Seasonal dynamics of functional diversity of soil microbial communities in Castanopsis kawakamii forest gaps
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摘要:目的 林窗作为森林生态系统中的小尺度干扰，对森林土壤养分循环与微生物群落功能多样性维持起着重要作用。明确不同林窗大小土壤养分和微生物群落功能多样性及其季节动态响应机制，有助于改善格氏栲林土壤生态环境。方法 以格氏栲天然林林窗为对象，采用Biolog微平板法，研究不同林窗大小土壤理化性质、碳源利用能力和代谢特征的季节动态变化规律。结果 （1）林窗生长季土壤温度、速效钾含量显著高于非生长季，pH值、速效磷含量显著低于非生长季。林窗形成促进了土壤pH值升高与速效钾积累，加速了微生物对碱解氮与速效磷的吸收和利用。（2）林窗生长季土壤微生物平均颜色变化率在培养的中后期显著高于非生长季。林窗内土壤微生物在生长季中主要利用碳源为羧酸、多聚物和氨基酸类，在非生长季中主要利用碳源为氨基酸类和羧酸类。（3）林窗土壤微生物功能多样性指数中，生长季Shannon-Wiener和Pielou指数显著高于非生长季，Simpson指数显著低于非生长季。林窗面积对生长季土壤微生物群落功能多样性指数影响不明显。非生长季小林窗Simpson指数最高，中林窗Shannon-Wiener指数和Pielou指数最高。（4）广义线性模型表明，林窗生长季的土壤温度和土壤速效钾含量升高有利于土壤微生物常见种优势度升高；非生长季土壤碱解氮含量降低，速效钾含量升高有利于微生物功能多样性升高。结论 生长季林窗内土壤微生物群落功能多样性保持较高水平，土壤养分循环效率高；非生长季非林窗土壤环境有利于土壤微生物生长代谢，维持土壤生态系统功能。林窗形成导致土壤温度与速效养分含量的异质性是影响土壤微生物群落代谢特征和功能多样性的主要因素。
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关键词:
- 林窗 /
- 土壤微生物 /
- 功能多样性 /
- Biolog微平板法 /
- 格氏栲林
Abstract:Objective As a small-scale disturbance in the forest ecosystem, the forest gap plays an important role in nutrient cycling and functional diversity of microbial community in the soil of forest. Clarifying the seasonal dynamic response mechanism of soil nutrients and functional diversity of microbial community to the forest size will help to improve the soil ecological environment of Castanopsis kawakamii forests.Method The gaps of C. kawakamii natural forests were used as research objects, and the Biolog micro-plate method was used, combining the regularity of soil physical and chemical properties, carbon source utilization capacity and metabolic characteristics with different size of forest gaps during growing season and non-growing season.Result (1) The soil temperature (ST), available potassium (AK) content in the growing season of the gaps were significantly higher than those in the non-growing season, and pH value, available phosphorus (AP) content were significantly lower than those in the non-growing season. The formation of forest gaps promoted the increase of soil pH value and the accumulation of available potassium (AK), and accelerated the absorption and utilization of alkali-hydrolyzable nitrogen (HN) and available phosphorus (AP) by microorganisms. (2) The average well color development (AWCD) of soil microorganisms in the growing season of the forest gaps was significantly higher than that in the non-growing season in the middle and late cultivation periods. The soil microorganisms of the forest gaps mainly used carboxylic acid, polymer, and amino acid carbon sources in the growing season, and mainly used the carbon sources as amino acid and carboxylic acid carbon sources in the non-growing season. (3) In soil microbial functional diversity indexes of forest gaps, Shannon-Wiener and Pielou indices in the growing season of the forest gaps were significantly higher than those in the non-growing season, and the Simpson index was significantly lower than in the non-growing season. Gap size had no significant effect on the functional diversity index of microbial community during the growing season. The Simpson index of the soil microbial community in the non-growing small forest window was the highest. The Shannon-Wiener index and Pielou index of the medium gaps were the highest. (4) Generalized linear model (GLM) analysis showed that increasing soil temperature (ST) and soil available potassium (AK) in the growing season of the forest gaps were conducive to increasing the dominance of common soil microorganism species. The reduction of soil alkaline nitrogen (HN) content and the increase of available potassium (AK) content in non-growing seasons were conducive to the increase of microbial diversity and uniformity.Conclusion Functionaldiversity of soil microbial community of the forest gaps during the growing season is maintained at a high level, and the soil nutrient cycling efficiency is high. The soil environment of non-gaps during the non-growing season is conducive to the growth and metabolism of soil microorganisms and maintains the function of the soil ecosystem. Heterogeneity of soil temperature and available nutrient content caused by the formation of forest gaps is the main factor affecting the metabolic characteristics and functional diversity of soil microbial communities. -
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图 1 不同林窗内生长季与非生长季土壤微生物AWCD值
LG1为生长季大林窗；MG1为生长季中林窗；SG1为生长季小林窗；NG1为生长季对照；LG2为非生长季大林窗；MG2为非生长季中林窗；SG2为非生长季小林窗；NG2为非生长季对照。同一列数据上不同字母表示差异显著（P < 0.05）。下同。LG1, large forest gap in growing season; MG1, medium forest gap in growing season; SG1, small forest gap in growing season; NG1, control in growing season; LG2, large forest gap in non-growing season; MG2, medium forest gap in non-growing season; SG2, small forest gap in non-growing season; NG2, control in non-growing season. Different letters above the same column data indicate significant differences （P < 0.05）.The same below.
