不同密度杉木林对林下植被和土壤微生物群落结构的影响

doi:10.17521/cjpe.2020.0158

摘要/Abstract

摘要：

为揭示土壤养分和细菌群落对林下植被调控的响应机制, 调查了浙江开化3种林分密度(高密度(KH)、中密度(KM)和低密度(KL))的17年生杉木人工林林下植被和生物量, 测定土壤理化性质, 并基于16S rDNA高通量测序技术分析细菌群落结构变化。结果表明, 3种密度的杉木林下植被地上部分总生物量为0.10-2.10 t·hm^-2, 且优势植物物种差异显著。理化性质测定分析发现, 高密度与低密度林分的土壤pH、有效磷含量差异显著。相关性分析表明, 土壤pH与林下植被中草本、灌木生物量及总生物量均呈显著正相关关系, 土壤有机质含量与灌木植被生物量及林下植被总生物量呈显著正相关关系, 速效钾含量与灌木植被生物量呈显著正相关关系。土壤微生物群落结构分析可知, 3种密度杉木林地土壤中酸杆菌门、变形菌门、放线菌门和绿弯菌门为优势菌群, 总相对丰度占比超过80%。冗余分析(RDA)表明土壤pH、碱解氮、有效磷和速效钾含量是土壤细菌群落结构变化的关键影响因素。酸杆菌门的优势亚群为Gp2、Gp1、Gp3和Gp6, 占酸杆菌群的51.32%-57.38%, 且随林分密度降低, 林下植被增多, Gp1占比增大, Gp2和Gp6占比下降; Gp6相对丰度与pH呈极显著负相关关系。可见, 杉木纯林经营中适度降低林分密度有利于林下植被生长和良好细菌群落结构保持, 有利于维持杉木林地土壤肥力, 实现可持续经营。

关键词: 杉木, 林下植被, 生物量, 土壤理化性质, 细菌, 酸杆菌

Abstract:

Aims The aim of the present study was to investigate the responses of understory vegetation to soil nutrients and bacterial communities.
Methods This study investigated the understory vegetation biomasses and species composition as well as soil physical and chemical properties in 17-year-old Chinese fir plantations with three densities (high-density (KH), medium-density (KM), and low-density (KL)) in Kaihua, Zhejiang. The changes of bacterial community structures were analyzed via 16S rDNA high-throughput sequencing techniques.
Important findings The result showed that the total above-ground biomass of the understory vegetation ranged from 0.10 to 2.10 t·hm^-2 and the dominant plant species varied in three Chinese fir plantations. The soil pH and available phosphorus content were significantly different between high-density and low-density forest stands. Correlation analysis showed that soil pH was positively correlated with the biomass of herbs, shrubs and the total of understory vegetation, while the content of soil organic matter was just positively related with the last two factors, and the available potassium content was only affected by the biomass of shrub. Based on the analysis of the soil microbial community, the Acidobacteria, Proteobacteria, Actinobacteria, and Chloroflexi were the dominant phyla in the three Chinese fir plantations. Redundancy analysis showed that soil pH, available nitrogen, available phosphorus and available potassium contents played a crucial role in regulating the soil bacterial community structures. Gp2, Gp1, Gp3 and Gp6 were the dominant subgroups of Acidobacteria, accounting for 51.32%- 57.38% of the Acidobacteria. With the decline of the Chinese fir density, the biomass of understory vegetation and the proportion of Gp1 increased, while the proportion of Gp2 and Gp6 decreased and the relative abundance of Gp6 was negatively correlated with soil pH. Obviously, the moderate reduction in stand density of pure Chinese fir forests was beneficial in the growth of understory vegetation and in maintaining a reasonable bacterial community structure, which helps to maintain the soil fertility of the Chinese fir forests and to achieve sustainable management in the long run.

