模拟增温和丛枝菌根对门源草原毛虫幼虫生长发育的影响
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摘要
采用开顶式生长室(Open top chamber, OTC)模拟增温和苯菌灵抑制丛枝菌根研究上述两种因素综合作用下门源草原毛虫幼虫的生长速率、蛹化时间和蛹重。结果表明,增温和丛枝菌根抑制及其交互作用均对门源草原毛虫幼虫生长速率产生了显著影响。相比对照组而言,增温使该指标升高了34%。丛枝菌根抑制未对上述指标产生显著影响。增温和丛枝菌根抑制的交互作用使门源草原毛虫幼虫生长速率较对照组升高了16%,而较增温组降低了13%。增温处理下雌、雄幼虫的蛹化时间分别为204、218 d,而不增温处理下分别为212、223 d。增温使得雌、雄幼虫的蛹化时间较不增温处理分别提前了2%和4%。增温和不增温处理下的雌、雄虫蛹化时间差分别为15、12 d。增温将上述时间差扩大了25%。此外,增温及其与丛枝菌根抑制的交互作用对门源草原毛虫雌虫蛹重的影响显著,而对于雄虫的蛹重来说,仅增温处理的影响显著。增温和增温丛枝菌根抑制处理,使得雌蛹重较对照组增大了22%和8%。增温使雄蛹重增大了18%。首次研究了增温和丛枝菌根对植食性昆虫的综合影响。
It is known that global warming, an extremely threatening problem for human, changes the ecological processes and ecosystem in an unprecedented way. There is no doubt that climate warming can significantly affect insects that are more sensitive to the changes in environment and are regarded as the bio-indicator of ecosystem to disturbances. Herbivores attract more attention than other insects owing to their advantages in terms of quantity and function. Although the general effects of warming on herbivores are positive, recent studies suggest that they are highly variable across species and depend on the optimum temperature. In addition, there are many abiotic and biotic factors, such as soil fertilization, precipitation, wind, below ground herbivores, arbuscular mycorrhiza fungi, and enemies of herbivores affecting the interspecific interactions of predation. They can probably alter the responses of herbivores to warming. This hypothesis had been validated by some field studies. Arbuscular mycorrhiza fungi(AMF), one of the most widely distributed soil organisms, develop root symbiotic associations with nearly 85% of terrestrial plants. They can help roots uptake mineral elements, such as nitrogen(N) and phosphorus(P), in exchange for photosynthetic products from host plant. By this pathway, AMF can interact with their host plants, and the outcome of these interactions in plants can cascade up to insect herbivores. Despite little consensuses on the effects of AMF on predation, several studies have suggested that they cannot be ignored. Surprisingly, there is no study on the role of AMF in regulating the responses of herbivores to warming. To close this gap, we conducted an in situ experiment in Tibetan Plateau(TP), the highest and largest plateau on earth, where temperatures increase more rapidly than the global average. We used an open top chamber(OTC), benomyl simulating warming, and AMF control to investigate their influences on the growth rate(RG) and pupation time(PT) of Gynaephora menyuanensis larvae, an endemic generalist species in northeastern TP. The larvae acted an ideal object of study to solve the above problem. The results showed that the colonization rate of AMF with fungicide treatment was 29% lower than that in no fungicide treatment group, indicating that fungicide significantly suppressed mycorrhiza symbiosis. The mean GRs under no warming without fungicide(NWNF), no warming with fungicide(NWF), warming without fungicide(WNF), and warming with fungicide(WF) treatments were 1.63, 2.17, 1.57, and 1.88 mg/d. The effects of warming, fungicide, and their interaction on the GR of caterpillars were significant. The GRs under WNF and WF treatments increased by 34% and 16% compared with that under NWNF treatment. The GR under WF treatment decreased by 13% compared with that under WNF treatment. Besides, warming significantly affected the PT of both female and male. The advancement of pupation time of female(2%) and male(4%) under warming(W) treatment was higher than that under no warming(NW) treatment. The difference between the pupation of female and male caterpillars under W treatment increased by 25% compared with that under NW treatment. The effects of warming, fungicide, and their interaction on the pupal weight(PW) of female caterpillars were significant, but PW of males were only altered by warming. The PWs of females under WNF and WF treatments increased by 22% and 8% compared with that under NWNF treatment. And the PWs of males under W treatment was increased by 18% compared with that under NWNF treatment. This is the first report on the effects of warming and arbuscular mycorrhiza fungi on herbivores.
