川西亚高山不同演替阶段天然次生林土壤碳氮含量及酶活性特征

doi:10.17521/cjpe.2020.0203

植物生态学报 ›› 2020, Vol. 44 ›› Issue (9): 973-985.DOI: 10.17521/cjpe.2020.0203

所属专题：碳储量

川西亚高山不同演替阶段天然次生林土壤碳氮含量及酶活性特征

胡宗达¹^,^*(), 刘世荣³, 罗明霞¹, 胡璟⁴, 刘兴良², 李亚非¹, 余昊¹, 欧定华¹

¹四川农业大学资源学院, 成都 611130
²四川省林业科学研究院, 森林和湿地生态恢复与保育四川重点实验室, 成都 610081
³中国林业科学研究院森林生态环境与保护研究所, 北京 100091
⁴湖南农业大学资源环境学院, 长沙 410128

收稿日期:2020-06-19 修回日期:2020-08-06 出版日期:2020-09-20 发布日期:2020-09-03
通讯作者: * 胡宗达 E-mail: (huzd98@163.com )
基金资助:
森林和湿地生态恢复与保育四川重点实验室开放课题(2019KFKT03);国家重点研发计划(2017YFC0505004);四川省环境治理与生态保护重大科技专项(2018SZDZX0031)

Characteristics of soil carbon and nitrogen contents and enzyme activities in sub-alpine secondary forests with different successional stages in Western Sichuan, China

HU Zong-Da¹^,^*(), LIU Shi-Rong³, LUO Ming-Xia¹, HU Jing⁴, LIU Xing-Liang², LI Ya-Fei¹, YU Hao¹, OU Ding-Hua¹

¹College of Resources, Sichuan Agricultural University, Chengdu 611130, China
²Ecological Restoration and Conservation on Forest and Wetland Key Laboratory of Sichuan Province, Sichuan Academy of Forestry, Chengdu 610081, China
³Research Institute of Forest Ecology, Environment and Protection, Chinese Academy of Forestry, Beijing 100091, China
⁴College of Resources and Environment, Hunan Agricultural University, Changsha 410128, China

Received:2020-06-19 Revised:2020-08-06 Online:2020-09-20 Published:2020-09-03
Contact: HU Zong-Da
Supported by:
Supported by Forest and Wetland Ecological Restoration and Conservation of Open Project from Key Laboratory of Sichuan Province(2019KFKT03);the National Key R&D Program of China(2017YFC0505004);the Environmental Governance and Ecological Protection Major Project of Sichuan Province(2018SZDZX0031)

摘要/Abstract

摘要：

为了解次生林自然更新演替过程中土壤碳氮含量及酶活性的变化规律, 采用空间代替时间的方法, 在川西亚高山米亚罗林区选取环境条件基本一致的20世纪60、70和80年代采伐迹地自然更新演替的次生林(60-NSF、70-NSF和80-NSF)和岷江冷杉(Abies faxoniana)原始林(对照, CK)为对象, 研究了表层(0-20 cm)土壤碳氮含量和土壤酶活性的关系。结果表明: 表层土壤有机碳(SOC)、可溶性有机碳(DOC)、轻组有机碳(LFOC)含量均随森林植被更新演替呈显著降低趋势, 而全氮(TN)和可溶性有机氮(DON)含量则表现为60-NSF < 80-NSF < 70-NSF, 但70-NSF和80-NSF间差异不显著; 次生林表层土壤有机碳氮及其活性组分含量均低于CK, 其中80-NSF的DOC和DON含量与CK差异不显著。次生林的β-葡萄糖苷酶(βG)、β-N-乙酰氨基葡萄糖苷酶(NAG)和多酚氧化酶(PHO)活性均显著低于CK, 纤维素水解酶(CBH)和过氧化物酶(PEO)活性与CK无显著差异; 天然次生林中, 60-NSF的βG和CBH活性显著低于70-NSF和80-NSF; 80-NSF的NAG活性显著高于60-NSF和70-NSF; 4种林型之间的PEO活性无显著性差异。Pearson相关分析和冗余分析显示, 土壤TN、LFOC和DOC含量与土壤酶活性显著相关, 其中TN含量解释了酶活性变化的65.4%, 说明土壤氮含量变化可能会影响到土壤碳的水解酶活性, 同时也表明土壤微生物优先利用易分解碳和氮。因此, 次生林近60年的天然更新演替引起了TN、LFOC及DOC含量的显著下降, 导致表层土壤某些胞外酶(如βG、CBH和NAG)活性降低。从土壤酶活性角度看, 岷江冷杉原始林比早期演替阶段的次生林(<60 a)更有利于川西亚高山高海拔森林生态系统的碳氮循环。

