川西亚高山森林凋落物去除对土壤腐殖质动态的影响

doi:10.17521/cjpe.2021.0166

植物生态学报 ›› 2022, Vol. 46 ›› Issue (3): 330-339.DOI: 10.17521/cjpe.2021.0166

所属专题：凋落物

川西亚高山森林凋落物去除对土壤腐殖质动态的影响

刘谣, 焦泽彬, 谭波, 李晗, 王丽霞, 刘思凝, 游成铭, 徐振锋, 张丽^*()

长江上游林业生态工程四川省重点实验室, 长江上游森林资源保育与生态安全国家林业和草原局重点实验室, 高山森林生态系统定位研究站, 四川农业大学生态林业研究所, 成都 611130

收稿日期:2021-04-30 接受日期:2021-09-01 出版日期:2022-03-20 发布日期:2021-10-15
通讯作者: 张丽
作者简介:* (zhangli19830116@hotmail.com)
基金资助:
国家自然科学基金(31901295);国家自然科学基金(31800519);国家自然科学基金(32001165);国家自然科学基金(31700542);国家重点研发计划(2016YFC0502505);国家重点研发计划(2017YFC0505003);四川省杰出青年科技人才计划项目(2020JDJQ0052)

Litter removal effects on dynamics of soil humic substances in subalpine forests of western Sichuan, China

LIU Yao, JIAO Ze-Bin, TAN Bo, LI Han, WANG Li-Xia, LIU Si-Ning, YOU Cheng-Ming, XU Zhen-Feng, ZHANG Li^*()

Forestry Ecological Engineering in the Upper Reaches of Yangtze River Key Laboratory of Sichuan Province, National Forestry and Grassland Administration Key Laboratory of Forest Resources Conservation and Ecological Safety on the Upper Reaches of the Yangtze River, Long-term Research Station of Alpine Forest Ecosystems, Institute of Ecology & Forestry, Sichuan Agricultural University, Chengdu 611130, China

Received:2021-04-30 Accepted:2021-09-01 Online:2022-03-20 Published:2021-10-15
Contact: ZHANG Li
Supported by:
National Natural Science Foundation of China(31901295);National Natural Science Foundation of China(31800519);National Natural Science Foundation of China(32001165);National Natural Science Foundation of China(31700542);National Key R&D Program of China(2016YFC0502505);National Key R&D Program of China(2017YFC0505003);Program of Sichuan Excellent Youth Sci-Tech Foundation(2020JDJQ0052)

摘要/Abstract

摘要：

森林凋落物作为森林土壤腐殖质的主要来源, 在土壤腐殖质的形成中发挥着重要作用, 但不同森林类型凋落物因其含量、组成等的不同, 对土壤腐殖质的影响也不同。该研究以川西亚高山针叶林、阔叶林和针阔混交林3种不同森林类型为对象, 采用凋落物原位控制实验, 对比研究不同关键期凋落物去除对土壤可提取腐殖质、胡敏酸和富里酸含量及胡敏酸/富里酸、胡敏酸/可提取腐殖质的影响。主要结果: (1)土壤可提取腐殖质、胡敏酸和富里酸含量在不同森林类型中差异显著。土壤可提取腐殖质含量总体表现为针叶林>针阔混交林>阔叶林, 胡敏酸含量总体表现为针阔混交林>针叶林>阔叶林, 而富里酸含量则表现为针叶林>阔叶林>针阔混交林, 其中3种林型中土壤腐殖质的主要成分为富里酸, 总体均表现为富里酸型。不同采样时期也显著影响了土壤可提取腐殖质、胡敏酸和富里酸含量, 总体均表现为先升高后下降的趋势。除个别采样时期外, 凋落物去除总体降低了土壤可提取腐殖质、胡敏酸和富里酸的含量。(2)胡敏酸/富里酸和胡敏酸/可提取腐殖质的结果显示3种林型土壤总体腐殖化程度均较低, 整体表现为针阔混交林>阔叶林>针叶林, 凋落物去除在一定程度上有利于提高阔叶林与针阔混交林的腐殖质品质。(3)相关分析表明不同凋落物处理间土壤可提取腐殖质与土壤有机碳含量、全氮含量和土壤含水量呈显著正相关关系, 与温度呈显著负相关关系。综上所述, 短期的凋落物去除会降低土壤腐殖物质的含量, 但不同林型间由于凋落物类型差异会导致土壤腐殖质的不同变化, 说明土壤腐殖质的动态变化受凋落物类型以及环境因素的综合调控。因此, 关于凋落物变化对土壤腐殖质的影响还需进一步的长期研究。

