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毛竹(Phyllostachys edulis (Carr.) H. de Lehaie)是我国分布最广、面积最大和经济价值最高的竹种。第八次全国森林资源清查(2009—2013年)数据表明,我国竹林面积达601万hm2,其中毛竹林面积为443万hm2,占竹林总面积的73%,并且毛竹林面积还将继续增加[1-2]。毛竹具有生长速度快、经济效益好等优点,是山区农民的重要收入来源。为了获得更大的经济效益,近年来大面积马尾松(Pinus massoniana Lamb.)林以及杉木(Cunninghamia lanceolata (Lamb.) Hook.)林等被毛竹林所替代。然而,已有研究表明,天然林受到毛竹入侵或人为改造成毛竹林后,毛竹易形成单优群落,导致生物多样性明显下降[3-4]。同时,毛竹叶能释放化感物质抑制林下植被的生长,引起植物群落结构及物种多样性的改变[5]。近年来,毛竹林生物多样性降低、生产力下降等现象开始引起人们的广泛关注。
土壤微生物是土壤养分循环的核心驱动力,在土壤形成和发育过程以及维护土壤生态系统稳定性方面发挥着至关重要的作用[6]。地上植被对土壤微生物群落结构形成起着至关重要的作用[7],植物种类改变会引起凋落物种类、根系分泌物以及养分吸收特性等方面的改变,从而导致土壤物理、化学和生物学性质发生改变[8-9]。Chang等[10]研究表明,毛竹入侵日本柳杉(Cryptomeria japonica (L. f.) D. Don)林后显著改变了土壤微生物活性及群落结构。徐秋芳等[11]对比了马尾松林和毛竹林土壤微生物特性,发现马尾松林土壤微生物量较高,多样性更为丰富;而孙棣棣等[12]发现天然马尾松林改种毛竹林后,土壤微生物量和多样性变化不大,但微生物群落结构发生了变化。目前,关于马尾松林改造成毛竹林以及长期粗放经营后毛竹林土壤微生物量及群落结构的演变趋势的研究结果尚不统一,而导致土壤微生物群落发生改变的关键控制因子也并不清楚,开展长期粗放经营毛竹林土壤微生物群落演变趋势及其控制因素的研究,有助于深入了解土壤微生物群落对环境因子的响应机理,对于保护森林生态系统稳定性具有重要意义。
本研究在浙江安吉选取4个不同粗放经营年限的毛竹林,同时选取附近的马尾松林作为对照,通过分析土壤微生物生物量及群落结构的演变趋势,筛选影响土壤微生物群落的关键环境因子,揭示土壤微生物群落对毛竹林长期粗放经营的响应机理,为评估毛竹林生态系统稳定性,促进毛竹林可持续发展提供理论依据。
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研究结果表明,毛竹林土壤含水量均显著低于马尾松林(P<0.05),其中18 a毛竹林土壤水分含量最低(表 1)。毛竹林在粗放经营初期,土壤pH值呈现出逐渐升高的趋势,9 a毛竹林土壤pH值显著高于天然马尾松林和其它年份毛竹林(P<0.05),但长期粗放经营后土壤pH值又逐渐下降,18 a毛竹林与马尾松林没有显著差异。在改种毛竹的初始几年,土壤有机质、有效磷、速效钾等含量迅速增长,5 a毛竹林土壤有机质、有效磷、速效钾含量均显著高于马尾松林(P<0.05),但随着粗放经营年限的延长土壤有机质及碱解氮及有效磷又呈现出下降趋势,其中18 a毛竹林土壤有机质含量显著低于马尾松林(P<0.05)。与其它养分不同的是,随着粗放经营年限的延长,土壤碱解氮含量呈现出下降趋势,18 a毛竹林土壤碱解氮含量显著低于马尾松林(P<0.05)。
表 1 不同粗放经营年限毛竹林土壤理化性质
Table 1. Soil physic-chemical properties under bamboo stands with different culture histories
处理
Treatment含水量
Soil Moisture/%pH
(H2O)有机质
Organic matter/(g·kg-1)碱解氮
Alkalytic N/(mg·kg-1)有效磷
Available P/(mg·kg-11)速效钾
Available K/(mg·kg-1)MP 23.71±0.60a 4.51±0.05c 43.28±2.18b 115.21±21.88a 0.64±0.15b 40.0±7.2b 5a 18.91±1.45b 4.61±0.08bc 57.84±1.60a 102.20±3.9ab 1.97±0.41a 67.0±8.2a 9a 20.42±1.00b 4.88±0.15a 35.75±1.25bc 93.12±3.7ab 1.95±0.43a 41.0±5.0b 15a 19.08±1.99b 4.71±0.12b 41.87±2.13bc 93.95±20.4ab 1.49±0.85ab 45.7±3.2b 18a 15.51±2.13c 4.63±0.06bc 35.01±4.22c 70.15±5.1b 1.23±0.25ab 44.7±5.9b 注: 数据为3个重复的平均值及标准差(mean±SD);同列中不同字母表示处理间差异达5%显著水平。MP表示马尾松林;5 a、9 a、15 a、18 a表示毛竹林粗放经营年限为5年、9年、15年、18年。下同。
