亚热带森林不同植物及器官来源的可溶性有机质输入对土壤激发效应的影响及其作用机理

doi:10.17521/cjpe.2021.0288

植物生态学报 ›› 2022, Vol. 46 ›› Issue (7): 797-810.DOI: 10.17521/cjpe.2021.0288

所属专题：稳定同位素生态学

亚热带森林不同植物及器官来源的可溶性有机质输入对土壤激发效应的影响及其作用机理

甘子莹¹, 王浩¹, 丁驰¹, 雷梅¹, 杨晓刚², 蔡敬琰¹, 丘清燕¹^,^*(), 胡亚林¹

¹福建农林大学林学院森林生态稳定同位素研究中心, 福州 350002
²内蒙古赤峰市克什克腾旗林业和草原局, 内蒙古赤峰 024005

收稿日期:2021-08-11 接受日期:2021-11-05 出版日期:2022-07-20 发布日期:2022-07-15
通讯作者: 丘清燕
作者简介:* (qingyan_qiu@126.com)
基金资助:
国家自然科学基金促进海峡两岸科技合作联合基金(U1805243);国家自然科学基金(42077094);国家自然科学基金(41703066)

Effects of dissolved organic matter derived from different plant and tissues in a subtropical forest on soil priming effect and the underlying mechanisms

GAN Zi-Ying¹, WANG Hao¹, DING Chi¹, LEI Mei¹, YANG Xiao-Gang², CAI Jing-Yan¹, QIU Qing-Yan¹^,^*(), HU Ya-Lin¹

¹Forest Ecological Stable Isotope Research Center, Forestry College, Fujian Agricultural and Forestry University, Fuzhou 350002, China
²Forestry and Grassland Bureau of Keshiketeng Banner, Chifengshi, Inner Mongolia, Chifeng, Nei Mongol 024005, China

Received:2021-08-11 Accepted:2021-11-05 Online:2022-07-20 Published:2022-07-15
Contact: QIU Qing-Yan
Supported by:
National Natural Science Foundation of China for Promoting Cross Strait Scientific and Technological Cooperation(U1805243);National Natural Science Foundation of China(42077094);National Natural Science Foundation of China(41703066)

摘要/Abstract

摘要：

外源有机物的输入可以通过正负激发效应影响土壤有机碳(SOC)的矿化。然而, 当前的研究较少考虑不同植物及器官来源可溶性有机质(DOM)输入对土壤激发效应的影响及其作用机理。该研究以武夷山森林土壤为研究对象, 以室内培养的方式向土壤中添加¹³C标记青冈(Cyclobalanopsis glauca)、杉木(Cunninghamia lanceolata)、木莲(Manglietia fordiana)和相思(Acacia confusa)这4种植物的根和叶来源DOM, 研究不同植物及器官来源DOM输入对土壤激发效应的影响及其作用机理。主要结果: 不同植物及器官来源DOM添加初期加快了SOC的矿化, 呈现正激发效应, 随后转为负激发效应。从整个培养期(90天)的累积激发效应来看, DOM的输入均抑制了SOC的矿化, 使其矿化量减少22%-49%, 其中青冈根DOM输入使SOC的矿化量减少最多, 而由木莲叶DOM输入减少的SOC矿化量最少。DOM输入引起的土壤激发效应强度受不同植物器官影响明显, 具体表现在植物根来源DOM输入所引起的土壤激发效应强度显著高于植物叶来源DOM输入所引起的激发效应强度(相思除外)。DOM的输入总体上提高了土壤微生物生物量碳(MBC)含量、土壤β-葡萄糖苷酶活性、纤维素酶活性以及土壤有效氮含量, 而对微生物群落组成无明显影响。从结构方程模型来看, DOM输入所引起的土壤激发效应主要受土壤微生物对外源碳的利用(¹³C-MBC)、纤维素酶活性以及土壤有效氮含量的影响, 这些因子的变化可解释植物叶来源DOM和根来源DOM添加处理下土壤激发效应变化的68%和86%。该研究结果表明在土壤氮充足的条件下, DOM的输入可以通过提高微生物生物量、土壤酶活性来加快分解所添加的外源有机物, 从而减少了对SOC的分解。因此, 在该研究中“底物优先利用”是土壤激发效应的主要作用机理。

关键词: 可溶性有机质, 激发效应, 酶活性, 土壤有机碳矿化, 亚热带植物

Abstract:

Aims The input of exogenous organic matter can affect the mineralization of soil organic carbon (SOC) through positive or negative priming effects. However, few studies have considered the effect of dissolved organic matter (DOM) derived from different plant and tissues on soil priming effect and revealed the underlying mechanisms.