关键词: 腐殖质, 胡敏酸, 富里酸, 凋落物, 亚高山森林

Abstract:

Aims Forest litter are the main sources of soil humic substances, and different litter types can have differential effects on the formation and composition of soil humic substances. The aim of this study was to determine how variations in litter input would affect the dynamics of soil humic substances in subalpine forests of western Sichuan.
Methods A field in-situ litter manipulation experiment incorporating litter removal was established on sites of coniferous, broad-leaved, and mixed coniferous and broad-leaved forest stands. Measurements were made on the contents of soil extractable humic substances, humic acid, and fulvic acid; ratios of humic acid to fulvic acid and humic acid to humic substances were also computed.
Important findings The contents of extractable humic substances, humic acid and fulvic acid of soils varied significantly among forest types. The extractable humic substances content followed a descending order of coniferous forest > mixed coniferous and broad-leaved forest > broad-leaved forest, and the humic acid content of mixed coniferous and broad-leaved forest > coniferous forest > broad-leaved forest, and the fulvic acid content of coniferous forest > broad-leaved forest > mixed coniferous and broad-leaved forest. Fulvic acid was the predominant fraction of soil humic substances in all the three forest types. Time of measurements also significantly affected the contents of extractable humic substances, humic acid and fulvic acid, with an increasing trend for up to 1.5-year followed by decreases thereafter. With few exceptions, litter removal generally reduced the contents of extractable humic substances, humic acid and fulvic acid. The ratios of humic acid/fulvic acid and humic acid/extractable humic substances indicate low degree of humification for all the three forest types, which followed a descending order of mixed coniferous and broad-leaved forest > broad-leaved forest > coniferous forest. Litter removal improved the humic substances quality in the broad-leaved and mixed coniferous and broad-leaved forests to some extent. Correlation analysis showed significantly positive correlations of soil extractable humic substances with soil organic carbon, total nitrogen and soil water content, and a significantly negative correlation with temperature. In summary, short-term litter removal reduced the soil humic substances content, with differential effects by different litter types. We draw conclusion that the dynamics of soil humic substances are comprehensively regulated by litter type and environmental factors. Therefore, the impact of litter changes on soil humic substances needs further long-term research.

Key words: humic substances, humic acid, fulvic acid, litter, subalpine forest

刘谣, 焦泽彬, 谭波, 李晗, 王丽霞, 刘思凝, 游成铭, 徐振锋, 张丽. 川西亚高山森林凋落物去除对土壤腐殖质动态的影响. 植物生态学报, 2022, 46(3): 330-339. DOI: 10.17521/cjpe.2021.0166

LIU Yao, JIAO Ze-Bin, TAN Bo, LI Han, WANG Li-Xia, LIU Si-Ning, YOU Cheng-Ming, XU Zhen-Feng, ZHANG Li. Litter removal effects on dynamics of soil humic substances in subalpine forests of western Sichuan, China. Chinese Journal of Plant Ecology, 2022, 46(3): 330-339. DOI: 10.17521/cjpe.2021.0166

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0166

https://www.plant-ecology.com/CN/Y2022/V46/I3/330

图/表 7

表1 川西亚高山森林样地主要信息

Table 1 Basic information on sampling plots in subalpine forests of western Sichuan

森林类型 Forest type	海拔 Altitude (m)	优势乔木 Dominant tree	主要林下植物 Major understory species
针叶林 Coniferous forest	3 000	岷江冷杉 Abies fargesii var. faxoniana	茜草 Rubia cordifolia 高山冷蕨 Cystopteris montana 高山杜鹃 Rhododendron lapponicum
针阔混交林 Mixed coniferous and broad-leaved forest	2 985	岷江冷杉 Abies fargesii var. faxoniana 红桦 Betula albosinensis	高山杜鹃 Rhododendron lapponicum 箭竹 Fargesia spathacea 高山冷蕨 Cystopteris montana
阔叶林 Broad-leaved forest	3 010	红桦 Betula albosinensis	箭竹 Fargesia spathacea 蟹甲草 Parasenecio forrestii 高山冷蕨 Cystopteris montana

表2 凋落物处理下不同采样时期川西亚高山3种森林类型土壤基本理化指标(平均值±标准误)

Table 2 Soil physical and chemical variables at different measurement times for the three forest types under contrasting litter treatments in subalpine forests of western Sichuan (mean ± SE)