Figure 1. Soil microbial AWCD values of different forest gaps in growing season and non-growing season
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图 2 不同林窗内生长季与非生长季土壤微生物碳源利用能力
Figure 2. Utilization of carbon sources by soil microorganisms of different forest gaps in growing season and non-growing season
表 1 林窗研究样地概况
Table 1 General situation of the research plots of the forest gaps
林窗 Forest gap GS/m2 SHC/% LG 206.17 ± 4.53 0.85 MG 73.15 ± 1.93 0.67 SG 33.49 ± 2.48 0.50 NG 100 ± 0 0.77 注：LG为大林窗；MG为中林窗；SG为小林窗；NG为对照；GS为林窗面积；SHC为灌草覆盖度。数据为均值 ± 标准差，n = 3，下同。Notes: LG, large gap; MG, medium gap; SG, small gap; NG, control; GS, forest gap size; SHC, shrub and herb coverage. The data are mean ± standard deviation, n = 3. The same below. 
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表 2 林窗土壤特性的双因素方差分析
Table 2 Two-way ANOVA analysis of soil characteristics in different growing seasons and gap size
环境因子 Environmental factor 因子 Factor 自由度 df F P ST 季节 Season 1 2 707.332 < 0.001*** 林窗面积 Forest gap size 3 0.978 0.428 季节 × 林窗面积 Season × forest gap size 3 0.325 0.807 SWC 季节 Season 1 2.246 0.153 林窗面积 Forest gap size 3 0.298 0.826 季节 × 林窗面积 Season × forest gap size 3 0.077 0.972 pH 季节 Season 1 45.334 < 0.001*** 林窗面积 Forest gap size 3 5.256 0.010** 季节 × 林窗面积 Season × forest gap size 3 2.132 0.136 HN 季节 Season 1 3.219 0.092 林窗面积 Forest gap size 3 1.256 0.323 季节 × 林窗面积 Season × forest gap size 3 0.586 0.633 AK 季节 Season 1 84.533 < 0.001*** 林窗面积 Forest gap size 3 9.146 0.001** 季节 × 林窗面积 Season × forest gap size 3 0.733 0.548 AP 季节 Season 1 28.392 < 0.001*** 林窗面积 Forest gap size 3 18.916 < 0.001*** 季节 × 林窗面积 Season × forest gap size 3 7.052 0.003** SOC 季节 Season 1 1.439 0.248 林窗面积 Forest gap size 3 2.032 0.150 季节 × 林窗面积 Season × forest gap size 3 1.521 0.247 注：ST为土壤温度；SWC为土壤含水量；HN为碱解氮；AK为速效钾；AP为速效磷；SOC为有机碳，下同。*代表差异显著（P < 0.05）；**代表差异极显著（P < 0.01）；***代表差异极显著（P < 0.001），下同。Notes: ST, soil temperature; SWC, soil water content; HN, alkali-hydrolyzable nitrogen; AK, available potassium; AP, available phosphorus; SOC, soil organic carbon. * represents significant difference （P < 0.05）; ** represents extremely significant difference （P < 0.01）; *** represents extremely significant difference （P < 0.001）. The same below.