Methods In this study, we investigated the different priming effects of ¹³C-labeled DOM derived from roots and leaves of different plants (i.e., Cyclobalanopsis glauca, Cunninghamia lanceolata, Manglietia fordiana and Acacia confusa) on SOC mineralization and clarifying the underlying mechanisms via an incubation experiment of soils sampled from Wuyi Mountain.

Important findings Inputs of DOM derived from different plant and tissues all accelerated the mineralization of SOC, exhibiting a positive priming effect at the initial stage after DOM input, and then showed a negative priming effect. For the cumulative priming effect during the whole incubation period (90 d), the input of DOM inhibited the mineralization of SOC, with the reduction magnitude ranging from 22% to 49%. Among them, the input of DOM derived from roots of Cyclobalanopsis glauca had the most pronounced effect on the reduction of SOC mineralization, while the input of DOM derived from leaves of M. glauca had the least effect on reduction of SOC mineralization. The intensity of soil priming effect induced by DOM was significantly affected by different plant tissues. DOM derived from plant roots showed more pronounced negative priming effect than DOM derived from plant leaves. In general, DOM input increased soil microbial biomass carbon (MBC) and soil β-glucosidase activities and cellobiohydrolase activities and soil available nitrogen content, but had no significant effect on the composition of soil microbial community. The structural equation model showed that soil priming effect induced by DOM was mainly affected by soil ¹³C-MBC, cellobiohydrolase activity and soil available nitrogen content. Changes in these factors could explain 68% and 86% of the variation of priming effect induced by plant leaf-derived DOM and root-derived DOM, respectively. The results suggested that if the soil is rich in available nitrogen, DOM input can accelerate the decomposition of exogenous organic matter through increasing microbial biomass and soil enzyme activity, and thus reducing the decomposition of SOC. Therefore, “substrate preferential utilization” is the main mechanism of soil priming effect in this study.

Key words: dissolved organic matter, priming effect, enzyme activity, soil organic carbon mineralization, subtropical plants

甘子莹, 王浩, 丁驰, 雷梅, 杨晓刚, 蔡敬琰, 丘清燕, 胡亚林. 亚热带森林不同植物及器官来源的可溶性有机质输入对土壤激发效应的影响及其作用机理. 植物生态学报, 2022, 46(7): 797-810. DOI: 10.17521/cjpe.2021.0288

GAN Zi-Ying, WANG Hao, DING Chi, LEI Mei, YANG Xiao-Gang, CAI Jing-Yan, QIU Qing-Yan, HU Ya-Lin. Effects of dissolved organic matter derived from different plant and tissues in a subtropical forest on soil priming effect and the underlying mechanisms. Chinese Journal of Plant Ecology, 2022, 46(7): 797-810. DOI: 10.17521/cjpe.2021.0288

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0288

https://www.plant-ecology.com/CN/Y2022/V46/I7/797

图/表 10

表1 亚热带森林不同植物及器官来源可溶性有机质(DOM)的基本化学性质(平均值±标准误, n = 3)

Table 1 Basic chemical properties of dissolved organic matter (DOM) derived from different plant and tissues in a subtropical forest (mean ± SE, n = 3)

