外生菌根真菌对干旱条件下油松幼苗生长的影响
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摘要
通过对干旱条件下外生菌根真菌对油松苗木影响的研究,以期提高苗木成活率,促进树种更新和防治水土流失。采用盆栽控水模拟干旱胁迫,设对照(CK)、轻度干旱(LD)、中度干旱(MD)与重度干旱(SD) 4个水分处理,并接种木霉,测定油松幼苗的生长与生理指标。结果显示:1)接种木霉后,油松幼苗的生物量与根冠比均显著提高(P <0. 05);干旱胁迫下,油松幼苗株高和针叶长度都呈递减趋势;但接种木霉后,幼苗株高和针叶长度的降低趋势减弱,幼苗抗旱性显著增强(P <0. 01)。2)随着干旱胁迫加剧,油松幼苗针叶的可溶性蛋白质和淀粉含量持续降低,接种木霉后,相应指标虽亦随胁迫加剧递减,但较之未接种幼苗,其变化趋势减弱(P <0. 01)。3)干旱胁迫下,未接种和接种木霉的幼苗可溶性糖含量分别在轻度胁迫和中度胁迫达到最高值,即接种木霉后,幼苗的耐旱阈值增大。干旱胁迫下,外生菌根真菌可促进油松幼苗生长,中度干旱胁迫下木霉对油松幼苗生长的促进作用效果最显著。相同水分处理条件下,菌根苗比非菌根苗的抗旱能力更强。
[Background] It researched the effect of ectomycorrhizal fungi on Pinus tabulaeformis seedlings to increase the survival rate of seedlings,promote the regeneration of tree species,and prevent soil erosion. [Methods] Potted water was used to simulate drought stress,and four water treatments were designed: control( CK),light drought( LD),moderate drought( MD),severe drought( SD).Inoculating the seedlings of Pinus tabulaeformis with Trichoderma spp. was to study the effect of growth which ectomycorrhizal fungi had on P. tabulaeformis seedling under drought stress. To each treatment,It selected seedlings whose growth was basically the same, and measured the morphological and physiological indexes. The specific operation refered to Principles and Techniques of Plant Physiological and Biochemical Experiments written by Li Hesheng. [Results] The plant height and needle length of P. tabulaeformis seedlings were significantly affected by inoculation with Trichoderma spp. and drought stress treatment( P < 0. 01),but the interaction was not significant( P > 0. 05). The effects of inoculation with Trichoderma spp. and drought stress on the soluble protein,soluble sugar and starch content of P. tabulaeformis seedlings were significant( P < 0. 01). However,the interaction to the soluble protein and starch content of Pi. tabulaeformis seedlings was not significant( P > 0. 05),and the interaction to the soluble sugar content was significantly affected( P < 0. 01). The detailed analysis results to the morphological and physiological indexes of P. tabulaeformis seedlings were as follows:( 1)After inoculation with Trichoderma spp.,the biomass and root-shoot ratio of P. tabulaeformis seedlings were significantly increased( P < 0. 05). The mycorrhizal dependence on Trichoderma spp. was moderately dependent. Under drought stress,the plant height and needle length of P. tabulaeformis seedlings showed a decreasing trend. However,after inoculation with Trichoderma spp.,the decrease of plant height and needle length was weakened,and the drought resistance of seedling was significantly enhanced( P < 0. 01).( 2) With the increase of drought stress,the soluble protein and starch content of needles in P. tabulaeformis seedlings continued to decrease. After inoculation with Trichoderma spp.,the corresponding index showed a decreasing trend with increasing stress,but the trend of inoculated Trichoderma spp. seedlings was weaker than that of uninoculated seedlings.( P < 0. 01).( 3) Under drought stress,the soluble sugar content of needles in P. tabulaeformis seedlings showed a trend of "first rise and then fall". The soluble sugar content of seedlings uninoculated and inoculated Trichoderma spp.reached the highest value in LD and MD,respectively. In other words,the drought tolerance threshold of the seedlings increased. [Conclusions]The results showed that under drought stress,ectomycorrhizal fungi promoted the growth of P. tabulaeformis seedlings. Under moderate drought stress,Trichoderma spp. had the most significant effect on the growth of P. tabulaeformis seedlings. Under the same water treatment,the mycorrhizal seedlings had stronger drought resistance than non-mycorrhizal seedlings.
