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丛枝菌根真菌(Arbuscular mycorrhizal fungi, AMF)能够增强植物根系对养分的获取能力,而植物为其提供生长所需的碳源,两者形成互惠共生的丛枝菌根[1]。研究表明,AMF能与90%的维管植物形成丛枝菌根,其中AMF侵染率和孢子密度被用作表征共生关系的紧密程度[2-3]。AMF的定殖和产孢与营养元素和地下生物量存在密切的关系[4-5]。此外,AMF侵染率和孢子密度也可以一定程度反映AMF对生存策略的选择,对了解寄主植物的生长发育状况具有重要意义。以往研究主要集中在农田生态系统中,如冬小麦(Triticum aestivum Linn.)、蚕豆(Vicia faba Linn.)等[6-8]。由于森林根系较复杂,目前对森林生态系统的研究较少。
毛竹(Phyllostachys edulis (Carrière) J. Houz.)是我国亚热带地区分布最为广泛的竹种,其面积为467.8万hm2,占我国竹林面积的73.0%[9],且有不断增加的趋势[10]。毛竹具有独特复杂的根鞭系统,根冠比(0.37~0.50)远高于其他森林生态系统[11],这可能导致毛竹林土壤AMF特征与其他森林存在差异。因此,亟需加强对毛竹林土壤AMF特征的研究。此外,毛竹林分布区是我国氮沉降最严重的地区之一(30.7 kg·hm−2·yr−1)[12],大量的氮输入会导致土壤pH、有机质含量和C:N降低[13],势必会影响AMF的特征。然而,关于氮添加对AMF侵染率和孢子密度影响的研究存在争议[6,14-15]。例如,陈良华等[14]发现施氮显著降低了杉木(Cunninghamia lanceolata (Lamb.) Hook.)AMF侵染率,而汪鹞雄等[15]发现氮添加会显著提高杉木AMF侵染率,Wang等[6]发现长期施氮显著降低了玉米(Zea mays Linn.)和小麦的AMF侵染率和孢子密度。目前,氮添加对毛竹林AMF的影响知之甚少。
生物炭是由生物质在缺氧条件下热裂解而成的芳香类化合物[16],具有多孔道、表面积巨大的特性[17]。生物炭作为一种土壤添加剂,已被广泛用于农业生态系统中。研究表明,施用生物炭提高了土壤的孔隙度[18]、pH[19]和含水率[20],同时增加土壤中营养物质的含量,尤其是氮、磷含量[21]。王策等[22]研究发现添加生物炭可以有效提高土壤中有机质含量,改善土壤团粒结构和微生物群落结构。然而,在大气氮沉降导致的土壤酸化和养分失衡背景下,添加生物炭对毛竹林土壤AMF的影响及其调控机制尚不清楚,相关研究亟待开展。本文以毛竹林为研究对象,探讨了不同水平的氮和生物炭添加对毛竹林AMF侵染率和孢子密度以及土壤理化性质的影响,以期从AMF共生关系角度为全球变化背景下毛竹林的可持续经营提供科学参考。
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与对照相比,氮添加显著提高了土壤微生物量碳(38.5%~110.3%)、有效磷(75.2%~144.5%)和有效氮(27.9%~66.7%)含量,显著降低了土壤含水率(5.6%~15.2%)和pH(7.6%~11.0%)(P < 0.05,表1)。生物炭添加(BC20和BC40)显著提高了土壤pH和有效磷含量,显著降低了可溶性有机氮、微生物量碳和有效氮含量(P < 0.05)。在氮添加处理下,生物炭添加显著提高了土壤含水率(2.6%~18.5%)和pH(2.6%~11.0%),显著降低了土壤有效氮(2.4%~22.5%)含量(P < 0.05)。可重复双因素方差分析表明氮和生物炭添加及其交互作用均显著影响土壤含水率、pH、可溶性有机氮、微生物量碳、有效磷和有效氮(表2,P < 0.05)。
表 1 不同氮和生物炭添加处理下毛竹林的土壤特性
Table 1. Soil properties under different N and biochar addition treatments in the Moso bamboo plantations
处理
Treatment含水率
SWC/%酸碱度
pH可溶性有机氮
DON/(mg·kg−1)微生物量碳
MBC/(mg·kg−1)有效磷
AP/(mg·kg−1)有效氮
AN/(mg·kg−1)N0-BC0 32.68±1.77 Aa 4.62±0.02 Ab 42.07±0.81 Aa 554.18±8.19 Da 7.05±0.16 Dc 54.87±2.34 Ca N0-BC20 28.66±0.07 b 4.75±0.04 a 30.71±0.56 c 518.10±14.59 b 12.11±0.96 b 49.84±2.03 b N0-BC40 32.03±0.16 a 4.79±0.05 a 34.84±0.69 b 514.55±6.25 b 17.1±0.12 a 49.75±1.20 b N30-BC0 27.7±0.4 Cb 4.27±0.02 Bb 30.54±1.38 Ca 1 165.48±1.82 Aa 14.98±0.61 Bb 70.19±7.41 Ba N30-BC20 32.82±1.63 a 4.36±0.02 b 33.39±1.4 a 1 165.19±5.13 a 17.79±0.89 a 68.53±1.08 ab N30-BC40 30.82±0.75 a 4.52±0.09 a 32.28±2.21 a 944.24±16.77 b 12.22±1.83 c 60.67±2.12 b N60-BC0 30.84±0.23 Bb 4.19±0.07 Cb 40.39±0.56 Aab 971.07±23.59 Bb 12.35±0.59 Cc 