川西亚高山岷江冷杉幼苗生物量积累与分配的影响因子

doi:10.17521/cjpe.2024.0018

植物生态学报 ›› 2024, Vol. 48 ›› Issue (11): 1459-1470.DOI: 10.17521/cjpe.2024.0018 cstr: 32100.14.cjpe.2024.0018

川西亚高山岷江冷杉幼苗生物量积累与分配的影响因子

陶琼¹^,², 缪宁¹^,^*(), 岳喜明¹^,³, 罗建琼¹, 薛盼盼¹, 王晖⁴

¹四川大学生命科学学院, 教育部生物资源与生态环境重点实验室, 成都 610065
²四川省天然林保护中心, 成都 610081
³四川省泸州市生态环境保护综合行政执法支队, 四川泸州 646018
⁴中国林业科学研究院森林生态环境与自然保护研究所, 国家林业和草原局森林生态环境重点实验室, 北京 100091

收稿日期:2024-01-23 接受日期:2024-08-23 出版日期:2024-11-20 发布日期:2024-08-23
通讯作者: *缪宁(miaoning@scu.edu.cn)
基金资助:
国家重点研发计划(2022YFD2200504-02)

Influencing factors of biomass accumulation and allocation of Abies fargesii var. faxoniana seedlings in the subalpine region of western Sichuan, China

TAO Qiong¹^,², MIAO Ning¹^,^*(), YUE Xi-Ming¹^,³, LUO Jian-Qiong¹, XUE Pan-Pan¹, WANG Hui⁴

¹College of Life Sciences, Sichuan University, Key Laboratory of Bio-resource and Eco-environment of Ministry of Education, Chengdu 610065, China
²Sichuan Natural Forest Protection Center, Chengdu 610081, China
³Comprehensive Administrative Enforcement Detachment of Ecology and Environment Protection of Luzhou City, Sichuan Province, Luzhou, Sichuan 646018, China
⁴Key Laboratory of Forest Ecology and Environment of National Forestry and Grassland Administration, Ecology and Nature Conservation Institute, Chinese Academy of Forestry, Beijing 100091, China

Received:2024-01-23 Accepted:2024-08-23 Online:2024-11-20 Published:2024-08-23
Contact: *MIAO Ning (miaoning@scu.edu.cn)
Supported by:
National Key R&D Program of China(2022YFD2200504-02)

摘要/Abstract

摘要：

岷江冷杉(Abies fargesii var. faxoniana)的天然更新是川西亚高山地区天然次生林生态修复的关键, 对其幼苗生物量积累和分配的关键制约因子进行研究有助于深入理解其更新和生长机理。该研究对比了红桦(Betula albosinensis)阔叶林、红桦-岷江冷杉针阔混交林和岷江冷杉原始林3种林型中岷江冷杉幼苗的生物量分配格局和器官间的异速生长关系, 分析了幼苗的生物量分配与地形和微生境因子之间的关系, 并定量分解了各因子对生物量积累影响的贡献度。岷江冷杉原始林中幼苗的根生物量分配比例(30.3%)和茎生物量分配比例(43.3%)显著高于红桦-岷江冷杉针阔混交林和红桦阔叶林, 而叶生物量分配比例(26.4%)显著低于红桦-岷江冷杉针阔混交林(37.4%)和红桦阔叶林(42.3%)。红桦阔叶林中幼苗的根、茎和叶的生物量呈现等速生长关系, 红桦-岷江冷杉针阔混交林和岷江冷杉原始林中幼苗的叶-根间亦呈等速生长关系, 但叶-茎和根-茎间则为异速生长关系。相比于东北、北和西北坡, 西坡生境中的幼苗年平均生物量的积累最高。坡向(20.9%)对幼苗生物量积累和分配的贡献率大于海拔(18.1%), 基质类型(15.8%)、苔藓盖度(11.7%)及厚度(7.7%)和冠层盖度(7.4%)是微生境因子中对岷江冷杉幼苗生物量累积贡献度前四的因子。该研究可为川西亚高山次生林中岷江冷杉更新以及次生林的结构优化提供支撑。

关键词: 岷江冷杉幼苗, 生物量分配, 异速生长, 冗余分析, 微生境因子

Abstract:

Aims The natural regeneration of Abies fargesiivar. faxoniana is crucial for the ecological restoration of natural secondary forests in the subalpine region of western Sichuan. The study of the key factors in the biomass accumulation and allocation of seedlings is helpful to understand the mechanism of regeneration. Therefore, our objective is to reveal the critical factors for the biomass accumulation of A. fargesiivar. faxoniana seedlings in the Betula albosinensis broadleaf forests (BB), B. albosinensis- A. fargesii var. faxoniana needleleaf-broadleaf forests (BA), and A. fargesii var. faxoniana primary forests (AP).

