菌根真菌与氮磷对针茅草原群落生产力变化的作用机制
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摘要
丛枝菌根真菌(AMF)是陆地生态系统的重要组成部分,能够与约80%的陆地植物种类形成AMF-植物共生体。AMF为植物提供氮(N)磷(P)等土壤养分,植物为AMF提供光合产物。土壤NP水平及其平衡性(N：P),调节AMF与植物间的共生关系,影响AMF对植物生长的贡献。大量研究表明AMF影响植物养分吸收,调节物种间的相互作用关系和植被更新,作用于植物群落的生产力变化。然而,AMF与土壤NP水平及其平衡性互作对植物群落生产力变化的作用机制还知之甚少。
本研究采用原位控制试验和室内盆栽试验相结合的方法探讨以下三个内容：(1)原位施用真菌抑制剂(苯菌灵)和P肥对克氏针茅(Stipa krylovii)草原植物群落生产力、多样性和稳定性的影响；(2)AMF与土壤NP水平及其平衡性互作对草原植物群落生产力和多样性的影响;(3)AMF和建立菌丝网络的成株植物对幼苗定植的影响。主要结果如下：
(1)三年原位试验的结果表明：AMF对克氏针茅草原植物群落的物种多样性无显著影响,但有助于保持豆科植物的物种多样性。不添加P肥时,AMF对植物群落生产力无显著影响,添加P肥后,AMF降低了植物群落的生产力。较低P肥添加水平下,AMF对植物群落生产力的时间稳定性无显著影响,而较高P肥添加水平下,AMF增加了生产力的时间稳定性,也增加了群落中优势度较大的物种间的补偿性生长效应。
(2)盆栽试验结果表明：土壤N与AMF的交互作用对克氏针茅草原植物群落的Shannon多样性无显著影响,而高P水平下,N与AMF的交互作用显著影响了植物群落的生产力。在高P低N水平下,AMF降低了生产力,而在高P高N水平下AMF对生产力无显著影响。在低P水平下,AMF显著增加了植物群落的生产力,与N水平无关。土壤P与AMF的交互作用显著影响了植物群落的Shannon多样性。在低P水平下,AMF降低了多样性,随着P水平升高,AMF对多样性的负效应降低。在低N：P下,AMF降低了植物群落的生产力,而在较高N：P下AMF增加了生产力。AMF仅在中等N：P下显著降低了植物群落的Shannon多样性,而在较高或较低N：P下对多样性无显著影响。AMF增加了植物群落水平化学计量学的内稳性和在不同N：P下生产力的稳定性,降低了物种间的互补性生长效应。
(3)克氏针茅或羊草(Leymus chinensis)成株建立的菌丝网络更有利于其同种幼苗的P吸收。但是,冷蒿(Artemisia frigida)成株建立的菌丝网络更有利于克氏针茅和冰草(Agropyron cristatum)幼苗的P吸收。与其它植物成株建立的菌丝网络相比,冷蒿建植的菌丝网络更有利于冷蒿幼苗的生长。克氏针茅或羊草建立的菌丝网络显著抑制了冰草幼苗的生长,而冷蒿或冰草建立的菌丝网络抑制了羊草幼苗的生长。菌丝网络对植物幼苗P吸收的作用,与菌丝网络对其生长的作用不一致。
综上所述,AMF与土壤NP水平及其平衡性互作影响了克氏针茅草原植物群落的多样性、生产力和稳定性。AMF增加了群落化学计量学的内稳性和物种间的补偿性生长效应,降低了物种间的互补性生长效应,有助于提高针茅草地生态系统的稳定性。不同种类成株植物建立菌丝网络对周围幼苗的定植影响不同。
Arbuscular mycorrhizal fungi (AMF) are important components of terrestrial ecosystems, and can form mycorrhizal symbioses with about80%of all terrestrial plant species. AMF provide soil resources such as nitrogen (N) and phosphorus (P) for plants and receive photosynthates in return. Soil available N and phosphorus P and their balance (ecological stoichiometry, N:P) mediate AMF-plant symbiosis along the mutualism-antagonism spectrum and affect the mycorrhizal contribution to plant growth. Numerous studies show that AMF impact plant community productivity through mediating plant nutrients acquisition, plant-plant interactions and vegetation regeneration. However, relatively little is known about how AMF interact with soil N and P availability and their balance to influence the productivity changes of plant communities.