77.67±2.24 Ba N60-BC20 32.56±0.2 a 4.31±0.07 a 37.13±3.59 b 1 074.50±29.27 a 16.53±0.24 a 64.15±2.23 b N60-BC40 33.25±0.6 a 4.30±0.03 ab 42.84±0.74 a 955.33±8.37 b 13.95±0.24 b 63.6±2.84 b N90-BC0 30.39±0.5 Bb 4.11±0.02 Dc 37.04±1.66 Ba 767.8±26.62 Ca 17.24±0.32 Aa 91.46±2.43 Aa N90-BC20 31.96±0.49 a 4.19±0.01 b 35.86±0.93 a 747.54±27.41 a 16.77±0.53 a 83.94±4.08 b N90-BC40 31.19±1.03 ab 4.23±0.02 a 31.16±1.98 b 598.9±11.49 b 14.56±2.61 a 70.88±1.27 c 注:不同的大写字母表示在无生物炭添加处理下,不同氮添加处理之间存在显著差异(P < 0.05)。不同的小写字母表示在相同的氮添加处理下,不同生物炭添加处理之间存在显著差异(P < 0.05)
Notes: Different capital letters indicate significant differences among different N treatments without biochar addition treatments (P < 0.05). Different lowercase letters indicate significant differences among different biochar addition treatments under the same N addition treatments (P < 0.05)表 2 氮和生物炭添加对毛竹林土壤理化性质、AMF侵染率和孢子密度影响的双因素方差分析
Table 2. Two-way ANOVA of effects of N and biochar addition on soil properties, colonization rate and spore density of AMF in the Moso bamboo plantations
差异源
Source of
Variables氮添加
N addition生物炭
Biochar
addition交互作用
InteractionF P F P F P SWC 6.778 0.002 9.377 0.001 15.816 0.000 pH 259.735 0.000 41.902 0.000 2.606 0.043 DON 38.440 0.000 12.541 0.000 15.959 0.000 MBC 1 979.948 0.000 179.365 0.000 35.666 0.000 AP 24.463 0.000 22.993 0.000 28.372 0.000 AN 148.885 0.000 48.265 0.000 5.668 0.002 侵染率
Colonization rate30.763 0.000 4.319 0.025 19.014 0.000 孢子密度
Spore density20.873 0.000 11.595 0.000 12.888 0.000 -
与对照相比,氮和高生物炭添加均显著降低了AMF侵染率(P < 0.05,图2)。在N30处理下,BC40处理显著提高了AMF侵染率(108.2%,P < 0.05)。在N60和N90处理下,BC20和BC40处理对AMF侵染率没有显著影响(P > 0.05)。可重复双因素方差分析表明氮和生物炭添加及其交互作用均显著影响毛竹林土壤AMF侵染率(表2,P < 0.05)。
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与对照相比,氮和生物炭添加提高了AMF孢子密度,尤其是N30和BC40处理, 达到了统计意义上的显著性水平(P < 0.05,图3)。在N30处理下,BC20处理显著提高了孢子密度(23.7%,P < 0.05)。在N60和N90处理下,BC20和BC40处理对孢子密度没有显著影响(P > 0.05)。可重复双因素方差分析表明氮和生物炭添加及其交互作用均极显著影响毛竹林土壤AMF孢子密度(表2,P < 0.01)。
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Pearson相关性分析表明,AMF侵染率与土壤pH呈极显著正相关关系(P < 0.01),与AP和AN呈显著负相关关系(P < 0.05);孢子密度与pH和AP呈极显著正相关关系(P < 0.01),与可溶性有机氮(DON)呈显著负相关关系(P < 0.05)(表3)。
表 3 毛竹林土壤AMF侵染率、孢子密度与土壤理化特征的相关性分析
Table 3. Correlations between colonization rate and spore density of AMF and soil properties
SWC pH DON MBC AP AN 侵染率
Colonization rate−0.208 0.429** −0.207 0.067 −0.478** −0.403* 孢子密度
Spore density0.037 0.373** −0.358* 0.229 0.461** −0.286 注/Notes:*: P < 0.05; **: P < 0.01
氮和生物炭添加对毛竹林AMF侵染率和孢子密度的影响