Note: The data in the table are the means of 3 replicates and the standard error (mean±SD). Different letters in the same column indicate significant difference at 5% level. MP indicates masson pine. 5 a、9 a、15 a、18 a indicate moso bamboo stands with 5, 9, 15, 18 years of extensive management age, respectively. The same below. -
利用PLFA分析方法表征不同样地土壤微生物生物量及其群落,结果表明,马尾松林土壤微生物总PLFA含量以及细菌、真菌、放线菌等PLFA含量均显著高于毛竹林(P<0.05),而不同粗放经营年限毛竹林土壤总PLFA含量之间没有显著差异(表 2)。对于细菌,革兰氏阳性细菌PLFA含量变化趋势与总细菌相同,而15 a毛竹林土壤革兰氏阴性细菌PLFA含量与马尾松林没有显著差异。
表 2 不同粗放经营年限毛竹林土壤PLFA含量
Table 2. Soil PLFA contents under bamboo stands with different culture histories
nmol·g-1 处理
Treatment总PLFA
Total PLFA细菌
Bacteria革兰氏阳性细菌
G+ bacteria革兰氏阴性细菌
G- bacteria真菌
Fungi放线菌
Actinomycete原生动物
ProtozaMP 113.01±19.98a 63.97±13.92a 24.65±4.76a 22.55±5.84a 8.69±2.09a 11.28±4.74a 0.53±0.14a 5a 48.05±16.60b 28.74±10.20b 9.81±3.34b 11.30±4.49b 4.53±1.77b 5.75±1.96b 0.24±0.05b 9a 43.74±11.67b 26.86±7.48b 9.42±2.80b 10.53±2.91b 3.93±1.03b 4.30±2.21b 0.31±0.18b 15a 58.82±15.49b 34.87±9.72b 12.63±3.47b 16.10±5.01ab 5.36±1.29b 6.50±1.34b 0.25±0.04b 18a 46.61±4.28b 25.56±2.45b 9.53±0.60b 9.98±1.10b 5.14±0.55b 5.57±0.52b 0.26±0.04b 将土壤微生物总PLFA含量作为因变量,土壤理化性质作为自变量,进行逐步回归分析,以揭示对不同种类微生物PLFA含量影响最显著的一组变量(表 3)。结果表明,引入的变量中,碱解氮和有效磷对土壤总PLFA含量以及细菌(包括革兰氏阳性和阴性细菌)、真菌、原生动物PLFA含量等影响显著,均达到1%极显著水平(P<0.01)。对于真菌PLFA含量的影响,有效磷的贡献大于碱解氮。与土壤微生物总PLFA含量以及其它类群微生物相比,土壤放线菌PLFA含量主要受碱解氮、土壤pH以及有机质含量这组变量影响,同样达到极显著水平(P<0.01)。
表 3 逐步回归分析
Table 3. Stepwise regression analysis
因变量
Dependence自变量
IndependenceF值
F value显著性
Significance总Total AN, AP 26.703 0.000 细菌Bacteria AN, AP 26.348 0.000 真菌Fungi AP, AN 13.043 0.001 放线菌Actinomycete AN, pH, OM 44.152 0.000 原生动物Protoza AN, AP 8.672 0.005 G-细菌G- bacteria AN, AP 17.134 0.000 G+细菌G+ bacteria AN, AP 23.552 0.000 注:AN、AP、OM分别为碱解氮、有效磷和土壤有机质。
Note: AN, AP, and OM indicate alkalytic N, available P and soil organic matter, respectively. -
马尾松改种毛竹后,在短期内土壤微生物物种丰富度没有发生明显变化,但是在长期尺度上有下降的趋势,15 a和18 a毛竹林土壤微生物物种丰富度显著低于马尾松林以及5 a和9 a毛竹林(P<0.05),Shannon多样性指数也有相似的变化趋势(表 4)。所有毛竹林土壤微生物群落Simpson多样性指数均显著低于马尾松林(P<0.05)。与Shannon多样性指数及Simpson多样性指数相反,长期粗放经营毛竹林土壤微生物群落Shannon均匀度及Simpson均匀度呈现出先下降再逐渐升高的趋势,15 a和18 a毛竹林均显著高于5 a及9 a毛竹林(P<0.05),其中18 a毛竹林Shannon均匀度和Simpson均匀度显著高于对照马尾松林(P<0.05)。