In this study, we combined field and pot experiments to address the following three contents:(1) the effects of fungicide (benomyl) application and P addition on diversity, productivity and temporal stability of the Stipa krylovii Steppe community in a field experiment;(2) the interaction effects of AMF and soil N and P availability and their balance on plant community diversity and productivity;(3) the effects of AMF and the adult plant species establishing the mycorrhizal networks on seedling recruitment. The main results are as follows:
(1) In the three-year field experiment, AMF did not influence the species diversity of the plant communities, but AMF are essential for maintaining species richness of legumes. At the zero-P addition rate, AMF did not affect plant community productivity, but increased productivity under P addition treatments. At low P addition rates, AMF did not affect the temporal stability, but increased the temporal stability of plant community productivity and the compensatory effects between plants of high abundance under high P addition rates.
(2) In the pot experiment, the interaction between N addition and AMF inoculation did not significantly affect Shannon diversity of plant community, while there was a significant interaction between AMF inoculation and N addition found for community productivity at the highest P level. At high P and low N levels, AMF inoculation decreased plant community productivity, but did not affect the productivity at the highest P and N level. At low P levels, AMF inoculation increased plant community productivity independent of N addition. The interaction between P addition and AMF inoculation significantly affect Shannon diversity of plant community. AMF inoculation suppressed plant community diversity at low P levels, and P addition reduced the negative effect of AMF on the diversity. AMF inoculation decreased the community productivity at low N:P ratios, but increased it at high N:P ratios. Shannon diversity was reduced by AMF inoculation at the intermediate N:P ratios, while did not show significant response to AMF at low and high N:P ratios. AMF inoculation increased the stoichiometric homoeostasis of the plant community and the stability of plant community productivity at different N:P ratios, but reduced the complementary effects between plants.
(3) The mycorrhizal networks established by the adult plants of S. krylovii or Leymus chinensis provide more P benefit for the seedlings of the same species with the adult plants than the other species. However, the mycorrhizal networks established by the adult plants of Artemisia frigida provide more P benefit for S. krylovii and Agropyron cristatum seedlings. The mycorrhizal networks established by the adult plants of A. frigida showed more positive effects on the seedling growth of A. frigida than other species. The mycorrhizal networks established by the adult plants of S. krylovii or L. chinensis suppressed the seedling growth of A. cristatum, while the mycorrhizal networks established by the adult plants of A. cristatum or A. frigida suppressed the seedling growth of L. chinensis. The P benefit from AMF did not confirm the mycorrhizal growth response of both the seedlings plants.
In conclusion, AMF interact with soil N and P availablility and their balance to influence plant community productivity, diversity and ecosystem stability in the Stipa krylovii Steppe. AMF increased the stoichiometric homoeostasis of the plant community and compensatory effects between plants, and reduced the complementary effects between plants, which may contribute to the increase in stability of the Stipa Steppe ecosystem. Mycorrhizal networks established by various adult plant species differently affected the recruitment of surrounding seedling.
本研究采用原位控制试验和室内盆栽试验相结合的方法探讨以下三个内容：(1)原位施用真菌抑制剂(苯菌灵)和P肥对克氏针茅(Stipa krylovii)草原植物群落生产力、多样性和稳定性的影响；(2)AMF与土壤NP水平及其平衡性互作对草原植物群落生产力和多样性的影响;(3)AMF和建立菌丝网络的成株植物对幼苗定植的影响。主要结果如下：
(1)三年原位试验的结果表明：AMF对克氏针茅草原植物群落的物种多样性无显著影响,但有助于保持豆科植物的物种多样性。不添加P肥时,AMF对植物群落生产力无显著影响,添加P肥后,AMF降低了植物群落的生产力。较低P肥添加水平下,AMF对植物群落生产力的时间稳定性无显著影响,而较高P肥添加水平下,AMF增加了生产力的时间稳定性,也增加了群落中优势度较大的物种间的补偿性生长效应。
(2)盆栽试验结果表明：土壤N与AMF的交互作用对克氏针茅草原植物群落的Shannon多样性无显著影响,而高P水平下,N与AMF的交互作用显著影响了植物群落的生产力。在高P低N水平下,AMF降低了生产力,而在高P高N水平下AMF对生产力无显著影响。在低P水平下,AMF显著增加了植物群落的生产力,与N水平无关。土壤P与AMF的交互作用显著影响了植物群落的Shannon多样性。在低P水平下,AMF降低了多样性,随着P水平升高,AMF对多样性的负效应降低。在低N：P下,AMF降低了植物群落的生产力,而在较高N：P下AMF增加了生产力。AMF仅在中等N：P下显著降低了植物群落的Shannon多样性,而在较高或较低N：P下对多样性无显著影响。AMF增加了植物群落水平化学计量学的内稳性和在不同N：P下生产力的稳定性,降低了物种间的互补性生长效应。
(3)克氏针茅或羊草(Leymus chinensis)成株建立的菌丝网络更有利于其同种幼苗的P吸收。但是,冷蒿(Artemisia frigida)成株建立的菌丝网络更有利于克氏针茅和冰草(Agropyron cristatum)幼苗的P吸收。与其它植物成株建立的菌丝网络相比,冷蒿建植的菌丝网络更有利于冷蒿幼苗的生长。克氏针茅或羊草建立的菌丝网络显著抑制了冰草幼苗的生长,而冷蒿或冰草建立的菌丝网络抑制了羊草幼苗的生长。菌丝网络对植物幼苗P吸收的作用,与菌丝网络对其生长的作用不一致。