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表 3 不同面积林窗生长季与非生长季土壤特性
Table 3 Soil characteristics of forest gaps with different size in growing season and non-growing season
季节 Season 林窗
Forest gapST/℃ pH HN/（mg∙kg− 1） AK/（mg∙kg− 1） AP/（mg∙kg− 1） 生长季 Growing season LG 25.9 ± 0.65a 3.4 ± 0.08b 212.05 ± 34.94ab 120.89 ± 20.25a 5.89 ± 2.72d MG 25.7 ± 0.37a 3.25 ± 0.01cd 215.32 ± 7.09ab 86.51 ± 6.13ab 10.61 ± 0.30c SG 25.64 ± 0.14a 3.24 ± 0.08cd 182.57 ± 20.6b 93.56 ± 8.14b 12.57 ± 1.00abc NG 25.6 ± 0.59a 3.21 ± 0.03d 235.79 ± 12.76ab 94.85 ± 2.55b 12.64 ± 1.03abc 非生长季 Non-growing season LG 12.94 ± 0.36b 3.54 ± 0.09a 222.69 ± 55.27ab 75.46 ± 12.31cd 12.01 ± 0.01bc MG 12.63 ± 0.33b 3.37 ± 0.1bc 231.7 ± 49.14ab 56.01 ± 2.88e 12.08 ± 0.27bc SG 12.63 ± 0.07b 3.45 ± 0.1ab 234.15 ± 15.6ab 61.2 ± 6.91de 13.2 ± 0.64ab NG 12.02 ± 1.38b 3.53 ± 0.01a 248.07 ± 11.26a 59.89 ± 0.36de 14.33 ± 0.59a 注：同一列数据后不同字母表示差异显著（P < 0.05），下同。Notes: different letters in the same column indicate significant difference（P < 0.05）. The same below.
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表 4 林窗土壤平均颜色变化率值的双因素方差分析
Table 4 Two-way ANOVA on soil average well color development（AWCD）value of forest gaps
组间因子
Intergroup factordf F P 组内因子
Intragroup factor自由度
dfF P 季节 Season 1 250.338 < 0.001*** 时间 Time 8 989.012 < 0.001*** 林窗面积 Forest gap size 3 0.455 0.717 时间 × 季节 Time × season 8 224.237 < 0.001*** 季节 × 林窗面积
Season × forest gap size3 0.111 0.952 时间 × 林窗面积 Time × forest gap size 24 0.601 0.926 时间 × 季节 × 林窗面积
Time × season × forest gap size24 0.263 1.000
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表 5 不同生长季节与林窗面积的微生物功能多样性双因素分析结果
Table 5 Results of two-way ANOVA of functional diversity of microorganisms in different growing seasons and forest gap size
多样性指数 Diversity index 因子 Factor df F P Simpson 指数
Simpson index季节 Season 1 8.773 0.009** 林窗面积 Forest gap size 3 1.684 0.210 季节 × 林窗面积 Season × forest gap size 3 1.557 0.239 Shannon-Wiener指数
Shannon-Wiener index季节 Season 1 350.491 < 0.001*** 林窗面积 Forest gap size 3 4.298 0.021* 季节 × 林窗面积 Season × forest gap size 3 4.403 0.019* Pielou 指数
Pielou index季节 Season 1 350.665 < 0.001*** 林窗面积 Forest gap size 3 4.281 0.021* 季节 × 林窗面积 Season × forest gap size 3 4.406 0.019*
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表 6 不同面积林窗土壤微生物功能多样性
Table 6 Functional diversity of soil microorganisms of forest gaps in different size
季节
Season林窗
Forest gapSimpson指数
Simpson indexShannon-Wiener指数
Shannon-Wiener indexPielou指数
Pielou index生长季 Growing season LG 0.984 4 ± 0.000 2b 4.88 ± 0.02a 0.985 7 ± 0.004 3a MG 0.984 2 ± 0.000 3b 4.86 ± 0.01a 0.980 6 ± 0.002 2a SG 0.984 8 ± 0.000 3b 4.9 ± 0.01a 0.988 3 ± 0.001 4a NG 0.983 9 ± 0.001 1b 4.84 ± 0.05a 0.977 8 ± 0.010 9a 非生长季 Non-growing season LG 1.011 7 ± 0.006 7ab 4.49 ± 0.02c 0.906 5 ± 0.003 4c MG 0.996 7 ± 0.007b 4.63 ± 0.03b 0.935 3 ± 0.005 6b SG 1.031 0 ± 0.051 3a 4.54 ± 0.09c 0.917 1 ± 0.017 8c NG 0.987 0 ± 0.002b 4.48 ± 0.05c 0.904 0 ± 0.010 9c
