生物炭对华北冬小麦根系形态和内生真菌多样性的影响
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摘要
利用棉花秸秆生物炭,以冬小麦为试验对象,采用根系扫描法和Illumina MiSeq高通量测序技术探讨研究农田秸秆还田,施生物炭分别为4.5、9.0、13.5 t/hm~2后根系形态与内生真菌多样性的影响及其共生机制。试验结果表明,在生物炭施用量为4.5~13.5 t/hm~2时,在冬小麦成熟期提高初生根的直径和比根长,降低分支密度和生物量;降低次生根分支密度。在门水平,生物炭显著提高根内子囊菌门(Ascomycota)丰度,显著降低接合菌门(Zygomycota)、壶菌门(Chytridiomycota)和未鉴定杂菌(unidentified fungus)丰度,对球囊菌门影响不显著;在常见目水平,生物炭明显提高格孢菌目(Pleosporales)丰度,显著降低散囊菌目(Eurotiales)和被孢霉目(Mortierellales)丰度。在4.5~9.0 t/hm~2时,生物炭通过降低初生根内散囊菌目真菌丰度,促进初生根分支密度和生物量的生长;通过提高次生根内格孢菌目真菌丰度,抑制次生根直径和分支密度的生长。经综合比较,9.0 t/hm~2的生物炭处理对根系形态的优化效果优于其他处理,与对照处理相比,在成熟期,初生根直径和比根长分别提高5.00%、33.57%,分支密度和生物量分别降低67.26%、27.27%;次生根直径和分支密度分别降低13.16%、34.38%。
The winter wheat was selected to study the effect of biochar of cotton straw on root morphology and endophytic fungal communities by using the method of root scanning and Illumina Miseq highthroughput sequencing. The symbiosis mechanism was found by the test. The treatments were amount of 4. 5 t/hm~2,9. 0 t/hm~2 and 13. 5 t/hm~2 of straw biochar,respectively applied to the soil,and the straw returning was taken as test control in the field experiment. The results showed that root diameter and specific root length were increased significantly,and branching density and biomass were decreased significantly in the primary root,and only the root branching density was decreased in the secondary root,when the amount of biochar was 4. 5 ~ 13. 5 t/hm~2 in the mature period of winter wheat. At the fungal phyla level, the abundance of Ascomycota was increased significantly, and the abundances of Zygomycota,Chytridiomycota and unidentified fungus was reduced greatly, and the abundance of Glomeromycota was not significant. At the common fungal order level,the abundance of Pleosporales was increased significantly,and the abundance of Eurotiales and Mortierellales was reduced greatly. With the amount of 4. 5 ~ 9. 0 t/hm~2 biochar promoted the primary root growth of root branch density and biomass by decreasing the abundance of Eurotiales,and inhibited the secondary root growth of diameter and branch density by increasing the abundance of Pleosporales. By comprehensive comparison,the effect ofamount of 9. 0 t/hm~2 biochar on root morphology was better than that of other treatments. The treatment with amount of 9. 0 t/hm~2 significantly increased the primary root diameter by 5. 00% and specific root length by 33. 57%,and decreased root branch density by 67. 26% and biomass by 27. 27%,while also reduced the secondary root diameter by 13. 16% and branch density by 34. 38%,compared with the test control at maturity stage of winter wheat.
The winter wheat was selected to study the effect of biochar of cotton straw on root morphology and endophytic fungal communities by using the method of root scanning and Illumina Miseq highthroughput sequencing. The symbiosis mechanism was found by the test. The treatments were amount of 4. 5 t/hm~2,9. 0 t/hm~2 and 13. 5 t/hm~2 of straw biochar,respectively applied to the soil,and the straw returning was taken as test control in the field experiment. The results showed that root diameter and specific root length were increased significantly,and branching density and biomass were decreased significantly in the primary root,and only the root branching density was decreased in the secondary root,when the amount of biochar was 4. 5 ~ 13. 5 t/hm~2 in the mature period of winter wheat. At the fungal phyla level, the abundance of Ascomycota was increased significantly, and the abundances of Zygomycota,Chytridiomycota and unidentified fungus was reduced greatly, and the abundance of Glomeromycota was not significant. At the common fungal order level,the abundance of Pleosporales was increased significantly,and the abundance of Eurotiales and Mortierellales was reduced greatly. With the amount of 4. 5 ~ 9. 0 t/hm~2 biochar promoted the primary root growth of root branch density and biomass by decreasing the abundance of Eurotiales,and inhibited the secondary root growth of diameter and branch density by increasing the abundance of Pleosporales. By comprehensive comparison,the effect ofamount of 9. 0 t/hm~2 biochar on root morphology was better than that of other treatments. The treatment with amount of 9. 0 t/hm~2 significantly increased the primary root diameter by 5. 00% and specific root length by 33. 57%,and decreased root branch density by 67. 26% and biomass by 27. 27%,while also reduced the secondary root diameter by 13. 16% and branch density by 34. 38%,compared with the test control at maturity stage of winter wheat.