表 4 土壤微生物多样性指数
Table 4. Soil microbial diversity indexes
处理
Treatment物种丰富度
RichnessShannon多样性
Shannon diversitySimpson多样性
Simpson diversityShannon均匀度
Shannon evenessSimpson均匀度
Simpson evenessMP 45±6.81a 23.55±2.16a 17.03±1.65a 0.53±0.05bc 0.38±0.05bc 5a 44±6.66a 21.39±0.90ab 15.09±0.59b 0.49±0.07c 0.35±0.06c 9a 47±1.00a 21.36±1.12ab 14.72±0.74b 0.45±0.03c 0.31±0.02c 15a 35±3.21b 20.60±0.71b 15.17±0.34b 0.60±0.04ab 0.44±0.04ab 18a 32±0.58b 20.00±0.25b 15.28±0.54b 0.63±0.02a 0.48±0.03a 对马尾松林以及不同粗放经营年限毛竹林土壤微生物群落进行NMDS分析(图 1),马尾松林与毛竹林土壤微生物群落在NMDS第1排序轴上有明显的区分,而不同粗放经营年限毛竹林土壤微生物群落之间也存在差异,5 a、9 a毛竹林与15 a和18 a毛竹林在第2排序轴上区分明显。相似性检验结果表明,各处理土壤微生物群落之间存在显著差异(R=0.388 1, P=0.009)。典范对应分析结果表明,土壤含水量(F=5.401, P=0.001)、有效磷(F=2.494, P=0.022)、碱解氮(F=3.150, P=0.036)以及pH值(F=2.046, P=0.062)对土壤微生物群落结构的变异具有重要影响,合计解释了90.28%的变异量(图 2),其中土壤含水量、有效磷及碱解氮含量对土壤微生物群落结构变异影响显著(P<0.05)。
长期粗放经营毛竹林土壤微生物群落演变特征
Variation Patterns of Soil Microbial Community of Phyllostachys edulis Stands under Long-term Extensive Management
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摘要:
目的 通过分析土壤微生物生物量及群落结构的演变趋势,筛选影响土壤微生物群落的关键环境因子,揭示土壤微生物群落对毛竹林长期粗放经营的响应机理。 方法 选取不同粗放经营年限(5 a、9 a、15 a、18 a)毛竹林,以天然马尾松林(Masson pine,MP)作为对照,采用磷脂脂肪酸(Phospholipid fatty acids,PLFA)分析方法表征土壤微生物量及群落结构。 结果 结果表明,毛竹林土壤微生物总PLFA含量以及细菌、真菌、放线菌等PLFA含量均显著低于马尾松林(P<0.05),但不同经营年限毛竹林之间没有显著差异。土壤碱解氮及有效磷含量对土壤微生物总PLFA含量以及细菌、真菌、原生动物等PLFA含量影响显著(P<0.05),而土壤碱解氮、pH值以及有机质含量对放线菌PLFA含量影响显著(P<0.05)。长期粗放经营过程中毛竹林土壤微生物丰富度及多样性均呈逐渐下降趋势。非度量多维尺度转换排序(Non-metric multidimensional scaling,NMDS)分析结果表明,毛竹林土壤微生物群落结构与马尾松林有明显区分(R=0.388 1,P=0.009)。土壤含水量、碱解氮、有效磷以及pH值合计解释了90.28%的微生物群落结构变异量,其中土壤含水量、碱解氮、有效磷贡献显著(P<0.05)。 结论 长期粗放经营降低了毛竹林土壤微生物量,改变了群落结构,其生态风险还有待于进一步评估。 Abstract:Objective To investigate the effect of long-term extensive management on soil microbial communities and the related mechanisms. Method Moso bamboo (Phyllostachys edulis) stands with different extensive management time (5, 9, 15, and 18 years) were selected and compared with natural masson pine forest (Pinus massoniana Lamb.) as control. The phospholipid fatty acids (PLFA) were analyzed to indicate the soil microbial biomass and