杉木纯林林下补植模式对土壤质量和微生物群落的影响

doi:10.11707/j.1001-7488.LYKX20250050

Abstract

Abstract:

Objective: To address ecological issues such as soil degradation caused by long-term continuous monoculture of Cunninghamia lanceolata (Chinese fir), it is necessary to establish and transform plantations into multi-layered, uneven-aged mixed forests of Chinese fir and broadleaved species to enhance the sustainability of forest ecosystems. This study evaluated changes in soil quality under different understory enrichment planting modes of native tree species, aiming to identify suitable understory enrichment planting modes and provide a scientific basis for maintaining soil health and sustainable management of Chinese fir plantations. Method: In a Chinese fir plantation established in 1998, the third thinning was conducted in 2014 (retaining an average density of 225 tree·hm^–2). In 2015, four understory enrichment planting modes were conducted: a pure plantation served as the control without thinning or planting (M0), understory replanting of Phoebe bournei only (M1), understory replanting of Phoebe bournei and Taxus wallichiana var. chinensis (M2), and understory replanting of P. bournei, T. wallichiana var. chinensis, and Schima superba (M3). The proportion of Chinese fir to understory planted species was 3∶7 in all modes. In 2022, seven years after understory replanting, soil samples were collected from 0–60 cm depth to determine chemical properties, extracellular enzyme activities, and bacterial community characteristics (16S rRNA). A minimum data set (MDS) was selected to establish a soil quality index (SQI) model. Variance partitioning analysis (VPA) was used to quantify the contributions of biotic factors (enzyme activities, microbial metabolic limitations, and community structure) and abiotic factors (chemical properties and stoichiometric ratios) to soil quality. Structural equation model (SEM) was applied to analyze the relationships among “understory replanting mode–soil quality index”. Result: Compared with the pure plantation (M0), the multi-layered uneven-aged Chinese fir and broad-leaved species mixed forests (M1, M2, M3) formed by understory replanting significantly improved soil quality and microbial ecological functions. The content of key nutrients in the 0–20 cm soil layer, including total nitrogen (TN), total phosphorus (TP), available nitrogen (AN), available potassium (AK), and soil organic carbon (SOC), increased by 29.87%–72.62%, with the greatest improvements observed in M2. Understory enrichment planting drove carbon, nitrogen, and phosphorus cycling by optimizing soil enzyme activities. The soil sucrase (SUC) activity and glucosidase (GLU) activity of M1, and urease (URE) activity of M3 reached their peak in the 0–20 cm layer. In contrast, the pure plantation had higher acid phosphatase (ACP) and catalase (CAT) activities, reflecting differences in catabolic pathways. The average microbial phosphorus limitation in M2 decreased by 12.05% compared with the pure plantation. Additionally, M2 optimized nutrient utilization strategies by enhancing the relative activity of nitrogen hydrolase (RAN) in the 0–20 cm layer and alleviating microbial carbon limitation in the 40–60 cm layer. Microbial community analysis revealed that the established multi-layered uneven-aged mixed forests significantly increased the Shannon diversity index of soil microorganisms and altered the abundance of functional bacterial phyla such as Acidobacteriota and Proteobacteria. Understory enrichment planting modes indirectly affected soil quality by regulating total phosphorus (path coefficient: 0.68), nitrogen-to-phosphorus ratio (0.71), SOC (0.33), pH (0.34), nitrogen hydrolase activity (0.17), and soil microbial Shannon diversity (0.60). The SQI values of the three understory enrichment planting modes were higher than that of the pure plantation, with M2 exhibiting the highest soil quality in the 0–20 cm layer and the pure plantation showing the lowest soil quality in the 40–60 cm layer. Conclusion: Transforming pure Chinese fir plantations into multi-layered uneven-aged mixed forests by understory replanting native broadleaved species can enhance soil nutrient levels, balance stoichiometric limitations, optimize enzyme activities and microbial diversity, and improve overall soil quality. Among them, understory replanting of P. bournei and T. wallichiana var. chinensis (M2) demonstrates the most significant soil improvement effects. These findings provide a theoretical basis for the sustainable management of Chinese fir plantations.