[Background] It researched the effect of ectomycorrhizal fungi on Pinus tabulaeformis seedlings to increase the survival rate of seedlings,promote the regeneration of tree species,and prevent soil erosion. [Methods] Potted water was used to simulate drought stress,and four water treatments were designed: control( CK),light drought( LD),moderate drought( MD),severe drought( SD).Inoculating the seedlings of Pinus tabulaeformis with Trichoderma spp. was to study the effect of growth which ectomycorrhizal fungi had on P. tabulaeformis seedling under drought stress. To each treatment,It selected seedlings whose growth was basically the same, and measured the morphological and physiological indexes. The specific operation refered to Principles and Techniques of Plant Physiological and Biochemical Experiments written by Li Hesheng. [Results] The plant height and needle length of P. tabulaeformis seedlings were significantly affected by inoculation with Trichoderma spp. and drought stress treatment( P < 0. 01),but the interaction was not significant( P > 0. 05). The effects of inoculation with Trichoderma spp. and drought stress on the soluble protein,soluble sugar and starch content of P. tabulaeformis seedlings were significant( P < 0. 01). However,the interaction to the soluble protein and starch content of Pi. tabulaeformis seedlings was not significant( P > 0. 05),and the interaction to the soluble sugar content was significantly affected( P < 0. 01). The detailed analysis results to the morphological and physiological indexes of P. tabulaeformis seedlings were as follows:( 1)After inoculation with Trichoderma spp.,the biomass and root-shoot ratio of P. tabulaeformis seedlings were significantly increased( P < 0. 05). The mycorrhizal dependence on Trichoderma spp. was moderately dependent. Under drought stress,the plant height and needle length of P. tabulaeformis seedlings showed a decreasing trend. However,after inoculation with Trichoderma spp.,the decrease of plant height and needle length was weakened,and the drought resistance of seedling was significantly enhanced( P < 0. 01).( 2) With the increase of drought stress,the soluble protein and starch content of needles in P. tabulaeformis seedlings continued to decrease. After inoculation with Trichoderma spp.,the corresponding index showed a decreasing trend with increasing stress,but the trend of inoculated Trichoderma spp. seedlings was weaker than that of uninoculated seedlings.( P < 0. 01).( 3) Under drought stress,the soluble sugar content of needles in P. tabulaeformis seedlings showed a trend of "first rise and then fall". The soluble sugar content of seedlings uninoculated and inoculated Trichoderma spp.reached the highest value in LD and MD,respectively. In other words,the drought tolerance threshold of the seedlings increased. [Conclusions]The results showed that under drought stress,ectomycorrhizal fungi promoted the growth of P. tabulaeformis seedlings. Under moderate drought stress,Trichoderma spp. had the most significant effect on the growth of P. tabulaeformis seedlings. Under the same water treatment,the mycorrhizal seedlings had stronger drought resistance than non-mycorrhizal seedlings.
引文
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[4]王艺,丁贵杰.干旱胁迫下外生菌根真菌对马尾松幼苗生长和微量元素吸收的影响[J].浙江农林大学学报,2012,29(6):826.WANG Yi,DING Guijie.Growth and microelement absorption for Pinus massoniana seedlings with ecto-mycorrhizae and water stress[J].Journal of Zhejiang A&FUniversity,2012,29(6):826.
[5]邹红海.接种外生菌根菌繁育苗木技术的应用[J].中国林副特产,2015(1):56.ZOU Honghai.Application of inoculation of ectomycorrhizal fungi breeding seedlings[J].Forest By-Product and Speciality in China,2015(1):56.
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[7]张凤翔.油松的育苗及造林技术[J].吉林农业,2014(24):73.ZHANG Fengxiang.Seeding and afforestation techniques of Pinus tabulaeformis[J].Jilin Agriculture,2014(24):73.