时间 Time (a)	指标 Index	土壤含水量 Soil water content (%)	温度 Temperature (°C)	pH	有机碳含量 Organic carbon content (g·kg^-1)	全氮含量 Total nitrogen content (g·kg^-1)	全磷含量 Total phosphorus content (g·kg^-1)
1	BFI	54.94 ± 3.41^a	10.56 ± 0.05^a	5.82 ± 0.14^a	161.70 ± 25.4^a	0.25 ± 0.08^a	0.89 ± 0.07^a
	MFI	41.75 ± 9.09^a	10.41 ± 0.30^a	5.08 ± 0.11^b	121.83 ± 21.55^a	0.17 ± 0.07^a	0.87 ± 0.05^a
	CFI	55.61 ± 2.35^a	10.15 ± 0.06^a	4.76 ± 0.07^b	171.80 ± 20.08^a	0.28 ± 0.03^a	0.83 ± 0.05^a
	BFR	45.86 ± 7.70^a	10.45 ± 0.04^a	5.48 ± 0.06^a	96.46 ± 43.51^a	0.21 ± 0.06^a	0.86 ± 0.10^a
	MFR	53.82 ± 7.35^a	10.14 ± 0.15^a	5.50 ± 0.09^a	192.46 ± 44.59^a	0.30 ± 0.13^a	0.76 ± 0.06^a
	CFR	54.96 ± 1.50^a	9.75 ± 0.06^a	5.10 ± 0.25^a	166.63 ± 12.94^a	0.27 ± 0.06^a	0.78 ± 0.04^a
1.5	BFI	57.57 ± 2.10^a	5.00 ± 0.14^a	6.22 ± 0.14^a	186.73 ± 12.12^b	13.10 ± 1.59^a	0.76 ± 0.14^a
	MFI	57.59 ± 7.96^a	4.75 ± 0.11^a	5.37 ± 0.25^a	240.58 ± 18.48^a	11.36 ± 3.33^a	0.79 ± 0.05^a
	CFI	50.79 ± 1.88^a	4.65 ± 0.07^a	5.68 ± 0.45^a	211.02 ± 6.00^ab	11.10 ± 2.62^a	0.63 ± 0.13^a
	BFR	50.32 ± 3.02^a	4.70 ± 0.06^a	5.97 ± 0.35^a	116.24 ± 20.35^a	10.36 ± 0.84^a	0.74 ± 0.10^a
	MFR	53.32 ± 5.71^a	4.65 ± 0.07^a	5.54 ± 0.27^a	177.99 ± 51.16^a	10.37 ± 3.13^a	0.83 ± 0.08^a
	CFR	52.90 ± 4.42^a	4.56 ± 0.25^a	5.69 ± 0.24^a	228.29 ± 16.32^a	14.07 ± 1.11^a	0.71 ± 0.08^a
2	BFI	53.40 ± 8.82^a	10.24 ± 0.21^a	5.86 ± 0.21^a	121.61 ± 41.60^a	8.18 ± 2.54^a	0.47 ± 0.11^a
	MFI	58.46 ± 5.45^a	9.13 ± 0.05^a	5.81 ± 0.05^a	156.81 ± 31.94^a	7.75 ± 1.55^a	0.41 ± 0.06^a
	CFI	46.65 ± 3.35^a	9.09 ± 0.05^a	5.79 ± 0.05^a	99.19 ± 5.10^a	2.60 ± 1.19^a	0.22 ± 0.04^a
	BFR	46.29 ± 3.96^a	9.86 ± 0.05^a	5.74 ± 0.05^a	84.60 ± 18.86^a	3.29 ± 1.09^ab	0.44 ± 0.05^a
	MFR	48.75 ± 6.25^a	8.90 ± 0.12^a	5.91 ± 0.12^a	102.31 ± 21.54^a	5.25 ± 0.71^a	0.36 ± 0.04^ab
	CFR	43.88 ± 0.75^a	9.10 ± 0.14^a	5.62 ± 0.14^a	106.16 ± 10.74^a	0.82 ± 0.13^b	0.26 ± 0.02^b

表2 凋落物处理下不同采样时期川西亚高山3种森林类型土壤基本理化指标(平均值±标准误)

Table 2 Soil physical and chemical variables at different measurement times for the three forest types under contrasting litter treatments in subalpine forests of western Sichuan (mean ± SE)