Effects of Nitrogen and Biochar Addition on Arbuscular Mycorrhizal Fungi Colonization Rate and Spore Density in Moso Bamboo Plantations
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摘要:
目的 探讨氮沉降和生物炭添加对毛竹林丛枝菌根真菌特征的影响,为全球变化背景下毛竹林的可持续经营提供科学参考。 方法 以毛竹林为研究对象,探讨了不同梯度的氮沉降(0、30、60和90 kg·hm−2·yr−1)和生物炭添加(0、20和40 t ·hm−2)及其复合作用对毛竹林AMF侵染率和孢子密度以及土壤理化性质的影响。 结果 与对照(0 kg·hm−2·yr−1 N + 0 t·hm−2 BC)相比,氮添加显著降低了AMF侵染率(16.1%~51.7%)。高生物炭添加(40 t·hm−2)显著降低了AMF侵染率(46.0%),但提高了孢子密度(162.5%)。在氮添加处理下,生物炭添加提高了AMF侵染率,对孢子密度无显著影响。AMF侵染率与土壤pH呈显著正相关关系,与有效氮和有效磷呈显著负相关关系。AMF孢子密度与有效磷呈显著正相关关系。 结论 在氮添加处理下,生物炭添加提高了AMF侵染率,增强了毛竹与AMF的共生关系,表明在氮沉降背景下生物炭添加是实现毛竹林可持续经营的有效措施之一。 Abstract:Objective To investigate the effects of nitrogen (N) deposition and biochar addition on arbuscular mycorrhizal fungi (AMF) characteristics in Moso bamboo (Phyllostachys edulis (Carrière) J. Houz.) plantations for providing scientific reference for sustainable management of Moso bamboo plantations under global change. Methods We investigated the effects of different N addition rates (0, 30, 60, and 90 kg N·hm−2·yr−1), biochar addition rates (0, 20, and 40 t BC·hm−2), and their combination on the AMF colonization rate, spore density and soil properties of Phyllostachys edulis plantations. Results Compared with the control (0 kg N·hm−2·yr−1 + 0 t BC·hm−2), N addition significantly reduced the colonization rate of AMF (16.1%~51.7%). High biochar addition (40 t BC·hm−2) significantly reduced the colonization rate of AMF (46.0%), but increased the spore density (162.5%). Under the N addition treatments, biochar addition increased the colonization rate of AMF, and had no significant effect on spore density. The colonization rate of AMF was significantly positively correlated with soil pH, and negatively correlated with available N and available phosphorus. The spore density of AMF was significantly positively correlated with available phosphorus. Conclusion Under the N addition treatments, biochar addition increased the colonization rate of AMF and enhanced the symbiotic relationship between Moso bamboo and AMF, indicating that biochar addition is an effective measure to achieve sustainable management of Moso bamboo plantations under the background of N deposition. -
表 1 不同氮和生物炭添加处理下毛竹林的土壤特性
Table 1. Soil properties under different N and biochar addition treatments in the Moso bamboo plantations
处理
Treatment含水率
SWC/%酸碱度
pH可溶性有机氮
DON/(mg·kg−1)微生物量碳
MBC/(mg·kg−1)有效磷
AP/(mg·kg−1)有效氮