[8]ZHANG Haoqiang,TANG Ming,CHEN Hui,et al.Communities of arbuscular mycorrhizal fungi and bacteria in the rhizosphere of Caragana korshinkii and Hippophae rhamnoides in Zhifanggou watershed[J].Plant&Soil,2010,326(1/2):415.
[9]WINDHAM MT,ELAD Y,BAKER R.A mechanism for increased plant growth induced by Trichoderma spp[J].Phytopathology,1986,76(5):520.
[10]林武星,黄雍容,朱炜,等.干旱胁迫对台湾栾树幼苗生长和生理生化指标的影响[J].中国水土保持科学,2014,12(5):54.LIN Wuxing,HUANG Yongrong,ZHU Wei,et al.Effects of drought stress on the growth and physiological and biochemical characteristics of Koelreuteria elegans seedlings[J].Science of Soil&Water Conservation,2014,12(5):54.
[11]陈志成,王志伟,王荣荣,等.3种阔叶树种对持续干旱的生理响应及抗旱性评价[J].中国水土保持科学,2013,11(2):67.CHEN Zhicheng,WANG Zhiwei,WANG Rongrong,et al.Physiological response of three broadleaved tree species to drought stress and evaluation of drought resistance[J].Science of Soil&Water Conservation,2013,11(2):67.
[12]弓明钦,王凤珍,陈羽,等.西南桦对菌根的依赖性及其接种效应研究[J].林业科学研究,2000,13(1):8.GONG Mingqian,WANG Fengzhen,CHEN Yu,et al.Study on the dependence of Betula alnoides on mycorrhiza and its inoculation effect[J].Forest Research,2000,13(1):8.
[13]李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000:183.LI Hesheng.Principles and techniques of plant physiological and biochemical experiments[M].Beijing:Higher Education Press,2000:183.
[14]刘锦春,钟章成,何跃军.干旱胁迫及复水对喀斯特地区柏木幼苗活性氧清除系统的影响[J].应用生态学报,2011,22(11):2838.LIU Jinchun,ZHONG Zhangcheng,HE Yuejun.Effects of drought stress and re-watering on the active oxygen scavenging system of Cupressus funebris seedlings in Karst area[J].Chinese Journal of Applied Ecology,2011,22(11):2838.
[15]董丽佳,桑卫国.模拟增温和降水变化对北京东灵山辽东栎种子出苗和幼苗生长的影响[J].植物生态学报,2012,36(8):819.DONG Lijia,SANG Weiguo.Effects of simulated warming and precipitation changes on seed emergence and seedling growth of Quercus mongolica in Dongling Mountain,Beijing,China[J].Chinese Journal of Plant E-cology,2012,36(8):819.
[16]简在友,许桂芳,孟丽.微量元素对红豆杉菌根菌生长的影响[J].生态环境,2007,16(1):201.JIAN Zaiyou,XU Guifang,MENG Li.Effect of trace elements on growth of Taxus chinensis var.mairei mycorrhizal fungi[J].Ecology and Environment,2007,16(1):201.
[17]童琳,唐旭利,张静,等.菌根形成对不同成熟度的森林优势树种磷吸收的影响[J].生态科学,2015,34(4):96.TONG Lin,TANG Xuli,ZHANG Jing,et al.Influence of mycorrhizal formulation on phosphorus acquisition of dominant tree species in young forest and old-growth forest[J].Ecological Science,2015,34(4):96.
[18]韩旭,宋述尧.矮生菜豆叶片衰老过程中碳氮代谢指标的变化[J].长江蔬菜,2009(10):42.HAN Xu,SONG Shurao.Changes of carbon and nitrogen metabolism indexes in the process of leaf senescence of dwarf bean[J].Journal of Changjiang Vegetables,2009(10):42.