时间 Time (a)	指标 Index	土壤含水量 Soil water content (%)	温度 Temperature (°C)	pH	有机碳含量 Organic carbon content (g·kg^-1)	全氮含量 Total nitrogen content (g·kg^-1)	全磷含量 Total phosphorus content (g·kg^-1)
1	BFI	54.94 ± 3.41^a	10.56 ± 0.05^a	5.82 ± 0.14^a	161.70 ± 25.4^a	0.25 ± 0.08^a	0.89 ± 0.07^a
	MFI	41.75 ± 9.09^a	10.41 ± 0.30^a	5.08 ± 0.11^b	121.83 ± 21.55^a	0.17 ± 0.07^a	0.87 ± 0.05^a
	CFI	55.61 ± 2.35^a	10.15 ± 0.06^a	4.76 ± 0.07^b	171.80 ± 20.08^a	0.28 ± 0.03^a	0.83 ± 0.05^a
	BFR	45.86 ± 7.70^a	10.45 ± 0.04^a	5.48 ± 0.06^a	96.46 ± 43.51^a	0.21 ± 0.06^a	0.86 ± 0.10^a
	MFR	53.82 ± 7.35^a	10.14 ± 0.15^a	5.50 ± 0.09^a	192.46 ± 44.59^a	0.30 ± 0.13^a	0.76 ± 0.06^a
	CFR	54.96 ± 1.50^a	9.75 ± 0.06^a	5.10 ± 0.25^a	166.63 ± 12.94^a	0.27 ± 0.06^a	0.78 ± 0.04^a
1.5	BFI	57.57 ± 2.10^a	5.00 ± 0.14^a	6.22 ± 0.14^a	186.73 ± 12.12^b	13.10 ± 1.59^a	0.76 ± 0.14^a
	MFI	57.59 ± 7.96^a	4.75 ± 0.11^a	5.37 ± 0.25^a	240.58 ± 18.48^a	11.36 ± 3.33^a	0.79 ± 0.05^a
	CFI	50.79 ± 1.88^a	4.65 ± 0.07^a	5.68 ± 0.45^a	211.02 ± 6.00^ab	11.10 ± 2.62^a	0.63 ± 0.13^a
	BFR	50.32 ± 3.02^a	4.70 ± 0.06^a	5.97 ± 0.35^a	116.24 ± 20.35^a	10.36 ± 0.84^a	0.74 ± 0.10^a
	MFR	53.32 ± 5.71^a	4.65 ± 0.07^a	5.54 ± 0.27^a	177.99 ± 51.16^a	10.37 ± 3.13^a	0.83 ± 0.08^a
	CFR	52.90 ± 4.42^a	4.56 ± 0.25^a	5.69 ± 0.24^a	228.29 ± 16.32^a	14.07 ± 1.11^a	0.71 ± 0.08^a
2	BFI	53.40 ± 8.82^a	10.24 ± 0.21^a	5.86 ± 0.21^a	121.61 ± 41.60^a	8.18 ± 2.54^a	0.47 ± 0.11^a
	MFI	58.46 ± 5.45^a	9.13 ± 0.05^a	5.81 ± 0.05^a	156.81 ± 31.94^a	7.75 ± 1.55^a	0.41 ± 0.06^a
	CFI	46.65 ± 3.35^a	9.09 ± 0.05^a	5.79 ± 0.05^a	99.19 ± 5.10^a	2.60 ± 1.19^a	0.22 ± 0.04^a
	BFR	46.29 ± 3.96^a	9.86 ± 0.05^a	5.74 ± 0.05^a	84.60 ± 18.86^a	3.29 ± 1.09^ab	0.44 ± 0.05^a
	MFR	48.75 ± 6.25^a	8.90 ± 0.12^a	5.91 ± 0.12^a	102.31 ± 21.54^a	5.25 ± 0.71^a	0.36 ± 0.04^ab
	CFR	43.88 ± 0.75^a	9.10 ± 0.14^a	5.62 ± 0.14^a	106.16 ± 10.74^a	0.82 ± 0.13^b	0.26 ± 0.02^b

图1 川西亚高山3种林型凋落物自然输入与去除土壤的可提取腐殖质、胡敏酸和富里酸含量(平均值±标准误)。不同小写字母表示同一林型相同处理下不同采样时间土壤可提取腐殖质、胡敏酸和富里酸含量差异显著(p < 0.05)。*, 同一采样时间凋落物自然输入与凋落物去除之间差异显著(p < 0.05)。

Fig. 1 Soil extractable humic substances content, humic acid content and fulvic acid content as affected by litter input in broad-leaved forest, mixed coniferous and broad-leaved forest and coniferous forest in subalpine forests of western Sichuan (mean ± SE). Different lowercase letters indicate that the differences of extractable humic substances content, humic acid content and fulvic acid content between different measurement times within the same treatment are significant (p < 0.05). *, significant differences in soil humic substances content between litter treatments in the same measurement time (p < 0.05).