综上所述,AMF与土壤NP水平及其平衡性互作影响了克氏针茅草原植物群落的多样性、生产力和稳定性。AMF增加了群落化学计量学的内稳性和物种间的补偿性生长效应,降低了物种间的互补性生长效应,有助于提高针茅草地生态系统的稳定性。不同种类成株植物建立菌丝网络对周围幼苗的定植影响不同。
Arbuscular mycorrhizal fungi (AMF) are important components of terrestrial ecosystems, and can form mycorrhizal symbioses with about80%of all terrestrial plant species. AMF provide soil resources such as nitrogen (N) and phosphorus (P) for plants and receive photosynthates in return. Soil available N and phosphorus P and their balance (ecological stoichiometry, N:P) mediate AMF-plant symbiosis along the mutualism-antagonism spectrum and affect the mycorrhizal contribution to plant growth. Numerous studies show that AMF impact plant community productivity through mediating plant nutrients acquisition, plant-plant interactions and vegetation regeneration. However, relatively little is known about how AMF interact with soil N and P availability and their balance to influence the productivity changes of plant communities.
In this study, we combined field and pot experiments to address the following three contents:(1) the effects of fungicide (benomyl) application and P addition on diversity, productivity and temporal stability of the Stipa krylovii Steppe community in a field experiment;(2) the interaction effects of AMF and soil N and P availability and their balance on plant community diversity and productivity;(3) the effects of AMF and the adult plant species establishing the mycorrhizal networks on seedling recruitment. The main results are as follows:
(1) In the three-year field experiment, AMF did not influence the species diversity of the plant communities, but AMF are essential for maintaining species richness of legumes. At the zero-P addition rate, AMF did not affect plant community productivity, but increased productivity under P addition treatments. At low P addition rates, AMF did not affect the temporal stability, but increased the temporal stability of plant community productivity and the compensatory effects between plants of high abundance under high P addition rates.
(2) In the pot experiment, the interaction between N addition and AMF inoculation did not significantly affect Shannon diversity of plant community, while there was a significant interaction between AMF inoculation and N addition found for community productivity at the highest P level. At high P and low N levels, AMF inoculation decreased plant community productivity, but did not affect the productivity at the highest P and N level. At low P levels, AMF inoculation increased plant community productivity independent of N addition. The interaction between P addition and AMF inoculation significantly affect Shannon diversity of plant community. AMF inoculation suppressed plant community diversity at low P levels, and P addition reduced the negative effect of AMF on the diversity. AMF inoculation decreased the community productivity at low N:P ratios, but increased it at high N:P ratios. Shannon diversity was reduced by AMF inoculation at the intermediate N:P ratios, while did not show significant response to AMF at low and high N:P ratios. AMF inoculation increased the stoichiometric homoeostasis of the plant community and the stability of plant community productivity at different N:P ratios, but reduced the complementary effects between plants.
(3) The mycorrhizal networks established by the adult plants of S. krylovii or Leymus chinensis provide more P benefit for the seedlings of the same species with the adult plants than the other species. However, the mycorrhizal networks established by the adult plants of Artemisia frigida provide more P benefit for S. krylovii and Agropyron cristatum seedlings. The mycorrhizal networks established by the adult plants of A. frigida showed more positive effects on the seedling growth of A. frigida than other species. The mycorrhizal networks established by the adult plants of S. krylovii or L. chinensis suppressed the seedling growth of A. cristatum, while the mycorrhizal networks established by the adult plants of A. cristatum or A. frigida suppressed the seedling growth of L. chinensis. The P benefit from AMF did not confirm the mycorrhizal growth response of both the seedlings plants.
In conclusion, AMF interact with soil N and P availablility and their balance to influence plant community productivity, diversity and ecosystem stability in the Stipa krylovii Steppe. AMF increased the stoichiometric homoeostasis of the plant community and compensatory effects between plants, and reduced the complementary effects between plants, which may contribute to the increase in stability of the Stipa Steppe ecosystem. Mycorrhizal networks established by various adult plant species differently affected the recruitment of surrounding seedling.
引文
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