[19]刘灵娣,李存东,孙红春,等.干旱胁迫对不同铃重基因型棉花叶片碳水化合物代谢的影响[C].沈阳:全国作物生理学研讨会,2006(9):252.LIU Lingdi,LI Cundong,SUN Hongchun,et al.Effect of water stress on carbohydrate metabolism of different boll weight genotypes in cotton[C].Shenyang:National Crop Physiology Symposium,2006(9):252.
[20]MAI Li,GUNTER Hoch,CHRISTIAN Korner.Source/sink removal affects mobile carbohydrates in Pinus cembra at the Swiss treeline[J].Trees,2002,16(4/5):335.
[21]郭华军.水分胁迫过程中的渗透调节物质及其研究进展[J].安徽农业科学,2010,38(15):7751.GUO Huajun.Research progress on osmotic adjustment material under water stress[J].Journal of Anhui Agricultural Sciences,2010,38(15):7751.
[22]潘昕,邱权,李吉跃,等.干旱胁迫对青藏高原6种植物生理指标的影响[J].生态学报,2014,34(13):3558.PAN Xin,QIU Quan,LI Jiyue,et al.Physiological indexes of six plant species from the Tibetan Plateau under drought stress[J].Acta Ecologica Sinica,2014,34(13):3558.
[2]高润梅,石晓东,王林,等.当年生华北落叶松幼苗的耐旱性[J].林业科学,2015,51(7):149.GAO Runmei,SHI Xiaodong,WANG Lin,et al.Drought resistance of one-year-old seedlings of Larix principis-rupprechtii[J].Scientia Silvae Sinicae,2015,51(7):149.
[3]王巧,刘秀梅,王华田,等.干旱和水涝胁迫对幼龄油松生长及光合作用的影响[J].中国水土保持科学,2015,13(6):40.WANG Qiao,LIU Xiumei,WANG Huatian,et al.Effects of drought and waterlogging on growth and photosynthesis of potted young Pinus tabulaeformis[J].Science of Soil and Water Conservation,2015,13(6):40.
[4]王艺,丁贵杰.干旱胁迫下外生菌根真菌对马尾松幼苗生长和微量元素吸收的影响[J].浙江农林大学学报,2012,29(6):826.WANG Yi,DING Guijie.Growth and microelement absorption for Pinus massoniana seedlings with ecto-mycorrhizae and water stress[J].Journal of Zhejiang A&FUniversity,2012,29(6):826.
[5]邹红海.接种外生菌根菌繁育苗木技术的应用[J].中国林副特产,2015(1):56.ZOU Honghai.Application of inoculation of ectomycorrhizal fungi breeding seedlings[J].Forest By-Product and Speciality in China,2015(1):56.
[6]刘润进,陈应龙.菌根学[M].北京:科学出版社,2007:161.LIU Runjin,CHEN Yinglong.Mycorrhizology[M].Beijing:Science Press,2007:161.
[7]张凤翔.油松的育苗及造林技术[J].吉林农业,2014(24):73.ZHANG Fengxiang.Seeding and afforestation techniques of Pinus tabulaeformis[J].Jilin Agriculture,2014(24):73.
[8]ZHANG Haoqiang,TANG Ming,CHEN Hui,et al.Communities of arbuscular mycorrhizal fungi and bacteria in the rhizosphere of Caragana korshinkii and Hippophae rhamnoides in Zhifanggou watershed[J].Plant&Soil,2010,326(1/2):415.
[9]WINDHAM MT,ELAD Y,BAKER R.A mechanism for increased plant growth induced by Trichoderma spp[J].Phytopathology,1986,76(5):520.
[10]林武星,黄雍容,朱炜,等.干旱胁迫对台湾栾树幼苗生长和生理生化指标的影响[J].中国水土保持科学,2014,12(5):54.LIN Wuxing,HUANG Yongrong,ZHU Wei,et al.Effects of drought stress on the growth and physiological and biochemical characteristics of Koelreuteria elegans seedlings[J].Science of Soil&Water Conservation,2014,12(5):54.