It is known that global warming, an extremely threatening problem for human, changes the ecological processes and ecosystem in an unprecedented way. There is no doubt that climate warming can significantly affect insects that are more sensitive to the changes in environment and are regarded as the bio-indicator of ecosystem to disturbances. Herbivores attract more attention than other insects owing to their advantages in terms of quantity and function. Although the general effects of warming on herbivores are positive, recent studies suggest that they are highly variable across species and depend on the optimum temperature. In addition, there are many abiotic and biotic factors, such as soil fertilization, precipitation, wind, below ground herbivores, arbuscular mycorrhiza fungi, and enemies of herbivores affecting the interspecific interactions of predation. They can probably alter the responses of herbivores to warming. This hypothesis had been validated by some field studies. Arbuscular mycorrhiza fungi(AMF), one of the most widely distributed soil organisms, develop root symbiotic associations with nearly 85% of terrestrial plants. They can help roots uptake mineral elements, such as nitrogen(N) and phosphorus(P), in exchange for photosynthetic products from host plant. By this pathway, AMF can interact with their host plants, and the outcome of these interactions in plants can cascade up to insect herbivores. Despite little consensuses on the effects of AMF on predation, several studies have suggested that they cannot be ignored. Surprisingly, there is no study on the role of AMF in regulating the responses of herbivores to warming. To close this gap, we conducted an in situ experiment in Tibetan Plateau(TP), the highest and largest plateau on earth, where temperatures increase more rapidly than the global average. We used an open top chamber(OTC), benomyl simulating warming, and AMF control to investigate their influences on the growth rate(RG) and pupation time(PT) of Gynaephora menyuanensis larvae, an endemic generalist species in northeastern TP. The larvae acted an ideal object of study to solve the above problem. The results showed that the colonization rate of AMF with fungicide treatment was 29% lower than that in no fungicide treatment group, indicating that fungicide significantly suppressed mycorrhiza symbiosis. The mean GRs under no warming without fungicide(NWNF), no warming with fungicide(NWF), warming without fungicide(WNF), and warming with fungicide(WF) treatments were 1.63, 2.17, 1.57, and 1.88 mg/d. The effects of warming, fungicide, and their interaction on the GR of caterpillars were significant. The GRs under WNF and WF treatments increased by 34% and 16% compared with that under NWNF treatment. The GR under WF treatment decreased by 13% compared with that under WNF treatment. Besides, warming significantly affected the PT of both female and male. The advancement of pupation time of female(2%) and male(4%) under warming(W) treatment was higher than that under no warming(NW) treatment. The difference between the pupation of female and male caterpillars under W treatment increased by 25% compared with that under NW treatment. The effects of warming, fungicide, and their interaction on the pupal weight(PW) of female caterpillars were significant, but PW of males were only altered by warming. The PWs of females under WNF and WF treatments increased by 22% and 8% compared with that under NWNF treatment. And the PWs of males under W treatment was increased by 18% compared with that under NWNF treatment. This is the first report on the effects of warming and arbuscular mycorrhiza fungi on herbivores.
引文
[1] O′connor M I.Warming strengthens an herbivore-plant interaction.Ecology,2009,90(2):388- 398.
[2] Cao H,Zhao X Q,Wang S P,Zhao L,Duan J C,Zhang Z H,Ge S D,Zhu X X.Grazing intensifies degradation of a Tibetan Plateau alpine meadow through plant-pest interaction.Ecology and Evolution,2015,5(12):2478- 2486.
[3] Xi X Q,Griffin J N,Sun S C.Grasshoppers amensalistically suppress caterpillar performance and enhance plant biomass in an alpine meadow.Oikos,2013,122(7):1049- 1057.
[4] 严林.草原毛虫属的分类、地理分布及门源草原毛虫生活史对策的研究[D].兰州:兰州大学,2006.