Methods We compared the biomass allocation patterns of A. fargesiivar. faxoniana seedlings in different forest types and investigated allometric relationships between different organs by a standardized major axis regression analysis. We revealed the relationships between seedlings’ biomass and habitat factors through a redundancy analysis followed by a quantitative decomposition of the contribution of influencing factors.

Important findings The root biomass fraction (30.3%) and stem biomass fraction (43.3%) in the AP forest were significantly higher than that in the BA and BB forests. Correspondingly, leaf biomass fraction (26.4%) in the AP forest was significantly lower than that in the BA and BB forests. Seedlings in the BB forest showed an isometric relationship among three organs: roots, stems, and leaves. In the BA and AP forests, seedlings exhibited an isometric relationship between leaves and roots. In contrast, allometric relationships were observed between leaves and stems and between roots and stems. The annual biomass increment of seedlings was the highest in the western slope habitats compared to those on the northeast, north, and northwest slopes. The contribution rate of the slope aspect (20.9%) to seedling biomass accumulation and allocation surpassed that of altitude (18.1%), with substrate type (15.8%), moss cover density (11.7%) and thickness (7.7%), and canopy coverage (7.4%) ranking the top four among microhabitat factors. This study can assist in the management of Minjiang fir regeneration and the technical optimization of structures of secondary subalpine forests in western Sichuan.

Key words: Abies fargesii var. faxoniana seedlings, biomass allocation, allometric growth, redundancy analysis, microhabitat factors

陶琼, 缪宁, 岳喜明, 罗建琼, 薛盼盼, 王晖. 川西亚高山岷江冷杉幼苗生物量积累与分配的影响因子. 植物生态学报, 2024, 48(11): 1459-1470. DOI: 10.17521/cjpe.2024.0018

TAO Qiong, MIAO Ning, YUE Xi-Ming, LUO Jian-Qiong, XUE Pan-Pan, WANG Hui. Influencing factors of biomass accumulation and allocation of Abies fargesii var. faxoniana seedlings in the subalpine region of western Sichuan, China. Chinese Journal of Plant Ecology, 2024, 48(11): 1459-1470. DOI: 10.17521/cjpe.2024.0018

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图/表 8

表1 川西亚高山样点的林分特征和岷江冷杉幼苗生物量样本量

Table 1 Stand characteristics of sample sites and sample size of Abies fargesii var. faxoniana seedlings biomass in subalpine area of western Sichuan

林分类型 Forest type	样点 Sample site	样点编号 Site code	背景 Background	乔木层 Tree layer	灌木层 Shrub layer	草本层 Herb layer	坡向 Aspect slope	坡度 Slope (°)	海拔 Altitude (m)	样本量 Sample size
BB	SHG	1	盖度 Coverage (%)	70	35	85	NW	25-28	3 396	45
BB	SHG	1	平均高度 Average height (m)	13	3.5	0.4	NW	25-28	3 396	45
BA	272	2	盖度 Coverage (%)	60	35	70	N	25-35	3 258	72
BA	272	2	平均高度 Average height (m)	14	3.5	0.4	N	25-35	3 258	72
AP	189	3	盖度 Coverage (%)	70	20	45	N	35-37	3 995	44
	189	4	平均高度 Average height (m)	12	2.0	0.3	N	35-40	3 793	44
	SJB	5	盖度 Coverage (%)	47	25	27	W	32-35	3 958	44
	SJB	6	平均高度 Average height (m)	12	2.0	0.2	W	27-30	3 793	45
	295	7	盖度 Coverage (%)	60	50	40	NW	30-33	3 974	43
	295	8	平均高度 Average height (m)	10	4.0	0.3	NW	30-33	3 812	45
	JBG	9	盖度 Coverage (%)	65	25	10	NE	35-38	4 008	45
	JBG	10	平均高度 Average height (m)	13	5.0	0.1	NE	30-35	3 780	45

图1 川西亚高山岷江冷杉幼苗样点位置相对关系。E, 东坡; N, 北坡; NE, 东北坡; NW, 西北坡; W, 西坡。1-10为采样样点编号, 详细信息见表1。

Fig. 1 Relative sample site locations of Abies fargesii var. faxoniana seedlings in subalpine area of western Sichuan. E, east slope aspect; N, north slope aspect; NE, northeast slope aspect; NW, northwest slope aspect; W, west slope aspect. 1-10 are sample site codes, detailed information see Table 1.