Key words: Chinese fir plantation, Chinese fir-broadleaf mixed forest, soil quality, soil enzyme activity, soil microbial community

CLC Number:

S714

Yuxin Hu,Yihang Jiang,Zhenhua Liu,Guangyu Zhu,Jianguo Zhang,Xiongqing Zhang. Effects of Understory Enrichment Planting Modes in Pure Chinese fir Forests on Soil Quality and Microbial Communities[J]. Scientia Silvae Sinicae, 2026, 62(3): 61-73.

Figures/Tables 10

Table 1

Table 2

Table 3

Soil chemical properties and stoichiometric ratios of different understory enrichment planting modes and soil layers"

指标Index	M0			M1			M2			M3
指标Index	0~20 cm	20~40 cm	40~60 cm	0~20 cm	20~40 cm	40~60 cm	0~20 cm	20~40 cm	40~60 cm	0~20 cm	20~40 cm	40~60 cm
TN/(g·kg^–1)	0.94± 0.07bA	1.00± 0.02abA	0.72± 0.04bB	1.14± 0.07aA	1.04± 0.06aA	0.91± 0.10aB	1.18± 0.09aA	0.92± 0.07bB	0.76± 0.03bC	1.22± 0.08aA	0.97± 0.04abB	0.79± 0.03bC
TP /(g·kg^–1)	0.26± 0.08bA	0.24± 0.06bA	0.22± 0.04bB	0.28± 0.03bA	0.31± 0.04abA	0.31± 0.03bA	0.44± 0.03aA	0.42± 0.01aAB	0.33± 0.05aB	0.36± 0.07abA	0.33± 0.10abA	0.32± 0.08bA
TK/(g·kg^–1)	16.89± 2.04aA	17.97± 1.37aA	17.28± 0.72aA	17.87± 2.23aA	17.45± 3.68aA	17.83± 2.52aA	16.05± 0.85aA	14.98± 1.25aA	14.52± 0.29aA	15.38± 1.66aA	15.63± 1.82aA	14.64± 2.50aA
AN/(mg·kg^–1)	91.85± 4.83cA	80.11± 13.07bA	55.18± 13.17bB	126.46± 8.19bA	109.24± 1.84aB	92.64± 7.07aC	153.08± 2.53aA	124.67± 2.15aB	95.17± 24.04aC	122.83± 6.85bA	91.94± 10.73bB	75.47± 15.07abB
AP/(mg·kg^–1)	0.97± 0.30aA	0.85± 0.04aA	0.70± 0.21aA	1.22± 0.14aA	0.86± 0.35aA	0.78± 0.27aA	1.53± 0.38aA	1.19± 0.33aAB	0.66± 0.06aB	1.02± 0.20aA	0.77± 0.29aA	0.80± 0.39aA
AK /(mg·kg^–1)	47.88± 6.23bA	38.44± 1.51aB	33.29± 2.62aB	52.62± 2.99abA	39.87± 8.30aA	38.63± 9.24aA	67.99± 5.62aA	45.90± 4.92aB	40.24± 3.52aB	59.91± 12.52abA	42.96± 9.78aAB	35.53± 10.40aB
pH	4.89± 0.09bA	4.94± 0.13bA	5.03± 0.09aA	4.96± 0.08abA	4.95± 0.04bA	5.03± 0.01aA	5.15± 0.12aA	5.14± 0.05aA	5.08± 0.12aA	5.05± 0.04abA	4.99± 0.06abA	4.99± 0.03aA
AFe/(mg·kg^–1)	20.34± 3.9aA	16.13± 5.57aA	7.66± 2.41bB	24.49± 3.70aA	20.36± 3.48aA	19.75± 4.69aA	21.82± 1.76aB	17.7± 1.63aB	16.6± 1.06abA	20.09± 4.44aA	13.53± 3.58aAB	10.28± 3.89bcB
SOC/(g·kg^–1)	9.5± 0.84bA	7.07± 1.33bB	4.36± 0.97cC	15.19± 0.93aA	11.58± 1.10aB	10.29± 1.59aB	16.4± 2.26aA	11.98± 0.09abB	9.18± 0.65bB	14.53± 2.27aA	10.27± 3.13abAB	7.55± 1.30bB
C∶N	10.05± 0.12bA	7.09± 1.45bB	6.08± 1.25bB	13.32± 0.59abA	11.15± 1.15aAB	11.33± 1.13aB	13.96± 2.52aA	13.1± 0.91aA	12.2± 0.60aA	11.85± 1.09abA	10.57± 2.74abA	9.60± 2.02aA
N∶P	3.73± 0.83aA	4.38± 0.93aA	3.36± 0.78aA	4.08± 0.67aA	3.46± 0.61abA	2.96± 0.59aA	2.68± 0.24aA	2.34± 0.16bA	2.31± 0.26aA	3.52± 0.96aA	3.15± 1.09abA	2.59± 0.60aA
C∶P	37.41± 7.93aA	31.96± 12.14aA	20.51± 6.26aA	54.16± 7.44aA	38.58± 7.91aB	33.52± 7.60aB	37.45± 8.11aA	25.28± 0.88aB	24.75± 3.83aB	42.45± 15.73aA	35.29± 21.46aA	25.52± 11.19aA