引文
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2强学彩.秸秆还田量的农田生态效应研究[D].北京:中国农业大学,2003.QIANG Xuecai.Study on ecological effect in farm system of different amount of crop residues incorporation[D].Beijing:China Agricultural University,2003.(in Chinese)
3李素娟,陈继康,陈阜,等.华北平原免耕冬小麦生长发育特征研究[J].作物学报,2008,34(2):290-296.LI Sujuan,CHEN Jikang,CHEN Fu,et al.Characteristics of growth and development of winter wheat under zero-tillage in North China Plain[J].Acta Agrononica Sinica,2008,34(2):290-296.(in Chinese)
4 QI Y,ZHEN W,LI H.Allelopathy of decomposed maize straw products on three soil-born diseases of wheat and the analysis by GC-MS[J].Journal of Integrative Agriculture,2015,14(1):88-97.
5甄文超.玉米秸秆还田对冬小麦土传病害影响的研究[C]∥中国植物病理学会2009年学术年会论文集,2009.ZHEN Wenchao.The influence of maize straw amendment on the soil-borne disease of winter wheat[C]∥Chinese Proceedings of the Annual Meeting of Chinese Society for Plant Pathology,2009.(in Chinese)
6 PETRINI O.Fungal endophytes of tree leaves[M]∥ANDREWS J H,HIRANO S S.Microbial ecology of leaves.New York:Springer-Vedag,1991:179-197.
7 SMITH S E,READ D J.Mycorrhizal symbiosis[M].Cambridge,UK:Academic Press,2008.
8 WARNOCK D D,LEHMANN J,KUYPER T W,et al.Mycorrhizal responses to biochar in soil-concepts and mechanisms[J].Plant and Soil,2007,300(1-2):9-20.
9 BLACKWELL P.Can biochar help drive biological farming?[J].Western Mineral Fertilisers,2008(8):11.
10 SOLAIMAN Z M,BLACKWELL P,ABBOTT L K,et al.Direct and residual effect of biochar application on mycorrhizal root colonisation,growth and nutrition of wheat[J].Australian Journal of Soil Research,2010,48(7):546-554.
11 RILLIG M C,WAGNER M,SALEM M,et al.Material derived from hydrothermal carbonization:effects on plant growth and arbuscular mycorrhiza[J].Applied Soil Ecology,2010,45(3):238-242.
12 HAMMER E C,BALOGH-BRUNSTAD Z,JAKOBSEN I,et al.A mycorrhizal fungus grows on biochar and captures phosphorus from its surfaces[J].Soil Biology&Biochemistry,2014,77:252-260.
13 BLACKWELL P,KRULL E,BUTLER G,et al.Effect of banded biochar on dryland wheat production and fertiliser use in southwestern Australia:an agronomic and economic perspective[J].Australian Journal of Soil Research,2010,48(7):531-545.
14 WALLER F,AEHATZ B,BALTRUSNHAT H,et al.The endophytic fungus Piriformospora indica reprograms barley to salt-stress tolerance,disease resistance,and higher yield[J].Proceedings of the National Academy of Sciences,2005,102(38):13386-13391.
15 RODRIGUEZ R J,HENSON J M,VAN VOLKENBURGH E,et al.Stress tolerance in plants via habitat-adapted symbiosis[J].International Society for Microbial Ecology,2008,2:404-416.
16 MALINOWSKI D P,ALLOUSH G A,BELESKY D P.Evidence for chemical changes on the root surface of tall fescue in response to infection with the fungal endophyte Neotyphodium coenophialum[J].Plant and Soil,1998,205(1):1-12.