表2 不同林型内岷江冷杉幼苗所处微生境的特征

Table 2 Traits of microhabitats around Abies fargesii var. faxoniana seedlings in three forest types

变量 Variable	缩写 Abbreviation	林型 Forest type	平均值±标准误 Mean ± SE
郁闭度 Canopy coverage (%)	CC	AP	46 ± 2^c
		BA	59 ± 2^b
		BB	72 ± 1^a
灌木盖度 Shrub coverage (%)	SC	AP	18 ± 1^a
		BA	7 ± 2^b
		BB	9 ± 2^b
灌木高度 Shrub height (m)	SH	AP	0.43 ± 0.04^b
		BA	0.88 ± 0.26^a
		BB	0.32 ± 0.05^b
箭竹盖度 Fargesia coverage (%)	FC	AP	-
		BA	29 ± 3^a
		BB	4 ± 1^b
箭竹高度 Fargesia height (m)	FH	AP	-
		BA	1.38 ± 0.16^a
		BB	0.22 ± 0.03^b
草本盖度 Herbage coverage (%)	HC	AP	22 ± 1^c
		BA	34 ± 3^a
		BB	27 ± 4^b
草本高度 Herbage height (m)	HH	AP	0.08 ± 0.01^b
		BA	0.23 ± 0.01^a
		BB	0.17 ± 0.02^a
苔藓盖度 Moss coverage (%)	MC	AP	88 ± 1^a
		BA	44 ± 4^b
		BB	32 ± 5^b
苔藓厚度 Moss thicknesses (cm)	MT	AP	6.62 ± 0.19^a
		BA	2.35 ± 0.16^b
		BB	2.32 ± 0.24^b
凋落物盖度 Litter coverage (%)	LC	AP	25 ± 1^b
		BA	48 ± 2^a
		BB	22 ± 2^b
凋落物厚度 Litter thicknesses (cm)	LT	AP	1.26 ± 0.07^b
		BA	2.73 ± 0.17^a
		BB	2.64 ± 0.29^a

图2 不同林型岷江冷杉幼苗生物量年增量(A)和器官生物量比例(B) (平均值±标准误)。不同大写字母表示不同器官间差异显著(p < 0.05), 不同小写字母表示不同林型间差异显著(p < 0.05)。AP, 岷江冷杉原始林; BA, 红桦-岷江冷杉针阔混交林; BB, 红桦阔叶林。

Fig. 2 Annual biomass increment (A) and organ biomass fraction (B) of Abies fargesii var. faxoniana seedlings in different forest types (mean ± SE). Different uppercase letters indicate significant differences between organs (p < 0.05), different lowercase letters indicate significant differences between the forest types (p < 0.05). AP, Abies fargesii var. faxoniana primary forest; BA, Betula albosinensis - Abies fargesii var. faxoniana needleleaf-broadleaf forest; BB, Betula albosinensis broadleaf forest.

图3 不同林型岷江冷杉幼苗的总生物量与年龄的关系。所有回归模型均在α = 0.001水平上显著。AP, 岷江冷杉原始林; BA, 红桦-岷江冷杉针阔混交林; BB, 红桦阔叶林。

Fig. 3 Relationships between total seedling biomass and seedling age of Abies fargesii var. faxoniana in different forest types. All regression models were significant at the α = 0.001 level. AP, Abies fargesii var. faxoniana primary forest; BA, Betula albosinensis - Abies fargesii var. faxoniana needleleaf-broadleaf forest; BB, Betula albosinensis broadleaf forest.