[11]陈志成,王志伟,王荣荣,等.3种阔叶树种对持续干旱的生理响应及抗旱性评价[J].中国水土保持科学,2013,11(2):67.CHEN Zhicheng,WANG Zhiwei,WANG Rongrong,et al.Physiological response of three broadleaved tree species to drought stress and evaluation of drought resistance[J].Science of Soil&Water Conservation,2013,11(2):67.
[12]弓明钦,王凤珍,陈羽,等.西南桦对菌根的依赖性及其接种效应研究[J].林业科学研究,2000,13(1):8.GONG Mingqian,WANG Fengzhen,CHEN Yu,et al.Study on the dependence of Betula alnoides on mycorrhiza and its inoculation effect[J].Forest Research,2000,13(1):8.
[13]李合生.植物生理生化实验原理和技术[M].北京:高等教育出版社,2000:183.LI Hesheng.Principles and techniques of plant physiological and biochemical experiments[M].Beijing:Higher Education Press,2000:183.
[14]刘锦春,钟章成,何跃军.干旱胁迫及复水对喀斯特地区柏木幼苗活性氧清除系统的影响[J].应用生态学报,2011,22(11):2838.LIU Jinchun,ZHONG Zhangcheng,HE Yuejun.Effects of drought stress and re-watering on the active oxygen scavenging system of Cupressus funebris seedlings in Karst area[J].Chinese Journal of Applied Ecology,2011,22(11):2838.
[15]董丽佳,桑卫国.模拟增温和降水变化对北京东灵山辽东栎种子出苗和幼苗生长的影响[J].植物生态学报,2012,36(8):819.DONG Lijia,SANG Weiguo.Effects of simulated warming and precipitation changes on seed emergence and seedling growth of Quercus mongolica in Dongling Mountain,Beijing,China[J].Chinese Journal of Plant E-cology,2012,36(8):819.
[16]简在友,许桂芳,孟丽.微量元素对红豆杉菌根菌生长的影响[J].生态环境,2007,16(1):201.JIAN Zaiyou,XU Guifang,MENG Li.Effect of trace elements on growth of Taxus chinensis var.mairei mycorrhizal fungi[J].Ecology and Environment,2007,16(1):201.
[17]童琳,唐旭利,张静,等.菌根形成对不同成熟度的森林优势树种磷吸收的影响[J].生态科学,2015,34(4):96.TONG Lin,TANG Xuli,ZHANG Jing,et al.Influence of mycorrhizal formulation on phosphorus acquisition of dominant tree species in young forest and old-growth forest[J].Ecological Science,2015,34(4):96.
[18]韩旭,宋述尧.矮生菜豆叶片衰老过程中碳氮代谢指标的变化[J].长江蔬菜,2009(10):42.HAN Xu,SONG Shurao.Changes of carbon and nitrogen metabolism indexes in the process of leaf senescence of dwarf bean[J].Journal of Changjiang Vegetables,2009(10):42.
[19]刘灵娣,李存东,孙红春,等.干旱胁迫对不同铃重基因型棉花叶片碳水化合物代谢的影响[C].沈阳:全国作物生理学研讨会,2006(9):252.LIU Lingdi,LI Cundong,SUN Hongchun,et al.Effect of water stress on carbohydrate metabolism of different boll weight genotypes in cotton[C].Shenyang:National Crop Physiology Symposium,2006(9):252.
[20]MAI Li,GUNTER Hoch,CHRISTIAN Korner.Source/sink removal affects mobile carbohydrates in Pinus cembra at the Swiss treeline[J].Trees,2002,16(4/5):335.
[21]郭华军.水分胁迫过程中的渗透调节物质及其研究进展[J].安徽农业科学,2010,38(15):7751.GUO Huajun.Research progress on osmotic adjustment material under water stress[J].Journal of Anhui Agricultural Sciences,2010,38(15):7751.
[22]潘昕,邱权,李吉跃,等.干旱胁迫对青藏高原6种植物生理指标的影响[J].生态学报,2014,34(13):3558.PAN Xin,QIU Quan,LI Jiyue,et al.Physiological indexes of six plant species from the Tibetan Plateau under drought stress[J].Acta Ecologica Sinica,2014,34(13):3558.