[5] 郑莉莉,宋明华,尹谭凤,于飞海.青藏高原高寒草甸门源草原毛虫取食偏好及其与植物C、N含量的关系.生态学报,2016,36(8):2319- 2326.
[6] Smith D S,Schweitzer J A,Turk P,Bailey J K,Hart S C,Shuster S M,Whitham T G.Soil-mediated local adaptation alters seedling survival and performance.Plant and Soil,2012,352(1/2):243- 251.
[7] Koricheva J,Gange A C,Jones T.Effects of mycorrhizal fungi on insect herbivores:a meta-analysis.Ecology,2009,90(8):2088- 2097.
[8] Kula A A R,Hartnett D C.Effects of mycorrhizal symbiosis on aboveground arthropod herbivory in tallgrass prairie:an in situ experiment.Plant Ecology,2015,216(4):589- 597.
[9] 赵新全.高寒草甸生态系统与全球变化[M].北京:科学出版社,2009.
[10] Debevec E M,MacLean S F Jr.Design of greenhouses for the manipulation of temperature in tundra plant communities.Arctic and Alpine Research,1993,25(1):56- 62.
[11] 余欣超,陈珂璐,姚步青,马真,王文颖,王慧春,赵新全,周华坤.模拟增温下门源草原毛虫幼虫生长发育特征.生态学报,2016,36(24):8002- 8007.
[12] Hartnett D C,Wilson G W T.The role of mycorrhizas in plant community structure and dynamics:lessons from grasslands.Plant and Soil,2002,244(1/2):319- 331.
[13] Smith M D,Hartnett D C,Rice C W.Effects of long-term fungicide applications on microbial properties in tallgrass prairie soil.Soil Biology and Biochemistry,2000,32(7):935- 946.
[14] 刘润进,陈应龙.菌根学[M].北京:科学出版社,2007.
[15] 石福孙,陈华峰,吴宁.增温对川西北亚高山高寒草甸植物群落碳、氮含量的影响.植物研究,2008,28(6):730- 736.
[16] 曹慧,朱文琰,赵新全.试验增温和放牧对门源草原毛虫生长发育的影响.草业学报,2016,25(1):268- 272.
[17] 张棋麟,袁明龙.草原毛虫研究现状与展望.草业科学,2013,30(4):638- 646.
[18] 曹慧.门源草原毛虫-植物相互关系对增温和放牧的响应[D].北京:中国科学院大学,2015.
[19] 余欣超.不同增温梯度对门源草原毛虫生长发育与繁殖的影响[D].北京:中国科学院大学,2015.
[20] Tammaru T,Kaitaniemi P,Ruohom?ki K.Realized fecundity in Epirrita autumnata (Lepidoptera:Geometridae):relation to body size and consequences to population dynamics.Oikos,1996,77(3):407- 416.
[21] Fitter A H,Helgason T,Hodge A.Nutritional exchanges in the arbuscular mycorrhizal symbiosis:implications for sustainable agriculture.Fungal Biology Reviews,2011,25(1):68- 72.
[22] Veresoglou S D,Chen B D,Rillig M C.Arbuscular mycorrhiza and soil nitrogen cycling.Soil Biology and Biochemistry,2012,46:53- 62.
[23] Cahill J F Jr,Elle E,Smith G R,Shore B H.Disruption of a belowground mutualism alters interactions between plants and their floral visitors.Ecology,2008,89(7):1791- 1801.
[24] 包玉英,闫伟.内蒙古中西部草原主要植物的丛枝菌根及其结构类型研究.生物多样性,2004,12(5):501- 508.
[25] 孙永芳,付娟娟,褚希彤,杨云贵,许岳飞,呼天明.丛枝菌根真菌对垂穗披碱草吸收不同氮源效率的影响.草地学报,2015,23(2):294- 301.
[2] Cao H,Zhao X Q,Wang S P,Zhao L,Duan J C,Zhang Z H,Ge S D,Zhu X X.Grazing intensifies degradation of a Tibetan Plateau alpine meadow through plant-pest interaction.Ecology and Evolution,2015,5(12):2478- 2486.