关键词: 土壤有机碳, 土壤全氮, 酶活性, 演替阶段, 天然次生林, 土壤碳循环, 土壤氮循环

Abstract:

Aims Regeneration of sub-alpine forests have an capacity to sequester carbon and nitrogen. Our objectives were to quantify variations of soil organic carbon and nitrogen content and enzyme activities at different successional stages of natural secondary forests, and to better understand the underlying mechanisms of carbon and nitrogen sequestration in these sub-alpine forests.
Methods We used the space-for-time substitution method and selected four sub-alpine forests in Miyaluo forest of Western Sichuan, China. The secondary forests were at different successional stages with natural regeneration on cutting-blanks in 1960s (60-NSF), 1970s (70-NSF) and 1980s (80-NSF), and the Abies faxoniana primary forest was used as control (CK). The soil samples were taken from 0 to 20 cm depths in each forest in late July, 2019, and were transferred to the laboratory. Soil organic carbon (SOC), soil total nitrogen (TN), dissolved organic carbon (DOC) and nitrogen (DON), light fraction organic carbon (LFOC) content and soil enzyme activities were measured. The activities of five soil extracellular enzymes related to soil carbon and nitrogen cycling were determined to explain their relationships with soil physico-chemical properties.
Important findings The contents of topsoil SOC, DOC, LFOC decreased with succession stage, whereas TN and DON were all in the order of 60-NSF < 80-NSF < 70-NSF, although 80-NSF and 70-NSF exhibited no significant difference. The topsoil organic carbon and nitrogen and their active fractions contents in natural secondary forests were lower than those in the primary forest, while no significant difference in DOC and DON contents was observed between 80-NSF and CK. The activities of soil β-4-glucosidase (βG), β-4-N-acetylglucosa- minidase (NAG) and polyphenol oxidase (PHO) in CK were significantly higher than those in natural secondary forests, whereas soil cellulose hydrolysis (CBH) and phenol oxidase (PEO) activities had no significant difference between secondary forests and CK. Activities of βG and CBH in 60-NSF were significantly lower than those in 70-NSF and 80-NSF, but activities of NAG in 80-NSF were significantly higher than those in 60-NSFand 70-NSF. There was no significant difference in PEO activities among different types of forest. Both Pearson correlation analysis and redundancy analysis showed that soil enzyme activities were significantly correlated with soil TN, LFOC and DOC contents. TN content explained 65.4% of the variations in enzyme activity, which implied that change in soil nitrogen content might affect C-related hydrolytic enzyme activities (e.g. βG, CBH and NAG). Meanwhile soil microorganisms prefer to use readily decomposable carbon and nitrogen. Therefore, activities of some soil enzymes such as βG, CBH and NAG in natural secondary forests decreased due to the declines in soil TN, LFOC and DOC contents. We conclude that soil enzyme activities could be more favorable to C and N cycling in the Abies faxoniana primary forest than in the secondary forest at the early-successional stages (<60 a) in high-altitude sub-alpine forest ecosystems in Western Sichuan, China.

Key words: soil organic carbon, soil total nitrogen, enzyme activity, succession stage, natural secondary forest, soil carbon cycle, soil nitrogen cycle

胡宗达, 刘世荣, 罗明霞, 胡璟, 刘兴良, 李亚非, 余昊, 欧定华. 川西亚高山不同演替阶段天然次生林土壤碳氮含量及酶活性特征. 植物生态学报, 2020, 44(9): 973-985. DOI: 10.17521/cjpe.2020.0203

HU Zong-Da, LIU Shi-Rong, LUO Ming-Xia, HU Jing, LIU Xing-Liang, LI Ya-Fei, YU Hao, OU Ding-Hua. Characteristics of soil carbon and nitrogen contents and enzyme activities in sub-alpine secondary forests with different successional stages in Western Sichuan, China. Chinese Journal of Plant Ecology, 2020, 44(9): 973-985. DOI: 10.17521/cjpe.2020.0203

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林型 Forest type	立地特征 Site characteristic				林型特征 Stand characteristic
林型 Forest type	平均海拔 Mean altitude (m)	坡向 Slope aspect	平均坡度 Mean slope (°)	坡位 Slope position	林分密度 Stand density (tree·hm^-2)	平均树高 Mean height (m)	平均胸径 Mean DBH (cm)	郁闭度 Canopy density	草本层盖度 Coverage of herb (%)
80-NSF	3 387	东北 NE	22	中坡 Middle slope	2 375	6.35	10.83	0.60	65.2
70-NSF	3 312	西北 NW	27	中坡 Middle slope	1 833	7.25	11.12	0.62	79.3
60-NSF	3 080	东北 NE	24	中坡 Middle slope	1 583	6.83	14.17	0.68	40.5
CK	3 189	东北 NE	25	中上坡 Upper-middle slope	1 633	10.56	26.70	0.76	43.3