AN/(mg·kg−1)N0-BC0 32.68±1.77 Aa 4.62±0.02 Ab 42.07±0.81 Aa 554.18±8.19 Da 7.05±0.16 Dc 54.87±2.34 Ca N0-BC20 28.66±0.07 b 4.75±0.04 a 30.71±0.56 c 518.10±14.59 b 12.11±0.96 b 49.84±2.03 b N0-BC40 32.03±0.16 a 4.79±0.05 a 34.84±0.69 b 514.55±6.25 b 17.1±0.12 a 49.75±1.20 b N30-BC0 27.7±0.4 Cb 4.27±0.02 Bb 30.54±1.38 Ca 1 165.48±1.82 Aa 14.98±0.61 Bb 70.19±7.41 Ba N30-BC20 32.82±1.63 a 4.36±0.02 b 33.39±1.4 a 1 165.19±5.13 a 17.79±0.89 a 68.53±1.08 ab N30-BC40 30.82±0.75 a 4.52±0.09 a 32.28±2.21 a 944.24±16.77 b 12.22±1.83 c 60.67±2.12 b N60-BC0 30.84±0.23 Bb 4.19±0.07 Cb 40.39±0.56 Aab 971.07±23.59 Bb 12.35±0.59 Cc 77.67±2.24 Ba N60-BC20 32.56±0.2 a 4.31±0.07 a 37.13±3.59 b 1 074.50±29.27 a 16.53±0.24 a 64.15±2.23 b N60-BC40 33.25±0.6 a 4.30±0.03 ab 42.84±0.74 a 955.33±8.37 b 13.95±0.24 b 63.6±2.84 b N90-BC0 30.39±0.5 Bb 4.11±0.02 Dc 37.04±1.66 Ba 767.8±26.62 Ca 17.24±0.32 Aa 91.46±2.43 Aa N90-BC20 31.96±0.49 a 4.19±0.01 b 35.86±0.93 a 747.54±27.41 a 16.77±0.53 a 83.94±4.08 b N90-BC40 31.19±1.03 ab 4.23±0.02 a 31.16±1.98 b 598.9±11.49 b 14.56±2.61 a 70.88±1.27 c 注:不同的大写字母表示在无生物炭添加处理下,不同氮添加处理之间存在显著差异(P < 0.05)。不同的小写字母表示在相同的氮添加处理下,不同生物炭添加处理之间存在显著差异(P < 0.05)
Notes: Different capital letters indicate significant differences among different N treatments without biochar addition treatments (P < 0.05). Different lowercase letters indicate significant differences among different biochar addition treatments under the same N addition treatments (P < 0.05)表 2 氮和生物炭添加对毛竹林土壤理化性质、AMF侵染率和孢子密度影响的双因素方差分析
Table 2. Two-way ANOVA of effects of N and biochar addition on soil properties, colonization rate and spore density of AMF in the Moso bamboo plantations
差异源
Source of
Variables氮添加
N addition生物炭
Biochar
addition交互作用
InteractionF P F P F P SWC 6.778 0.002 9.377 0.001 15.816 0.000 pH 259.735 0.000 41.902 0.000 2.606 0.043 DON 38.440 0.000 12.541 0.000 15.959 0.000 MBC 1 979.948 0.000 179.365 0.000 35.666 0.000 AP 24.463 0.000 22.993 0.000 28.372 0.000 AN 148.885 0.000 48.265 0.000 5.668 0.002 侵染率
Colonization rate30.763 0.000 4.319 0.025 19.014 0.000 孢子密度
Spore density20.873 0.000 11.595 0.000 12.888 0.000 表 3 毛竹林土壤AMF侵染率、孢子密度与土壤理化特征的相关性分析
Table 3. Correlations between colonization rate and spore density of AMF and soil properties
SWC pH DON MBC AP AN 侵染率
Colonization rate−0.208 0.429** −0.207 0.067 −0.478** −0.403* 孢子密度
Spore density0.037 0.373** −0.358* 0.229 0.461** −0.286 注/Notes:*: P < 0.05; **: P < 0.01 -
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