Table 3

Table 4

Soil enzyme activity and microbial metabolism of different understory enrichment planting modes and soil layers"

指标 Index	M0			M1			M2			M3
指标 Index	0~20 cm	20~40 cm	40~60 cm	0~20 cm	20~40 cm	40~60 cm	0~20 cm	20~40 cm	40~60 cm	0~20 cm	20~40 cm	40~60 cm
SUC	3.11± 1.18bA	1.25± 0.94cB	0.58± 0.33bC	3.31± 0.21aA	2.40± 0.97aB	1.61± 0.79aC	2.59± 1.08cA	1.25± 0.83cB	0.97± 0.92aB	2.17± 0.83dA	1.53± 0.81bB	1.19± 0.61abC
GLU	0.74± 0.12cA	0.37± 0.03cB	0.21± 0.05cB	1.02± 0.12aA	0.84± 0.24aB	0.78± 0.26aB	0.95± 0.09abA	0.56± 0.04bB	0.46± 0.15bC	0.88± 0.24bcA	0.64± 0.14bB	0.54± 0.08bB
CEL	0.15± 0.15bB	0.32± 0.33aA	0.26± 0.17aAB	0.33± 0.07aA	0.15± 0.07bB	0.13± 0.06bB	0.28± 0.10aA	0.13± 0.07bB	0.07± 0.03bB	0.28± 0.16aA	0.17± 0.09bB	0.11± 0.04bB
URE	0.42± 0.14aA	0.32± 0.04bA	0.13± 0.02bB	0.49± 0.04aA	0.45± 0.05aA	0.40± 0.11aA	0.47± 0.05aA	0.34± 0.06bB	0.34± 0.10aB	0.54± 0.00aA	0.45± 0.02aAB	0.36± 0.04aB
ACP	1.85± 0.24aA	1.84± 0.21aA	1.22± 0.22cA	1.56± 0.24cA	1.68± 0.22bcAB	1.67± 0.19aB	1.66± 0.16bcA	1.60± 0.26cA	1.49± 0.11bB	1.68± 0.20bA	1.73± 0.11abAB	1.60± 0.10aB
CAT	2.58± 0.41aA	1.87± 0.20bB	1.53± 0.47cB	2.18± 0.10bA	1.96± 0.15bbA	1.96± 0.33bA	2.50± 0.38aA	2.21± 0.68aB	2.11± 0.63aB	2.52± 0.45aA	2.16± 0.56aB	1.80± 0.70bC
RAC	0.68± 0.19aA	0.37± 0.34aA	–0.17± 0.61aA	0.78± 0.02bA	0.69± 0.12aA	0.54± 0.20aA	0.69± 0.17aA	0.43± 0.23aA	0.12± 0.47aA	0.68± 0.12aA	0.51± 0.23aA	0.42± 0.17aA
RAN	0.55± 0.35bA	–0.83± 0.22aA	–2.65± 0.92bB	–0.37± 0.05aA	–0.48± 0.11aA	–0.66± 0.29aA	–0.45± 0.13aA	–0.81± 0.20aA	–1.04± 0.47aA	–0.37± 0.13aA	–0.55± 0.11aA	–0.93± 0.22aB
RAP	0.36± 0.12aA	0.44± 0.14aA	0.20± 0.21aA	0.23± 0.08aA	0.31± 0.10aA	0.35± 0.10aA	0.30± 0.09aA	0.35± 0.14aA	0.36± 0.12aA	0.30± 0.06aA	0.37± 0.09aA	0.36± 0.02aA
E_C∶E_N	–1.65± 1.03aB	–0.53± 0.50aAB	0.01± 0.26aA	–2.18± 0.22aA	–1.53± 0.67acA	–1.06± 0.89aA	–1.96± 0.97aA	–0.60± 0.46aA	–0.35± 0.82aA	–2.03± 0.94aA	–1.01± 0.30aAB	–0.50± 0.35aB
E_C∶E_P	2.26± 1.55aA	1.07± 1.16aA	1.09± 1.26aA	3.77± 1.19aA	2.52± 1.3aA	1.74± 1.10aA	2.55± 1.30aA	1.48± 1.13aA	0.70± 1.85aA	2.31± 0.72aA	1.51± 0.93aA	1.18± 0.52aA
E_N∶E_P	–1.54± 0.65aA	–1.92± 0.39abA	3.75± 3.97aA	–1.77± 0.68aA	–1.60± 0.30aA	–1.92± 0.76aA	–1.55± 0.41aA	–2.41± 0.40bAB	–2.83± 0.85aB	–1.24± 0.35aA	–1.47± 0.06aA	–2.59± 0.55aB
RLC	3.83± 0.78aA	1.20± 1.26aA	2.66± 1.38aA	4.41± 0.89aA	2.96± 1.47aB	2.05± 1.39aB	3.08± 1.59aA	1.60± 1.23aB	1.30± 1.58aB	3.10± 1.07aA	1.82± 1.11aB	1.29± 0.61aB
N-P limitation	127.35± 10.34aA	118.05± 5.02aA	135.35± 13.48aA	121.52± 11.36aA	122.47± 5.23aA	119.24± 8.29aA	113.70± 6.54bA	112.89± 3.50bAB	107.55± 9.64aB	129.97± 8.98aA	124.16± 1.02aA	111.66± 4.62aB