[3] Xi X Q,Griffin J N,Sun S C.Grasshoppers amensalistically suppress caterpillar performance and enhance plant biomass in an alpine meadow.Oikos,2013,122(7):1049- 1057.
[4] 严林.草原毛虫属的分类、地理分布及门源草原毛虫生活史对策的研究[D].兰州:兰州大学,2006.
[5] 郑莉莉,宋明华,尹谭凤,于飞海.青藏高原高寒草甸门源草原毛虫取食偏好及其与植物C、N含量的关系.生态学报,2016,36(8):2319- 2326.
[6] Smith D S,Schweitzer J A,Turk P,Bailey J K,Hart S C,Shuster S M,Whitham T G.Soil-mediated local adaptation alters seedling survival and performance.Plant and Soil,2012,352(1/2):243- 251.
[7] Koricheva J,Gange A C,Jones T.Effects of mycorrhizal fungi on insect herbivores:a meta-analysis.Ecology,2009,90(8):2088- 2097.
[8] Kula A A R,Hartnett D C.Effects of mycorrhizal symbiosis on aboveground arthropod herbivory in tallgrass prairie:an in situ experiment.Plant Ecology,2015,216(4):589- 597.
[9] 赵新全.高寒草甸生态系统与全球变化[M].北京:科学出版社,2009.
[10] Debevec E M,MacLean S F Jr.Design of greenhouses for the manipulation of temperature in tundra plant communities.Arctic and Alpine Research,1993,25(1):56- 62.
[11] 余欣超,陈珂璐,姚步青,马真,王文颖,王慧春,赵新全,周华坤.模拟增温下门源草原毛虫幼虫生长发育特征.生态学报,2016,36(24):8002- 8007.
[12] Hartnett D C,Wilson G W T.The role of mycorrhizas in plant community structure and dynamics:lessons from grasslands.Plant and Soil,2002,244(1/2):319- 331.
[13] Smith M D,Hartnett D C,Rice C W.Effects of long-term fungicide applications on microbial properties in tallgrass prairie soil.Soil Biology and Biochemistry,2000,32(7):935- 946.
[14] 刘润进,陈应龙.菌根学[M].北京:科学出版社,2007.
[15] 石福孙,陈华峰,吴宁.增温对川西北亚高山高寒草甸植物群落碳、氮含量的影响.植物研究,2008,28(6):730- 736.
[16] 曹慧,朱文琰,赵新全.试验增温和放牧对门源草原毛虫生长发育的影响.草业学报,2016,25(1):268- 272.
[17] 张棋麟,袁明龙.草原毛虫研究现状与展望.草业科学,2013,30(4):638- 646.
[18] 曹慧.门源草原毛虫-植物相互关系对增温和放牧的响应[D].北京:中国科学院大学,2015.
[19] 余欣超.不同增温梯度对门源草原毛虫生长发育与繁殖的影响[D].北京:中国科学院大学,2015.
[20] Tammaru T,Kaitaniemi P,Ruohom?ki K.Realized fecundity in Epirrita autumnata (Lepidoptera:Geometridae):relation to body size and consequences to population dynamics.Oikos,1996,77(3):407- 416.
[21] Fitter A H,Helgason T,Hodge A.Nutritional exchanges in the arbuscular mycorrhizal symbiosis:implications for sustainable agriculture.Fungal Biology Reviews,2011,25(1):68- 72.
[22] Veresoglou S D,Chen B D,Rillig M C.Arbuscular mycorrhiza and soil nitrogen cycling.Soil Biology and Biochemistry,2012,46:53- 62.
[23] Cahill J F Jr,Elle E,Smith G R,Shore B H.Disruption of a belowground mutualism alters interactions between plants and their floral visitors.Ecology,2008,89(7):1791- 1801.
[24] 包玉英,闫伟.内蒙古中西部草原主要植物的丛枝菌根及其结构类型研究.生物多样性,2004,12(5):501- 508.
[25] 孙永芳,付娟娟,褚希彤,杨云贵,许岳飞,呼天明.丛枝菌根真菌对垂穗披碱草吸收不同氮源效率的影响.草地学报,2015,23(2):294- 301.