Key words: Chinese fir, understory vegetation, biomass, soil physical and chemical properties, bacteria, Acidobacteria

丁凯, 张毓婷, 张俊红, 柴雄, 周世水, 童再康. 不同密度杉木林对林下植被和土壤微生物群落结构的影响. 植物生态学报, 2021, 45(1): 62-73. DOI: 10.17521/cjpe.2020.0158

DING Kai, ZHANG Yu-Ting, ZHANG Jun-Hong, CHAI Xiong, ZHOU Shi-Shui, TONG Zai-Kang. Effects of Chinese fir plantations with different densities on understory vegetation and soil microbial community structure. Chinese Journal of Plant Ecology, 2021, 45(1): 62-73. DOI: 10.17521/cjpe.2020.0158

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0158

https://www.plant-ecology.com/CN/Y2021/V45/I1/62

图/表 16

表1 不同密度杉木林试验样地的基本概况(平均值±标准差)

Table 1 Basic information of the test plots of Chinese fir plantations with different densities (mean ± SD)

类型 Type	林分密度 Stand density (plant·hm^-2)	平均胸径 Average DBH (cm)	平均树高 Average tree height (m)	郁闭度 Canopy density
KH	1 783.33 ± 76.38 ^a	14.3 ± 2.55 ^b	12.8 ± 0.69 ^c	0.96 ± 0.02 ^a
KM	1 616.67 ± 52.65 ^b	15.6 ± 1.40 ^b	14.4 ± 1.36 ^b	0.84 ± 0.02 ^b
KL	1 436.25 ± 28.87 ^c	17.4 ± 0.95 ^a	16.3 ± 0.31 ^a	0.78 ± 0.01 ^c

表2 不同密度杉木人工林林下植被种类

Table 2 Understory vegetation types of Chinese fir plantations with different densities

类型 Type		林下植被种类 Understory vegetation type
KH		檵木、豆腐柴、短尾越桔、狗脊 Loropetalum chinense, Premna microphylla, Vaccinium carlesii, Woodwardia japonica
KM		檵木、五节芒、黄瑞木、菝葜、狗脊、海金沙、三脉紫菀 L. chinense, Miscanthus floridulus, Adinandra millettii, Smilax china, W. japonica, Lygodium japonicum, Aster trinervius subsp.ageratoides
KL		檵木、芒萁、狗脊、渐尖毛蕨、淡竹叶、多花黄精、烟管头草、假福王草、大叶白纸扇、大青、寒莓、华紫珠、楤木 Loropetalum chinense, Dicranopteris pedata, W. japonica, Cyclosorus acuminatus, Lophatherum gracile, Polygonatum cyrtonema, Carpesium cernuum, Paraprenanthes sororia, Mussaenda shikokiana, Clerodendrum cyrtophyllum, Rubus buergeri, Callicarpa cathayana, Aralia chinensis

表3 不同密度杉木人工林林下植被生物量(平均值±标准差)

Table 3 Understory vegetation biomass of Chinese fir plantations with different densities (mean ± SD)

类型 Type	优势植物种类 Dominate species	地上部分生物量 Aboveground biomass (t·hm^-2)		总计 Total (t·hm^-2)
类型 Type	优势植物种类 Dominate species	灌木层 Shrub layer	草本层 Herb layer	总计 Total (t·hm^-2)
KH	檵木 Loropetalum chinense	0.09 ± 0.15 ^b	0.01 ± 0.02 ^b	0.10 ± 0.17 ^b
KM	檵木、五节芒 L. chinense, Miscanthus floridulus	0.26 ± 0.02 ^b	0.15 ± 0.02 ^b	0.40 ± 0.07 ^b
KL	檵木、芒萁、狗脊、渐尖毛蕨 L. chinense, Dicranopteris pedata, Woodwardia japonica, Cyclosorus acuminatus	0.87 ± 0.15 ^a	1.23 ± 0.15 ^a	2.10 ± 0.57 ^a

表4 不同密度杉木人工林土壤的理化性质(平均值±标准差)

Table 4 Soil physical and chemical properties of Chinese fir plantation with different densities (mean ± SD)