17 AZEVEDO M D,WEHY R E.A study of the funsal endophyte Acremonium coenophialum in the roots of tall fescue seedlings[J].Mycologia,1995,87(3):289-297.
18 RAFUNAN M H,SAIGA S.Endophytic fungi(Neotyphodium coenophialum)affect the growth and mineral uptake,transport and efficiency ratios in tall fescue(Festuca arundinacea)[J].Plant and Soil,2005,272(1-2):163-171.
19 BLANKE V,RENKER C,WANGER M,et al.Nitrogen supply affects arbuscular mycorrhizal colonization of Artemisia vulgaris in a phosphate polluted field site[J].New Phytologist,2005,166(3):981-992.
20 OMACINI M,SEMMARTIN M,PEREZ L I,et al.Grass-endophytic symbiosis:a neglected aboveground interaction with multiple belowground consequences[J].Applied Soil Ecology,2012,61:273-279.
21马元喜.小麦的根[M].北京:中国农业出版社,1999.
22 TYSLENKO A M.Effect of root number on yield in spring wheat[J].Byulleton Vsesoyuznogo Ordena Lenina Iordena Druzhby Narodov Institute Rastenievodstva Imenin I Vavibva,1981,114:15-17.
23 TIHONOV V E.The role of the number of embryonic roots in spring bread wheats in the semidesert conditions of northern priarale[J].Byulleton Vsesoyuznogo Ordena Lenina Institute Rastenievodstva Imenin I Vavibva,1973,33:3-7.
24薛丽华,段俊杰,王志敏,等.冬小麦调亏灌溉下次生根和初生根对植株生长的影响[J].华北农学报,2010,25(3):87-91.XUE Lihua,DUAN Junjie,WANG Zhimin,et al.The effect of secondary and seminal roots of winter wheat to plant growth under regulated deficit irrigation conditions[J].Acta Agriculturae Boreali-Sinica,2010,25(3):87-91.(in Chinese)
25贾银锁,张锦熙,刘钖山,等.冬小麦不同单位根的功能及分组的研究[J].核农学报,1990,4(1):25-30.JIA Yinsuo,ZHANG Jinxi,LIU Yangshan,et al.Study on the functions of different unit-roots and the grouping of roots in winter wheat[J].Acta Agriculturae Nucleatae Sinica,1990,4(1):25-30.(in Chinese)
26 SUN J Q,GUO L D,ZANG W,et al.Diversity and ecological distribution of endophytic fungi associated with medicinal plants[J].Science in China Series C:Life Sciences,2008,51:751-759.
27 CARON D A,COUNTWAY P D,SAVAI P,et al.Defining DNA-based operational taxonomic units for microbial-eukaryote ecology[J].Applied and environment Microbiology,2009,75(18):5797-5808.
28李中阳,齐学斌,樊向阳,等.生物质炭对冬小麦产量、水分利用效率及根系形态的影响[J].农业工程学报,2015,31(12):119-124.LI Zhongyang,QI Xuebin,FAN Xiangyang,et al.Influences of biochars on growth,yield,water use efficiency and root morphology of winter wheat[J].Transactions of the CSAE,2015,31(12):119-124.(in Chinese)
29张伟明,孟军,王嘉宇,等.生物炭对水稻根系形态与生理特性及产量的影响[J].作物学报,2013,39(8):1445-1451.ZHANG Weiming,MENG Jun,WANG Jiayu,et al.Effect of biochar on root morphological and physiological characteristics and yield in rice[J].Acta Agronomica Sinica,2013,39(8):1445-1451.(in Chinese)
30 ABIVEN S,HUND A,MARTINSEN V,et al.Biochar amendment increases maize root surface areas and branching:a shovelomics study in Zambia[J].Plant Soil,2015,395:45-55.