Table 4

Table 5

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Fig.2

Fig.3

Fig.4

Fig.5

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项目 Items	M1			M2			M3			M0
样地编号 Plot No.	M1-1	M1-2	M1-3	M2-1	M2-2	M2-3	M3-1	M3-2	M3-3	M0-1	M0-2	M0-3
树种比例 Tree species composition ratio	3∶7∶ 0∶0	3∶7∶ 0∶0	3∶7∶ 0∶0	3∶5∶ 2∶0	3∶6∶ 1∶0	3∶6∶ 1∶0	3∶4∶ 1∶2	3∶2∶ 4∶1	3∶3∶ 3∶1	10∶0∶ 0∶0	10∶0∶ 0∶0	10∶0∶ 0∶0
坡位 Slope position	中坡Mid-slope			中坡Mid-slope			中坡Mid-slope			中坡Mid-slope
坡向 Slope aspect	西南Southwest			西南Southwest			西南Southwest			西南Southwest
海拔 Elevation/m	216.11			160.68			176.61			163.94
杉木密度 Density of C. lanceolata/ (trees·hm^–2)	375			350			367			1675
杉木平均胸径 Mean DBH of C. lanceolata/cm	23.16			25.20			22.07			18.46
杉木平均高 Mean tree height of C. lanceolata/m	21.42			24.83			21.12			17.01
闽楠平均胸径 Mean DBH of P. bournei/cm	4.36			5.82			5.25
闽楠平均高 Mean tree height of P. bournei/m	5.58			6.64			6.37
红豆杉平均胸径 Mean DBH of T. wallichiana var. chinensis/cm				5.99			5.88
红豆杉平均高 Mean tree height of T. wallichiana var. chinensis/m				5.08			4.82
木荷平均胸径 Mean DBH of S. superba/cm							4.49
木荷平均高 Mean tree height of S. superba/m							5.86