类型 Type	pH	束缚水含量(CW) Water content (%)	碱解氮含量 Alkali-hydrolyzable N content (mg·kg^-1)	有效磷含量 Available P content (mg·kg^-1)	速效钾含量 Available K content (mg·kg^-1)	有机质含量 Organic matter content (g·kg^-1)
KH	3.44 ± 0.06 ^b	3.17 ± 0.44 ^a	142.18 ± 9.30 ^a	2.51 ± 0.18 ^b	54.24 ± 9.24 ^a	34.29 ± 4.11 ^a
KM	3.52 ± 0.06 ^ab	3.63 ± 0.45 ^a	156.88 ± 6.54 ^a	3.31 ± 0.45 ^a	66.91 ± 11.63 ^a	38.58 ± 5.17 ^a
KL	3.62 ± 0.03 ^a	3.34 ± 0.55 ^a	156.63 ± 1.42 ^a	3.40 ± 0.32 ^a	74.72 ± 3.90 ^a	44.58 ± 1.81 ^a

表5 不同密度杉木林林下植被地上生物量与土壤理化性质相关性分析

Table 5 Correlation analysis of aboveground biomass of understory vegetation and soil physical and chemical properties of Chinese fir plantations with different densities

类型 Type	pH	束缚水含量 Water content	碱解氮含量 Alkali-hydrolyzable N content	有效磷含量 Available P content	速效钾含量 Available K content	有机质含量 Organic matter content
草本生物量 Herb biomass	0.764^*	0.191	0.485	0.514	0.612	0.749
灌木生物量 Shrub biomass	0.874^*	0.193	0.564	0.648	0.760^*	0.820^*
总生物量 Total biomass	0.814^*	0.193	0.520	0.571	0.675	0.783^*

图1 不同处理可操作分类单元(OTU)数韦恩图。KH, 高密度; KL, 低密度; KM, 中密度。

Fig. 1 Venn diagram of different processing Operational Taxonomy Unit (OTU) numbers. KH, high-density; KL, low-density; KM, medium-density.

表6 不同密度杉木林土壤细菌群落多样性(平均值±标准差)

Table 6 Diversity of soil bacterial communities in different densities of Chinese fir forests (mean ± SD)

类型 Type	物种数 Observed species	Shannon指数 Shannon index	Chao1指数 Chao1 index	覆盖率 Coverage (%)
KH	4 939.80 ± 173.89 ^a	10.21 ± 0.04 ^a	6 223.62 ± 445.90 ^a	0.973 2 ± 0.004 ^a
KM	5 171.25 ± 320.72 ^a	10.27 ± 0.25 ^a	6 468.85 ± 109.98 ^a	0.965 3 ± 0.005 ^a
KL	5 277.67 ± 263.71 ^a	10.30 ± 0.14 ^a	6 619.39 ± 214.13 ^a	0.975 3 ± 0.006 ^a

图2 不同密度杉木林土壤优势细菌门。Acidobacteria, 酸杆菌门; Actinobacteria, 放线菌门; Actinomycetales, 放线菌目; Chloroflexi, 绿弯菌门; Firmicutes, 厚壁菌门; Gammaproteobacteria, γ-变形菌; Planctomycetes, 浮霉菌门; Proteobacteria, 变形菌门; Rhodospirillales, 红螺菌目; Spartobacteria, 斯巴杆菌纲; Verrucomicrobia, 疣微菌门。Gp, 酸杆菌门亚群; unclassified, 未分类细菌。KH, 高密度; KL, 低密度; KM, 中密度。

Fig. 2 Soil dominant bacteria in different densities of Chinese fir forests. Gp, Acidobacteria subgroup. KH, high-density; KL, low-density; KM, medium-density.

附录I 不同密度杉木林土壤细菌群落结构差异分析

Supplement I Analysis of differences in soil bacterial community structure of Chinese fir forests with different densities

计算距离 Calculate distance	F	p
未加权的 Unweighted_unifrac	1.12	0.19
加权的 Weighted_unifrac	1.09	0.41

表7 不同密度杉木林下土壤理化性质对细菌群落结构的影响

Table 7 Effects of soil physical and chemical properties on bacterial community structure of Chinese fir plantations with different densities