植物器官 Plant tissue	碳(C)含量 Carbon (C) content (g·kg^-1)	氮(N)含量 Nitrogen (N) content (g·kg^-1)	C:N	稳定碳同位素比值 δ¹³C (‰)
CGL	426.70 ± 1.17^a	3.79 ± 0.05^g	112.54 ± 0.13^a	32.16 ± 0.05^g
CGR	362.52 ± 6.80^c	4.71 ± 0.04^f	77.03 ± 0.72^c	26.83 ± 0.05^h
CLL	380.69 ± 17.98^b	4.43 ± 0.15^fg	85.96 ± 1.16^b	40.25 ± 0.01^e
CLR	355.38 ± 0.29^cd	17.21 ± 0.09^c	20.65 ± 0.12^e	36.37 ± 0.03^f
ACL	351.61 ± 0.81^cd	50.11 ± 0.56^a	7.02 ± 0.08^g	44.36 ± 0.10^d
ACR	342.23 ± 3.36^de	15.08 ± 0.08^d	22.69 ± 0.14^e	53.92 ± 0.10^a
MFL	362.64 ± 0.81^c	7.52 ± 0.24^e	48.28 ± 1.63^d	49.30 ± 0.20^c
MFR	329.10 ± 0.15^e	19.62 ± 0.10^b	16.77 ± 0.08^f	49.89 ± 0.20^b

图1 亚热带森林土壤有机碳(SOC)来源、植物可溶性有机质(DOM)来源累积CO2排放动态以及不同碳源对CO2排放的贡献(平均值±标准误, n = 3)。ACL, 相思叶; ACR, 相思根; CGL, 青冈叶; CGR, 青冈根; CK, 对照; CLL, 杉木叶; CLR, 杉木根; MFL, 木莲叶; MFR, 木莲根。**和*分别表示因子对该指标有极显著(p < 0.01)和显著(p < 0.05)影响, ns表示因子对该指标的影响不显著。不同小写字母表示不同处理下来自土壤的CO2排放量差异显著(p < 0.05), 不同大写字母则表示不同处理下来自DOM的CO2排放量差异显著(p < 0.05)。

Fig. 1 Dynamics of cumulative CO2 emissions derived from soil organic carbon (SOC) and dissolved organic matter (DOM) in a subtropical forest and contribution of different carbon sources to CO2 emissions (mean ± SE, n = 3). ACL, Acacia confusa leaf; ACR, Acacia confusa root; CGL, Cyclobalanopsis glauca leaf; CGR, Cyclobalanopsis glauca root; CK, control; CLL, Cunninghamia lanceolata leaf; CLR, Cunninghamia lanceolata root; MFL, Manglietia fordiana leaf; MFR, Manglietia fordiana root. ** and * represent the factor has significant effect on the index at p < 0.01 and p < 0.05, respectively. ns represents no significant effect. Different lowercase letters indicate CO2 emissions derived from soil was significantly different under different treatments (p < 0.05), and different uppercase letters indicate the CO2 emissions derived from DOM were significantly different under different treatments (p < 0.05).

图2 亚热带森林不同植物及器官来源可溶性有机质(DOM)添加对土壤激发速率与土壤累积激发效应的影响(平均值±标准误, n = 3)。ACL, 相思叶; ACR, 相思根; CGL, 青冈叶; CGR, 青冈根; CK, 对照; CLL, 杉木叶; CLR, 杉木根; MFL, 木莲叶; MFR, 木莲根。**和*分别表示因子对该指标有极显著(p < 0.01)和显著(p < 0.05)影响, ns表示因子对该指标的影响不显著。

Fig. 2 Effect of dissolved organic matter (DOM) derived from different plant and tissues in a subtropical forest on soil priming rate and soil cumulative priming effect (mean ± SE, n = 3). ACL, Acacia confusa leaf; ACR, Acacia confusa root; CGL, Cyclobalanopsis glauca leaf; CGR, Cyclobalanopsis glauca root; CK, control; CLL, Cunninghamia lanceolata leaf; CLR, Cunninghamia lanceolata root; MFL, Manglietia fordiana leaf; MFR, Manglietia fordiana root. ** and * represent the factor has significant effect on the index at p < 0.01 and p < 0.05, respectively. ns represents the factor has no significant effect on the index.

