生物生物量元素占土壤元素百分比(D)与演替时间线性关系斜率的平均值和95%置信区间。图中点和误差线分别表示斜率的平均值和95%置信区间。括号内数字表示报道该变量的演替序列个数。Csoil, 土壤有机C; Nsoil, 土壤全N; Psoil, 土壤全P; Cmic, 微生物生物量C; Nmic, 微生物生物量N; Pmic, 微生物生物量P; C:Nsoil, 土壤C:N; C:Psoil, 土壤C:P; N:Psoil, 土壤N:P; C:Nmic, 微生物生物量C:N; C:Pmic, 微生物生物量C:P; N:Pmic, 微生物生物量N:P; C:Nimb, C:N化学计量不平衡性; C:Pimb, C:P化学计量不平衡性; N:Pimb, N:P化学计量不平衡性; qMBC, 微生物生物量C占土壤有机C百分比; qMBN, 微生物生物量N占土壤全N百分比; qMBP, 微生物生物量P占土壤全P百分比。

Fig. 1 Means and 95% confidence interval of the slopes of the linear relationships between soil and microbial C, N, P concentrations (A), stoichiometric ratios (B), stoichiometric imbalances (C), fraction of the elements in microbial biomass over the total amounts in soils (D) and the successional age. The dots and error bars represent the means and 95% confidence intervals, respectively; values in brackets are the numbers of age sequences. Csoil, soil organic C; Nsoil, soil total N; Psoil, soil total P; Cmic, microbial biomass C; Nmic, microbial biomass N; Pmic, microbial biomass P; C:Nsoil, soil C to N ratio; C:Psoil, soil C to P ratio; N:Psoil, soil N to P ratio; C:Nmic, microbial biomass C to N ratio; C:Pmic, microbial biomass C to P ratio; N:Pmic, microbial biomass N to P ratio; C:Nimb, C:N stoichiometric imbalance; C:Pimb, C:P stoichiometric imbalance; N:Pimb, N:P stoichiometric imbalance; qMBC, fraction of microbial biomass C in soil C; qMBN, fraction of microbial biomass N in soil total N; qMBP, fraction of microbial biomass P in soil total P.

图2 演替初始有机C含量和土壤C:N:P与演替时间线性关系斜率的关系。

Fig. 2 Correlations between the initial contents of soil organic C and the slopes of linear relationships between soil C:N:P and successional age.

图3 不同演替序列土壤C:N:P和微生物生物量C:N:P线性关系斜率的平均值和95%置信区间。图中点和误差线分别表示斜率的平均值和95%置信区间。纵坐标括号内数字表示报道该变量的演替序列个数。C:Nsoil vs. C:Nmic, 土壤C:N与微生物C:N的线性关系; C:Psoil vs. C:Pmic, 土壤C:P与微生物C:P的线性关系; N:Psoil vs. N:Pmic, 土壤N:P与微生物N:P的线性关系。

Fig. 3 Means and 95% confidence interval of the slopes of the linear relationships between soil C:N:P and microbial C:N:P. The dots and error bars represent the means and 95% confidence intervals, respectively; the values in brackets are the numbers of age sequences. C:Nsoil vs. C:Nmic, relationship between soil C:N and microbial C:N; C:Psoil vs. C:Pmic, relationship between soil C:P and microbial C:P; N:Psoil vs. N:Pmic, relationship between soil N:P and microbial N:P.

表2 广义线性模型分析演替时间、化学计量不平衡性和个体序列对微生物熵影响

Table 2 Summary of the results from a general linear model showing the integrative effects of successional ages, stoichiometric imbalances and individual sequence on microbial quotient

分组 Group	因子 Factor	平方和 Sum of square	df	F	p	方差解释率 Explanatory rate of variance (%)
C:N	演替时间 Successional age	1.8	1	28.1	<0.001	2
	C:N化学计量不平衡性 C:N stoichiometric imbalance	31.9	1	484.8	<0.001	37
	个体序列 Individual sequence	46.5	17	41.6	<0.001	54
	误差 Error	6.2	95			7
C:P	演替时间 Successional age	0.4	1	15.1	<0.001	5
	C:P化学计量不平衡性 C:P stoichiometric imbalance	3.7	1	157.6	<0.001	57
	个体序列 Individual sequence	1.7	5	14.3	<0.001	26
	误差 Error	0.8	34			12
N:P	演替时间 Successional age	0.4	1	9.8	0.004	5
	C:N化学计量不平衡性 C:N stoichiometric imbalance	2.7	1	73.2	<0.001	41
	个体序列 Individual sequence	2.3	5	12.6	<0.001	35
	误差 Error	1.2	34			19

图4 微生物熵(qMBC)和土壤C:N:P以及C:N:P化学计量不平衡性的关系。C:Nsoil, 土壤C:N; C:Psoil, 土壤C:P; N:Psoil, 土壤N:P; C:Nimb, C:N化学计量不平衡性; C:Pimb, C:P化学计量不平衡性; N:Pimb, N:P化学计量不平衡性。方框中数据点为异常值。

Fig. 4 Relationships between microbial quotient (qMBC) and soil C:N:P and C:N:P stoichiometric imbalances. C:Nsoil, soil C to N ratio; C:Psoil, soil C to P ratio; N:Psoil, soil N to P ratio; C:Nimb, C:N stoichiometric imbalance; C:Pimb, C:P stoichiometric imbalance; N:Pimb, N:P stoichiometric imbalance. The dots in the black box are outliers.

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