理化指标 Physical and chemical indicator	Mds1	Mds2	R²	p
pH	-0.580 22	-0.814 46	0.159 7	0.747
束缚水含量 CW	-0.572 27	-0.820 07	0.131 1	0.768
碱解氮含量 AN	0.098 13	-0.995 17	0.803 1	0.049^*
有效磷含量 AP	0.128 86	-0.991 66	0.565 5	0.140
速效钾含量 AK	-0.121 23	-0.992 62	0.724 7	0.049^*
有机质含量 OM	-0.564 26	-0.825 60	0.472 8	0.267

图3 不同密度杉木林土壤理化性质与细菌门冗余分析(RDA)。AK, 速效钾含量; AN, 碱解氮含量; AP, 有效磷含量; CW, 束缚水含量; OM, 有机质含量。Acidobacteria, 酸杆菌门; Actinobacteria, 放线菌门; Chloroflexi, 绿弯菌门; Firmicutes, 厚壁菌门; Planctomycetes, 浮霉菌门; Proteobacteria, 变形菌门; Verrucomicrobia, 疣微菌门。

Fig. 3 Redundancy analysis (RDA) of soil physical and chemical properties and bacteria of Chinese fir plantations with different densities. AK, available K content; AN, alkali- hydrolyzable N content; AP, available P content; CW, water content; OM, organic matter content.

图4 不同密度杉木林土壤细菌属水平相对丰度。Acidobacteria, 酸杆菌门; Actinobacteria, 放线菌门; Bradyrhizobium, 慢生根瘤菌属; Chloroflexi, 绿弯菌门; Firmicutes, 厚壁菌门; Ktedonobacter, 纤线杆菌属; Planctomycetes, 浮霉菌门; Proteobacteria, 变形菌门; Rhodospirillales, 红螺菌目; Spartobacteria, 斯巴杆菌纲; Verrucomicrobia, 疣微菌门。Gp, 酸杆菌门亚群; other, 其他; unclassified, 未分类细菌。KH, 高密度; KL, 低密度; KM, 中密度。

Fig. 4 Relative abundance of bacteria genera in different densities of Chinese fir forests. Gp, Acidobacteria subgroup. KH, high-density; KL, low-density; KM, medium-density.

图5 不同密度杉木林土壤的酸杆菌门亚群(Gp)相对丰度聚类。KH, 高密度; KL, 低密度; KM, 中密度。

Fig. 5 Clustering of relative abundance of Acidobacteria subgroups (Gp) in different densities of Chinese fir plantations. KH, high-density; KL, low-density; KM, medium-density.

表8 不同密度杉木林酸杆菌优势亚群(Gp)与土壤理化性质的相关性

Table 8 Pearson correlation coefficients between dominant Acidobacteria subgroups (Gp) and soil physicochemical properties of Chinese fir plantations with different densities

分类 Item	pH	束缚水含量 Water content	碱解氮含量 Alkali-hydrolyzable N content	有效磷含量 Available P content	速效钾含量 Available K content	有机质含量 Organic matter content
Gp1	0.257	-0.043	0.118	0.455	0.228	-0.075
Gp2	-0.336	-0.077	0.266	0.316	0.142	0.416
Gp3	-0.006	0.164	0.573	0.693	0.293	0.353
Gp6	-0.890^**	0.178	-0.222	0.479	-0.642	-0.475

附录II 不同密度杉木林的指示物种(门水平)

Supplement II Indicator species of Chinese fir forest with different densities (phylum level)

类型 Type	指示物种 Indicator species	p
高密度 KH	Latescibacteria	0.048^*
低密度 KL	Candidatus_saccharibacteria	0.008^**

附录III 不同密度杉木林的指示物种(科水平)

Supplement III Indicator species of Chinese fir forest with different densities (family level)

类型 Type	指示物种 Indicator species	p
高密度 KH	Dehalococcoidaceae	0.038^*
	Phycisphaeraceae	0.036^*
	Geobacteraceae	0.027^*
低密度 KL	Candidatus_Adlerbacteria_unclassified	0.028^*
	Sphingobacteriaceae	0.048^*
	Saccharibacteria_genera_incertae_sedis	0.037^*
	Gemmatimonadaceae	0.033^*

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