指标Index	缩写 Abbreviation	测定方法 Determination methods
全氮 Total nitrogen	TN	半微量凯氏定氮法Semi-micro Kjeldahl method
全磷 Total phosphorus	TP	消煮?钒钼酸铵比色法Digestion-ammonium vanadomolybdate colorimetry
全钾 Total potassium	TK	氢氧化钠熔解?火焰分光光度法Sodium hydroxide fusion-flame spectrophotometry
碱解氮 Available nitrogen	AN	碱解蒸馏法Alkali hydrolysis-distillation method
有效磷 Available phosphorus	AP	碳酸氢钠浸提?钼锑抗比色法Sodium bicarbonate extraction-molybdenum antimony anti colorimetry
速效钾 Available potassium	AK	乙酸铵浸提?火焰分光光度法Ammonium acetate extraction-flame spectrophotometry
pH值 pH value	pH	水浸提电位法（水土比2.5∶1）Water extraction potentiometry （water-soil ratio 2.5∶1）
有机质 Organic matter	SOM	重铬酸钾氧化?外加热法Potassium dichromate oxidation-external heating method
有效铁 Available iron	AFe	DTPA浸提?原子吸收分光光度法DTPA extraction-atomic absorption spectrophotometry
土壤蔗糖酶 Soil sucrase	SUC	3,5-二硝基水杨酸比色法3,5-dinitrosalicylic acid colorimetry
葡萄糖苷酶 β-Glucosidase	GLU	消煮?钒钼酸铵比色法Digestion-ammonium vanadomolybdate colorimetry
纤维素酶 Cellulase	CEL	3,5-二硝基水杨酸比色法3,5-dinitrosalicylic acid colorimetry
酸性磷酸酶 Acid phosphatase	ACP	磷酸苯二钠比色法Disodium phenyl phosphate colorimetry
脲酶 Urease	URE	苯酚?次氯酸钠比色法Phenol-sodium hypochlorite colorimetry
过氧化氢酶 Catalase	CAT	高锰酸钾滴定法Potassium permanganate titration

林下补植模式 Understory enrichment planting models	土层深度 Soil layers/cm	Shannon	Simpson	Chao	Ace	Good’s_coverage
M0	0~20	8.47±0.11bA	0.99±0.00aA	2 732.87±129.22aA	2 977.44±146.10aA	0.98±0.00aA
	20~40	8.29±0.09bAB	0.99±0.00aA	2 582.40±56.97aA	2 799.72±90.72aA	0.98±0.00aA
	40~60	8.15±0.16bB	0.99±0.00aA	2 554.00±145.01aA	2 768.72±145.01aB	0.99±0.00aA
M1	0~20	8.58±0.13abA	0.99±0.00aA	2 598.92±149.10abA	2 681.04±149.10bA	0.99±0.00aA
	20~40	8.47±0.25bB	0.99±0.00aA	2 446.58±117.69abAB	2 518.04±118.00bAB	0.99±0.00aA
	40~60	8.27±0.20bB	0.99±0.00aA	2 243.27±202.61bB	2 308.59±212.31bB	0.99±0.00aA
M2	0~20	8.76±0.23aA	0.99±0.00aA	2 672.73±199.43abA	2 776.64±206.03bA	0.98±0.00aA
	20~40	8.61±0.12aA	0.99±0.00aA	2 552.18±118.48abA	2 645.41±124.25abA	0.98±0.00aA
	40~60	8.64±0.16aA	0.99±0.00aA	2 578.67±111.65aA	2 685.35±114.25aA	0.98±0.00aA
M3	0~20	8.49±0.14bA	0.99±0.00aA	2 607.91±198.17bA	2 712.45±198.40bA	0.98±0.00aA
	20~40	8.33±0.17bA	0.99±0.00aA	2 581.56±188.81bA	2 688.11±193.05bA	0.98±0.00aA
	40~60	8.51±0.15bA	0.99±0.00bA	2 525.03±133.22abA	2 623.19±138.30aA	0.98±0.00aA

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Effects of Understory Enrichment Planting Modes in Pure Chinese fir Forests on Soil Quality and Microbial Communities

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