林型 Forest type	立地特征 Site characteristic				林型特征 Stand characteristic
林型 Forest type	平均海拔 Mean altitude (m)	坡向 Slope aspect	平均坡度 Mean slope (°)	坡位 Slope position	林分密度 Stand density (tree·hm^-2)	平均树高 Mean height (m)	平均胸径 Mean DBH (cm)	郁闭度 Canopy density	草本层盖度 Coverage of herb (%)
80-NSF	3 387	东北 NE	22	中坡 Middle slope	2 375	6.35	10.83	0.60	65.2
70-NSF	3 312	西北 NW	27	中坡 Middle slope	1 833	7.25	11.12	0.62	79.3
60-NSF	3 080	东北 NE	24	中坡 Middle slope	1 583	6.83	14.17	0.68	40.5
CK	3 189	东北 NE	25	中上坡 Upper-middle slope	1 633	10.56	26.70	0.76	43.3

酶 Enzyme	缩写 Abbreviation	功能 Function	底物 Substrate
β-4-葡萄糖苷酶 β-4-glucosidase	βG	分解易降解碳 Decomposition of labile carbon	4-MUB-β-D-葡萄糖苷 4- MUB -β-D-glucoside
纤维素水解酶 Cellulose hydrolysis	CBH	分解易降解碳 Decomposition of labile carbon	羧甲基纤维素钠 Sodium carboxymethyl cellulose
β-4-N-乙酰基氨基葡萄糖苷酶 β-4-N-acetylglucosaminidase	NAG	分解氮 Hydrolyze nitrogen	4-MUB-β-D-乙酰基氨基葡萄糖苷 4-MUB-N-acetyl-β-D-glucosaminide
多酚氧化酶 Phenol oxidase	PHO	分解难降解碳 Decomposition of recalcitrant carbon	邻苯三酚 Pyrogallic acid
过氧化物酶 Peroxidase	PEO	分解难降解碳 Decomposition of recalcitrant carbon	邻苯三酚-过氧化氢 Pyrogallol-hydrogen peroxide

酶 Enzyme	缩写 Abbreviation	功能 Function	底物 Substrate
β-4-葡萄糖苷酶 β-4-glucosidase	βG	分解易降解碳 Decomposition of labile carbon	4-MUB-β-D-葡萄糖苷 4- MUB -β-D-glucoside
纤维素水解酶 Cellulose hydrolysis	CBH	分解易降解碳 Decomposition of labile carbon	羧甲基纤维素钠 Sodium carboxymethyl cellulose
β-4-N-乙酰基氨基葡萄糖苷酶 β-4-N-acetylglucosaminidase	NAG	分解氮 Hydrolyze nitrogen	4-MUB-β-D-乙酰基氨基葡萄糖苷 4-MUB-N-acetyl-β-D-glucosaminide
多酚氧化酶 Phenol oxidase	PHO	分解难降解碳 Decomposition of recalcitrant carbon	邻苯三酚 Pyrogallic acid
过氧化物酶 Peroxidase	PEO	分解难降解碳 Decomposition of recalcitrant carbon	邻苯三酚-过氧化氢 Pyrogallol-hydrogen peroxide

项目 Item	80-NSF	70-NSF	60-NSF	CK
pH	5.08 ± 0.06^a	5.24 ± 0.16^ab	5.42 ± 0.11^bc	5.54 ± 0.04^c
含水率 Soil water content (%)	57.65 ± 14.58^a	48.68 ± 2.99^a	23.73 ± 4.50^b	68.47 ± 9.99^a
有机碳含量 Soil organic carbon content (g·kg^-1)	98.07 ± 4.08^a	91.28 ± 5.90^a	47.29 ± 1.88^b	124.38 ± 14.70^c
全氮含量 Soil total nitrogen content (g·kg^-1)	8.22 ± 0.57^a	8.90 ± 0.60^a	3.28 ± 0.15^b	11.70 ± 1.03^c
碳氮比 C:N	11.95 ± 0.47^a	10.25 ± 0.03^b	14.40 ± 0.28^c	10.61 ± 0.35^b
全磷含量 Soil total phosphorus content (g·kg^-1)	0.78 ± 0.08^a	1.12 ± 0.14b^c	0.93 ± 0.02^ad	1.06 ± 0.05^cd
全钾含量 Soil total potassium content (g·kg^-1)	16.18 ± 0.47^a	21.69 ± 0.48^b	27.26 ± 0.87^c	23.86 ± 0.23^d
可溶性有机碳含量 Dissolved organic carbon content (mg·kg^-1)	123.57 ± 24.89^a	66.08 ± 5.52^b	45.19 ± 9.09^b	139.01 ± 38.74^a
轻组有机碳含量 Light fraction organic carbon content (g·kg^-1)	25.80 ± 9.00^a	20.85 ± 4.75^ab	14.02 ± 1.09^b	40.26 ± 5.99^c
可溶性有机氮含量 Dissolved organic nitrogen content (mg·kg^-1)	40.84 ± 5.35^a	48.29 ± 7.21^a	18.83 ± 5.64^b	39.03 ± 6.33^a