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表 7 土壤微生物功能多样性与环境因子间逐步回归结果
Table 7 Results of stepwise regression between functional diversity of soil microorganisms and environmental factors
季节 Season 多样性指数 Diversity index 环境因子 Environmental factor AIC 生长季 Growing season Simpson指数 Simpson index ST + SWC + pH + HN + AK + AP + SOC + GA − 423.13 ST + SWC + pH + AK + AP + SOC + GA − 424.92 ST + pH + AK + AP + SOC + GA − 426.74 ST + pH + AK + AP + SOC − 428.27 Shannon-Wiener指数 Shannon-Wiener index ST + SWC + pH + HN + AK + AP + SOC + GA − 181.88 ST + SWC + pH + HN + AK + AP + SOC − 183.66 ST + SWC + pH + HN + AP + SOC − 185.52 ST + SWC + pH + HN + SOC − 187.35 ST + SWC + HN + SOC − 188.68 ST + HN + SOC − 189.73 ST + SOC − 191.15 Pielou指数 Pielou index ST + SWC + pH + HN + AK + AP + SOC + GA − 277.94 ST + SWC + pH + HN + AK + AP + SOC − 279.72 ST + SWC + pH + HN + AP + SOC − 281.57 ST + SWC + pH + HN + SOC − 283.41 ST + SWC + HN + SOC 284.73 ST + HN + SOC − 285.79 ST + SOC − 287.21 非生长季 Non-growing season Simpson指数 Simpson index ST + SWC + pH + HN + AK + AP + SOC + GA − 284.55 ST + SWC + pH + HN + AK + SOC + GA − 286.52 ST + SWC + pH + HN + AK + SOC − 288.46 ST + SWC + HN + AK + SOC − 289.41 Shannon-Wiener指数 Shannon-Wiener index ST + SWC + pH + HN + AK + AP + SOC + GA − 84.2 SWC + pH + HN + AK + AP + SOC + GA − 86.11 SWC + pH + HN + AK + AP + GA − 87.56 Pielou指数 Pielou index ST + SWC + pH + HN + AK + AP + SOC + GA − 180.22 SWC + pH + HN + AK + AP + SOC + GA − 182.13 SWC + pH + HN + AK + AP + GA − 183.58 注：AIC为赤池信息准则。下同。Notes: AIC，akaike information criterion.The same below.
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表 8 土壤微生物功能多样性与环境因子间GLM分析
Table 8 GLM analysis of the relationship between functional diversity of soil microorganisms and environmental factors
季节
Season多样性指数
Diversity index环境因子
Environmental factorAIC β SE t P 生长季
Growing seasonSimpson指数
Simpson indexST − 428.27 1.25 × 10− 3 2.76 × 10− 4 4.525 0 0.000 9*** AK 3.12 × 10− 5 1.36 × 10− 5 2.298 0 0.038 6* Shannon-Wiener指数
Shannon-Wiener index— − 191.15 — — — — Pielou指数
Pielou index— − 287.21 — — — — 非生长季
Non-growing seasonSimpson指数
Simpson indexST − 289.41 3.47 × 10− 3 1.52 × 10− 3 2.289 0 0.031 2* SWC 2.46 × 10− 3 8.92 × 10− 4 2.754 0 0.011 0* HN 2.03 × 10− 4 3.96 × 10− 5 5.134 0 < 0.000 1*** AK − 7.74 × 10− 4 1.61 × 10− 4 − 4.808 0 < 0.000 1*** SOC − 1.44 × 10− 2 4.16 × 10− 3 − 3.471 0 0.002 0** Shannon-Wiener指数
Shannon-Wiener indexSWC − 87.56 − 0.089 1 0.025 9 − 3.455 0 0.002 2** AK 0.016 4 0.005 9 2.784 0 0.010 6* GA − 0.002 8 0.000 9 − 3.049 0 0.005 7** Pielou指数
Pielou indexSWC − 183.58 − 0.018 0 0.005 2 − 3.443 0 0.002 2** AK 0.003 3 0.001 2 2.784 0 0.010 6* GA − 0.000 6 0.000 2 − 3.050 0 0.005 7**
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