表3 凋落物、培养时间和林型对川西亚高山森林土壤可提取腐殖质、胡敏酸、富里酸含量、胡敏酸/富里酸及腐殖化度的三因素方差分析(F值)

Table 3 Results of three way ANOVA (F value) for testing the effects of litter, measurement time and forest type on soil extractable humic substances (EHS), humic acid (HA), fulvic acid (FA) content, and the ratios of humic acid/fulvic acid (HA/FA) and humic acid/extractable humic substances (PQ) in subalpine forests of western Sichuan

因子 Factor	可提取腐殖质 EHS	胡敏酸 HA	富里酸 FA	胡敏酸/富里酸 HA/FA	胡敏酸/可提取腐殖质 PQ
凋落物 Litter	11.571^**	0.827	5.659^*	2.745	0.098
培养时间 Time	73.835^**	28.137^**	38.713^**	10.653^**	4.263^*
林型 Type	10.808^**	31.479^**	15.783^**	5.963^**	19.403^**
凋落物×时间 Litter × Time	15.654^**	19.443^**	8.544^**	2.142	2.054
凋落物×林型 Litter × Type	3.328	9.650^**	3.853^*	5.657^*	10.340^**
时间×林型 Time × Type	8.584^**	8.011^**	12.489^**	5.838^**	7.266^**
凋落物×时间×林型 Litter × Time × Type	1.111	5.777^**	4.080^**	4.936^**	8.496^**

图2 川西亚高山3种林型凋落物自然输入与去除土壤的胡敏酸/富里酸(平均值±标准误)。不同小写字母表示同一林型相同处理下不同采样时间土壤胡敏酸/富里酸差异显著(p < 0.05)。*, 同一采样时间凋落物自然输入与凋落物去除之间差异显著(p < 0.05)。

Fig. 2 Humic acid/fulvic acid ratio as affected by litter input in broad-leaved forest, mixed coniferous and broad-leaved forest and coniferous forest in subalpine forests of western Sichuan (mean ± SE). Different lowercase letters indicate that the differences in humic acid/fulvic acid ratio between measurement times under given litter treatments are significant (p < 0.05). *, significant differences in humic acid/fulvic acid ratio between litter treatments in the same measurement time (p < 0.05).

图3 川西亚高山3种林型凋落物自然输入与去除土壤的胡敏酸/可提取腐殖质(平均值±标准误)。不同小写字母表示同一林型相同处理下不同采样时间土壤胡敏酸/可提取腐殖质差异显著(p < 0.05)。 *, 同一采样时间凋落物自然输入与凋落物去除之间差异显著(p < 0.05)。

Fig. 3 Humic acid/extractable humic substances ratio as affected by litter input in broad-leaved forest, mixed coniferous and broad-leaved forest and coniferous forest in subalpine forests of western Sichuan (mean ± SE). Different lowercase letters indicate that the differences in humic acid/extractable humic substances ratio between measurement times are significant (p < 0.05). *, significant differences in humic acid/extractable humic substances ratio between litter treatments in the same measurement time (p < 0.05).

表4 川西亚高山森林不同凋落物处理下土壤腐殖质与土壤理化性质之间的相关性

Table 4 Correlations of soil humic substances with soil physical and chemical properties under different litter treatments in subalpine forests of western Sichuan

指标 Index	凋落物输入 Soil retained litter					凋落物去除 Soil removed litter
指标 Index	EHS	HA	FA	HA/FA	PQ	EHS	HA	FA	HA/FA	PQ
SWC	0.157	0.223	0.089	0.168	-0.021	0.564^**	0.355	0.472^*	-0.389^*	-0.018
pH	-0.122	-0.213	-0.068	0.023	0.025	-0.194	-0.139	-0.154	-0.004	0.179
TM	-0.493^**	-0.547^**	-0.338	0.005	0.007	-0.512^**	-0.432^*	-0.370	0.062	0.015
SOC	0.578^**	0.707^**	0.366	0.105	0.025	0.755^**	0.409^*	0.656^**	-0.399^*	-0.107
TN	0.414^*	0.513^**	0.257	0.181	-0.012	0.658^**	0.418^*	0.546^**	-0.185	-0.127
TP	0.338	0.218	0.315	-0.181	-0.156	0.314	0.387^*	0.182	0.218	-0.055

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川西亚高山森林凋落物去除对土壤腐殖质动态的影响

Litter removal effects on dynamics of soil humic substances in subalpine forests of western Sichuan, China

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