川西亚高山不同演替阶段天然次生林土壤碳氮含量及酶活性特征

Characteristics of soil carbon and nitrogen contents and enzyme activities in sub-alpine secondary forests with different successional stages in Western Sichuan, China

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林型 Forest type	βG (nmol×g^-1×h^-1)	CBH (mg×g^-1×h^-1)	NAG (nmol×g^-1×h^-1)	PHO (mg×g^-1×h^-1)	PEO (mg×g^-1×h^-1)
80-NSF	22.64 ± 3.83^a	26.15 ± 2.49^a	9.76 ± 0.82^a	38.83 ± 1.59^a	36.19 ± 2.61^a
70-NSF	22.30 ± 1.23^a	25.66 ± 1.88^a	4.01 ± 0.36^b	35.09 ± 1.56^a	31.24 ± 2.91^a
60-NSF	6.20 ± 1.69^b	18.53 ± 0.95^b	2.37 ± 0.05^b	34.82 ± 1.04^a	36.69 ± 1.18^a
CK	31.54 ± 5.51^c	22.60 ± 1.03^ab	12.40 ± 0.56^c	46.02 ± 1.60^b	37.81 ± 1.54^a

项目 Item	DOC 含量 DOC content	DON 含量 DON content	SOC 含量 SOC content	TN含量 TN content	TP含量 TP content	TK含量 TK content	C:N	pH	土壤含水率 SWC	H	D	J	E
轻组有机碳含量 LFOC content	0.853^**	0.431	0.833^**	0.805^**	0.253	-0.189	-0.564	0.354	0.810^**	0.085	0.137	0.204	0.208
可溶性有机碳含量 DOC content	1	0.510	0.857^**	0.770^**	-0.079	-0.479	-0.456	-0.029	0.869^**	0.235	0.242	0.248	0.240
可溶性有机氮含量 DON content		1	0.712^**	0.754^**	0.311	-0.596^*	-0.837^**	-0.451	0.659^*	0.683^*	0.594^*	0.148	0.369
有机碳含量 SOC content			1	0.983^**	0.194	-0.450	-0.799^**	0.017	0.860^**	0.332	0.302	0.111	0.295
全氮含量 TN content				1	0.329	-0.392	-0.885^**	0.061	0.847^**	0.386	0.334	0.052	0.342
全磷含量 TP content					1	0.444	-0.486	0.408	0.157	0.420	0.282	-0.203	0.632^*
全钾含量 TK content						1	0.425	0.744^**	-0.483	-0.347	-0.390	-0.306	0.212
碳氮比 C:N							1	0.060	-0.710^**	-0.519	-0.431	0.040	-0.307
土壤pH Soil pH								1	0.013	-0.421	-0.410	-0.247	0.082
土壤含水率 SWC									1	0.340	0.280	0.051	0.222

项目 Item	β-4-葡萄糖苷酶 β-4-glucosidase	纤维素水解酶 Cellulose hydrolysis	β-4-N-乙酰基氨基葡萄糖苷酶 β-4-N-acetylglucosaminidase	多酚氧化酶 Phenol oxidase	过氧化物酶 Peroxidase
土壤pH Soil pH	0.010	-0.583^*	0.086	0.403	0.352
土壤含水率 SWC	0.804^**	0.498	0.809^**	0.749^**	0.298
全氮含量 TN content	0.939^**	0.474	0.787^**	0.715^**	-0.016
有机碳含量 SOC content	0.916^**	0.446	0.853^**	0.760^**	0.050
碳氮比 C:N	-0.867^**	-0.621^*	-0.508	-0.421	0.227
全磷含量 TP content	0.244	0.132	-0.124	0.237	-0.057
全钾含量 TK content	-0.456	-0.694^*	-0.447	-0.087	0.094
轻组有机碳含量 LFOC content	0.737^**	0.295	0.844^**	0.895^**	0.493
可溶性有机碳含量 DOC content	0.722^**	0.374	0.908^**	0.832^**	0.422
可溶性有机氮含量 DON content	0.682^*	0.814^**	0.388	0.222	-0.287
Shannon-Wiener指数 H	0.407	0.610^*	0.091	0.120	-0.243
Simpson指数 D	0.343	0.616^*	0.151	0.138	-0.224
Pielou均匀度指数 J	0.022	0.320	0.222	0.129	0.035
Margalef丰富度指数 E	0.330	0.119	0.074	0.317	0.047

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