community composition. Result Results showed that the total soil microbial PLFA contents as well as bacterial, fungal, actinomycic and protozoal PLFA contents under moso bamboo stands were all significant lower than that of masson pine forest (P < 0.05), while no significant difference was found among bamboo stands. The total PLFA contents of soil microbial community as well as bacterial, fungal and protozoal PLFA contents were all significantly correlated with soil alkalytic nitrogen (AN) and available phosphorus (AP) contents (P < 0.05), while the actinomycic PLFA content correlated with AN, organic matter contents and pH significantly (P < 0.05). Soil microbial diversity, i.e. richness, Shannon diversity and Simpson diversity indexes, decreased progressively with the extensive management time of moso bamboo. According to the results of non-metric multidimensional scaling (NMDS), the soil microbial communities under moso bamboo and masson pine forest were significantly different (R=0.388 1, P=0.009). Furthermore, the soil microbial communities were different between the moso bamboo stands with short-term (5a, 9a) and long-term (15a, 18a) extensive management. The soil moisture, AN, AP and pH explained 90.28% of the total microbial community variation, among which the soil moisture, AN and AP contributed significantly to the microbial community variation (P < 0.05). Conclusion The results indicate that long-term extensive management has a negative effect on soil microbial biomass and result in shifting of microbial community structure. However, the ecological risks of the forest substitution and long-term extensive management still need to be further assessed. -
表 1 不同粗放经营年限毛竹林土壤理化性质
Table 1. Soil physic-chemical properties under bamboo stands with different culture histories
处理
Treatment含水量
Soil Moisture/%pH
(H2O)有机质
Organic matter/(g·kg-1)碱解氮
Alkalytic N/(mg·kg-1)有效磷
Available P/(mg·kg-11)速效钾
Available K/(mg·kg-1)MP 23.71±0.60a 4.51±0.05c 43.28±2.18b 115.21±21.88a 0.64±0.15b 40.0±7.2b 5a 18.91±1.45b 4.61±0.08bc 57.84±1.60a 102.20±3.9ab 1.97±0.41a 67.0±8.2a 9a 20.42±1.00b 4.88±0.15a 35.75±1.25bc 93.12±3.7ab 1.95±0.43a 41.0±5.0b 15a 19.08±1.99b 4.71±0.12b 41.87±2.13bc 93.95±20.4ab 1.49±0.85ab 45.7±3.2b 18a 15.51±2.13c 4.63±0.06bc 35.01±4.22c 70.15±5.1b 1.23±0.25ab 44.7±5.9b 注: 数据为3个重复的平均值及标准差(mean±SD);同列中不同字母表示处理间差异达5%显著水平。MP表示马尾松林;5 a、9 a、15 a、18 a表示毛竹林粗放经营年限为5年、9年、15年、18年。下同。