图4 不同林型岷江冷杉幼苗各器官间(A, 叶-根; B, 茎-根; C, 叶-茎)生物量的异速生长方程。所有回归模型均在α = 0.001水平上显著。p表示斜率与1.0的差异显著性。AP, 岷江冷杉原始林; BA, 红桦-岷江冷杉针阔混交林; BB, 红桦阔叶林。

Fig. 4 Allometric relationships for different dimensions (A, leaf-root; B, stem-root; C, leaf-stem) of Abies fargesii var. faxoniana seedlings in different forest types. All regression models were significant at the α = 0.001 level. p indicates a significant difference in slope with 1.0. AP, Abies fargesii var. faxoniana primary forest; BA, Betula albosinensis - Abies fargesii var. faxoniana needleleaf-broadleaf forest; BB, Betula albosinensis broadleaf forest.

图5 岷江冷杉幼苗生物量与各生境因子的冗余分析(RDA) (A)和单个因子独立贡献率(B)。该模型在α = 0.001水平上显著。AL, 海拔; AMI, 平均年生物量增量; AS, 坡向; CC, 冠层盖度; FC, 箭竹盖度; FH, 箭竹高度; HC, 草本盖度; HH, 草本高度; LC, 凋落物盖度; LMF, 叶生物量分配比例; LT, 凋落物厚度; MC, 苔藓盖度; MT, 苔藓厚度; RMF, 根生物量分配比例; SC, 灌木盖度; SH, 灌木高度; SMF, 茎生物量分配比例; SU, 基质类型。log, 倒木; rock, 岩石; soil, 土壤; stump, 树桩。N, 北坡; NE, 东北坡; NW, 西北坡; W, 西坡。

Fig. 5 Redundancy analysis (RDA) diagram of all factors and biomass of Abies fargesii var. faxoniana seedlings (A) and the relative importance of individual factors (B). The model was significant at the α = 0.001 level. AL, altitude; AMI, annual biomass increment; AS, aspect slope; CC, canopy coverage; FC, Fargesia coverage; FH, Fargesia height; HC, herbage overage; HH, herbage height; LC, litter coverage; LMF, leaf biomass fraction; LT, litter thickness; MC, moss coverage; MT, moss thickness; RMF, root biomass fraction; SC, shrub coverage; SH, shrub height; SMF, stem biomass fraction; SU, substrate type. N, north slope aspect; NE, northeast slope aspect; NW, northwest slope aspect; W, west slope aspect.

表3 不同林分中岷江冷杉幼苗生物量与各生境因子的关系

Table 3 Relationship between Abies fargesii var. faxoniana seedling biomass and habitat factors in different forest types

林分类型 Forest type	响应变量 Response variable	解释变量 Explanatory variable	参数 Estimate	标准误 SE	t	p
岷江冷杉原始林 Abies fargesii var. faxoniana primary forest	RMF	AS (NW)	0.44	0.15	2.93	0.004^**
		SU (soil)	0.72	0.23	3.06	0.002^**
		SH	-0.17	0.08	-2.06	0.041^*
		HC	-0.14	0.07	-2.01	0.045^*
		MT	0.25	0.08	3.22	0.001^**
	SMF	AS (W)	0.51	0.14	3.52	<0.001^***
	LMF	AS (NW)	-0.47	0.17	-2.68	0.030^*
		AS (W)	-0.58	0.18	-3.12	0.013^*
		SU (rock)	-0.90	0.35	-2.54	0.012^*
		SU (soil)	-0.64	0.23	-2.80	0.005^**
		MT	-0.19	0.07	-2.75	0.008^**
	AMI	AS (W)	0.67	0.20	3.33	0.009^**
	AMI	SH	0.20	0.08	2.51	0.012^*
红桦-岷江冷杉针阔混交林 Betula. albosinensis - Abies fargesii var. faxoniana needleleaf-broadleaf forest	RMF	MC	0.49	0.15	3.27	0.002^**
	RMF	MT	-0.65	0.31	-2.06	0.044^*
	SMF	ns
	LMF	ns
	AMI	ns
红桦阔叶林 Betula albosinensis broadleaf forest	RMF	HH	0.34	0.14	2.48	0.018^*
	SMF	ns
	LMF	HC	-0.35	0.15	-2.24	0.031^*
	AMI	CC	-0.35	0.15	-2.40	0.021^*
		FC	1.59	0.49	3.24	0.002^**
		HH	-0.29	0.12	-2.41	0.020^*

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川西亚高山岷江冷杉幼苗生物量积累与分配的影响因子

Influencing factors of biomass accumulation and allocation of Abies fargesii var. faxoniana seedlings in the subalpine region of western Sichuan, China

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