图3 亚热带森林不同植物及器官来源可溶性有机质(DOM)添加对土壤β-葡萄糖苷酶和纤维素酶活性的影响(平均值±标准误, n = 3)。ACL, 相思叶; ACR, 相思根; CGL, 青冈叶; CGR, 青冈根; CK, 对照; CLL, 杉木叶; CLR, 杉木根; MFL, 木莲叶; MFR, 木莲根。**和*分别表示因子对该指标有极显著(p < 0.01)和显著(p < 0.05)影响, ns表示因子对该指标的影响不显著。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 3 Effects of dissolved organic matter (DOM) derived from different plant and tissues in a subtropical forest on soil β-glucosidase and cellobiohydrolase activities (means ± SE, n = 3). ACL, Acacia confusa leaf; ACR, Acacia confusa root; CGL, Cyclobalanopsis glauca leaf; CGR, Cyclobalanopsis glauca root; CK, control; CLL, Cunninghamia lanceolata leaf; CLR, Cunninghamia lanceolata root; MFL, Manglietia fordiana leaf; MFR, Manglietia fordiana root. ** and * represent the factor has significant effect on the index at p < 0.01 and p < 0.05, respectively. ns represents no significant effect. Different lowercase letters indicate significant difference among different treatments (p < 0.05).

图4 亚热带森林不同植物及器官来源可溶性有机质(DOM)添加对土壤微生物生物量碳与13C-MBC含量的影响(平均值±标准误, n = 3)。ACL, 相思叶; ACR, 相思根; CGL, 青冈叶; CGR, 青冈根; CK, 对照; CLL, 杉木叶; CLR, 杉木根; MFL, 木莲叶; MFR, 木莲根。**和*分别表示因子对该指标有极显著(p < 0.01)和显著(p < 0.05)影响, ns表示因子对该指标的影响不显著。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 4 Effects of dissolved organic matter (DOM) derived from different plant and tissues in a subtropical forest on soil MBC and 13C-MBC content (means ± SE, n = 3). ACL, Acacia confusa leaf; ACR, Acacia confusa root; CGL, Cyclobalanopsis glauca leaf; CGR, Cyclobalanopsis glauca root; CK, control; CLL, Cunninghamia lanceolata leaf; CLR, Cunninghamia lanceolata root; MFL, Manglietia fordiana leaf; MFR, Manglietia fordiana root. ** and * represent the factor has significant effect on the index at p < 0.01 and p < 0.05, respectively. ns represents no significant effect. Different lowercase letters indicate significant difference among different treatments (p < 0.05).

图5 亚热带森林不同植物及器官来源可溶性有机质(DOM)添加对土壤微生物磷酸脂肪酸(PLFAs)含量的影响(平均值±标准误, n = 3)。ACL, 相思叶; ACR, 相思根; CGL, 青冈叶; CGR, 青冈根; CK, 对照; CLL, 杉木叶; CLR, 杉木根; MFL, 木莲叶; MFR, 木莲根。**和*分别表示因子对该指标有极显著(p < 0.01)和显著(p < 0.05)影响, ns表示因子对该指标的影响不显著。

Fig. 5 Effects of dissolved organic matter (DOM) derived from different plant and tissues in a subtropical forest on soil microbial phosphate fatty acids (PLFAs) content (means ± SE, n = 3). G+, Gram-positive bacteria; G-, Gram-negative bacteria. ACL, Acacia confusa leaf; ACR, Acacia confusa root; CGL, Cyclobalanopsis glauca leaf; CGR, Cyclobalanopsis glauca root; CK, control; CLL, Cunninghamia lanceolata leaf; CLR, Cunninghamia lanceolata root; MFL, Manglietia fordiana leaf; MFR, Manglietia fordiana root. ** and * represent the factor has significant effect on the index at p < 0.01 and p < 0.05, respectively. ns represents no significant effect.

图6 亚热带森林不同植物及器官来源可溶性有机质(DOM)添加对土壤有效氮含量的影响(平均值±标准误, n = 3)。ACL, 相思叶; ACR, 相思根; CGL, 青冈叶; CGR, 青冈根; CK, 对照; CLL, 杉木叶; CLR, 杉木根; MFL, 木莲叶; MFR, 木莲根。**和*分别表示因子对该指标有极显著(p < 0.01)和显著(p < 0.05)影响, ns表示因子对该指标的影响不显著。不同小写字母表示不同处理间差异显著(p < 0.05)。

Fig. 6 Effects of dissolved organic matter (DOM) derived from different plant and tissues in a subtropical forest on soil available nitrogen (means ± SE, n = 3). ACL, Acacia confusa leaf; ACR, Acacia confusa root; CGL, Cyclobalanopsis glauca leaf; CGR, Cyclobalanopsis glauca root; CK, control; CLL, Cunninghamia lanceolata leaf; CLR, Cunninghamia lanceolata root; MFL, Manglietia fordiana leaf; MFR, Manglietia fordiana root. ** and * represent the factor has significant effect on the index at p < 0.01 and p < 0.05, respectively. ns represents no significant effect. Different lowercase letters indicate significant difference among different treatments (p < 0.05).