Note: The data in the table are the means of 3 replicates and the standard error (mean±SD). Different letters in the same column indicate significant difference at 5% level. MP indicates masson pine. 5 a、9 a、15 a、18 a indicate moso bamboo stands with 5, 9, 15, 18 years of extensive management age, respectively. The same below.表 2 不同粗放经营年限毛竹林土壤PLFA含量
Table 2. Soil PLFA contents under bamboo stands with different culture histories
nmol·g-1 处理
Treatment总PLFA
Total PLFA细菌
Bacteria革兰氏阳性细菌
G+ bacteria革兰氏阴性细菌
G- bacteria真菌
Fungi放线菌
Actinomycete原生动物
ProtozaMP 113.01±19.98a 63.97±13.92a 24.65±4.76a 22.55±5.84a 8.69±2.09a 11.28±4.74a 0.53±0.14a 5a 48.05±16.60b 28.74±10.20b 9.81±3.34b 11.30±4.49b 4.53±1.77b 5.75±1.96b 0.24±0.05b 9a 43.74±11.67b 26.86±7.48b 9.42±2.80b 10.53±2.91b 3.93±1.03b 4.30±2.21b 0.31±0.18b 15a 58.82±15.49b 34.87±9.72b 12.63±3.47b 16.10±5.01ab 5.36±1.29b 6.50±1.34b 0.25±0.04b 18a 46.61±4.28b 25.56±2.45b 9.53±0.60b 9.98±1.10b 5.14±0.55b 5.57±0.52b 0.26±0.04b 表 3 逐步回归分析
Table 3. Stepwise regression analysis
因变量
Dependence自变量
IndependenceF值
F value显著性
Significance总Total AN, AP 26.703 0.000 细菌Bacteria AN, AP 26.348 0.000 真菌Fungi AP, AN 13.043 0.001 放线菌Actinomycete AN, pH, OM 44.152 0.000 原生动物Protoza AN, AP 8.672 0.005 G-细菌G- bacteria AN, AP 17.134 0.000 G+细菌G+ bacteria AN, AP 23.552 0.000 注:AN、AP、OM分别为碱解氮、有效磷和土壤有机质。
Note: AN, AP, and OM indicate alkalytic N, available P and soil organic matter, respectively.表 4 土壤微生物多样性指数
Table 4. Soil microbial diversity indexes
处理
Treatment物种丰富度
RichnessShannon多样性
Shannon diversitySimpson多样性
Simpson diversityShannon均匀度
Shannon evenessSimpson均匀度
Simpson evenessMP 45±6.81a 23.55±2.16a 17.03±1.65a 0.53±0.05bc 0.38±0.05bc 5a 44±6.66a 21.39±0.90ab 15.09±0.59b 0.49±0.07c 0.35±0.06c 9a 47±1.00a 21.36±1.12ab 14.72±0.74b 0.45±0.03c 0.31±0.02c 15a 35±3.21b 20.60±0.71b 15.17±0.34b 0.60±0.04ab 0.44±0.04ab 18a 32±0.58b 20.00±0.25b 15.28±0.54b 0.63±0.02a 0.48±0.03a -
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