图7 亚热带森林植物叶和植物根来源可溶性有机质(DOM)输入对土壤激发效应影响的结构方程模型(SEM)。实线表示正向路径, 虚线为负向路径, 线段粗细表示因果关系的强弱。箭头相邻的数字表示关系的标准化路径系数(*, p < 0.05; **, p < 0.01; ***, p < 0.001), 实线箭头和虚线箭头分别表示拟合SEM中的正负相关。双向箭头表示变量之间的协方差。模型解释的总变异量用R2表示。AGFI, 调整拟合优度指数; 13C-MBC, 植物来源微生物生物量碳; GFI, 拟合优度指数; RMSEA, 近似均方根误差。

Fig. 7 Structural equation model (SEM) for the effect of dissolved organic matter (DOM) derived from roots and leaves of plant input on soil priming effect in a subtropical forest. Solid line is positive path and dashed line is negative path. Arrow line thickness indicates the strength of the causal relationship. Numbers adjacent to arrows represented standardized path coefficients of the relationships (*, p < 0.05; **, p < 0.01; ***, p < 0.001). Solid and dashed arrows indicated positive and negative relationships in a fitted SEM, respectively. Double-sided arrows represented covariance between variables. The total variation explained by the model is indicated by R2. AGFI, adjusted goodness of fit index; 13C-MBC, plant-derived microbial biomass carbon; G-, Gram-negative bacteria; GFI, goodness of fit index; RMSEA, root mean square error of approximation.

表2 亚热带森林土壤激发效应与土壤生化指标的关系

Table 2 Relationship between soil priming effect and soil biochemical indices in a subtropical forest

可溶性有机质来源 Dissolved organic matter source	土壤生化指标 Soil biochemical index	标准化直接效应值 Standardized direct effect	标准化间接效应值 Standardized indirect effect	标准化总效应值 Standardized total effect	p
叶 Leaf	土壤有效氮 Soil available nitrogen	0.000	0.466	0.466	<0.01
	细菌 Bacteria	-0.121	-0.078	-0.199	<0.05
	纤维素酶 Cellobiohydrolase	-0.547	0.000	-0.547	<0.01
	植物来源微生物生物量碳 ¹³C-MBC	-0.346	-0.233	-0.579	<0.01
根 Root	革兰氏阴性菌 G-	0.000	-0.122	-0.122	<0.01
	真菌 Fungi	0.162	0.000	0.162	<0.01
	植物来源微生物生物量碳 ¹³C-MBC	-0.483	-0.344	-0.827	<0.01
	纤维素酶 Cellobiohydrolase	-0.413	0.000	-0.413	<0.01
	土壤有效氮 Soil available nitrogen	0.248	0.000	0.248	<0.01

附录I 亚热带森林实验土壤(0-20 cm)基本理化性质

Supplement I Basic properties of the study soil in a subtropical forest (0-20 cm)

土壤基本理化性质 Basic properties of the study soil	平均值 Mean
土壤有机碳含量 SOC content (g·kg^-1)	12.30
总氮含量 TN content (g·kg^-1)	2.07
硝态氮含量 NO₃^--N content (mg N·kg^-1)	0.79
铵态氮含量 NH₄⁺-N content (mg N·kg^-1)	11.56
微生物生物量碳含量 MBC content (mg·kg^-1)	330.86
微生物生物量氮含量 MBN content (mg·kg^-1)	55.93
微生物碳/微生物氮 MBC:MBN	5.92
¹³C (‰)	-23.39

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亚热带森林不同植物及器官来源的可溶性有机质输入对土壤激发效应的影响及其作用机理

Effects of dissolved organic matter derived from different plant and tissues in a subtropical forest on soil priming effect and the underlying mechanisms

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