31李发虎,李明,刘金泉,等.生物炭对温室黄瓜根际土壤真菌丰度和根系生长的影响[J/OL].农业机械学报,2017,48(4):265-270,341.http:∥www.j-csam.org/jcsam/ch/reader/view_abstract.aspx?flag=1&file_no=20170434&journal_id=jcsam.DOI:10.6041/j.issn.1000-1298.2017.04.034.LI Fahu,LI Ming,LIU Jinquan,et al.Effect of biochar on fungal abundance of rhizosphere soil and cucumber root growth in greenhouse[J/OL].Transactions of the Chinese Society for Agricultural Machinery,2017,48(4):265-270,341.(in Chinese)
32李明,胡云,黄修梅,等.生物炭对设施黄瓜根际土壤养分和菌群的影响[J/OL].农业机械学报,2016,47(11):172-178.http:∥www.j-csam.org/jcsam/ch/reader/view_abstract.aspx?flag=1&file_no=20161123&journal_id=jcsam.DOI:10.6041/j.issn.1000-1298.2016.11.023.LI Ming,HU Yun,HUANG Xiumei,et al.Effect of biological carbon on nutrient and bacterial communities of rhizosphere soil of facility cucumber[J/OL].Transactions of the Chinese Society for Agricultural Machinery,2016,47(11):172-178.(in Chinese)
2强学彩.秸秆还田量的农田生态效应研究[D].北京:中国农业大学,2003.QIANG Xuecai.Study on ecological effect in farm system of different amount of crop residues incorporation[D].Beijing:China Agricultural University,2003.(in Chinese)
3李素娟,陈继康,陈阜,等.华北平原免耕冬小麦生长发育特征研究[J].作物学报,2008,34(2):290-296.LI Sujuan,CHEN Jikang,CHEN Fu,et al.Characteristics of growth and development of winter wheat under zero-tillage in North China Plain[J].Acta Agrononica Sinica,2008,34(2):290-296.(in Chinese)
4 QI Y,ZHEN W,LI H.Allelopathy of decomposed maize straw products on three soil-born diseases of wheat and the analysis by GC-MS[J].Journal of Integrative Agriculture,2015,14(1):88-97.
5甄文超.玉米秸秆还田对冬小麦土传病害影响的研究[C]∥中国植物病理学会2009年学术年会论文集,2009.ZHEN Wenchao.The influence of maize straw amendment on the soil-borne disease of winter wheat[C]∥Chinese Proceedings of the Annual Meeting of Chinese Society for Plant Pathology,2009.(in Chinese)
6 PETRINI O.Fungal endophytes of tree leaves[M]∥ANDREWS J H,HIRANO S S.Microbial ecology of leaves.New York:Springer-Vedag,1991:179-197.
7 SMITH S E,READ D J.Mycorrhizal symbiosis[M].Cambridge,UK:Academic Press,2008.
8 WARNOCK D D,LEHMANN J,KUYPER T W,et al.Mycorrhizal responses to biochar in soil-concepts and mechanisms[J].Plant and Soil,2007,300(1-2):9-20.
9 BLACKWELL P.Can biochar help drive biological farming?[J].Western Mineral Fertilisers,2008(8):11.
10 SOLAIMAN Z M,BLACKWELL P,ABBOTT L K,et al.Direct and residual effect of biochar application on mycorrhizal root colonisation,growth and nutrition of wheat[J].Australian Journal of Soil Research,2010,48(7):546-554.
11 RILLIG M C,WAGNER M,SALEM M,et al.Material derived from hydrothermal carbonization:effects on plant growth and arbuscular mycorrhiza[J].Applied Soil Ecology,2010,45(3):238-242.
12 HAMMER E C,BALOGH-BRUNSTAD Z,JAKOBSEN I,et al.A mycorrhizal fungus grows on biochar and captures phosphorus from its surfaces[J].Soil Biology&Biochemistry,2014,77:252-260.
13 BLACKWELL P,KRULL E,BUTLER G,et al.Effect of banded biochar on dryland wheat production and fertiliser use in southwestern Australia:an agronomic and economic perspective[J].Australian Journal of Soil Research,2010,48(7):531-545.
14 WALLER F,AEHATZ B,BALTRUSNHAT H,et al.The endophytic fungus Piriformospora indica reprograms barley to salt-stress tolerance,disease resistance,and higher yield[J].Proceedings of the National Academy of Sciences,2005,102(38):13386-13391.
15 RODRIGUEZ R J,HENSON J M,VAN VOLKENBURGH E,et al.Stress tolerance in plants via habitat-adapted symbiosis[J].International Society for Microbial Ecology,2008,2:404-416.
16 MALINOWSKI D P,ALLOUSH G A,BELESKY D P.Evidence for chemical changes on the root surface of tall fescue in response to infection with the fungal endophyte Neotyphodium coenophialum[J].Plant and Soil,1998,205(1):1-12.
17 AZEVEDO M D,WEHY R E.A study of the funsal endophyte Acremonium coenophialum in the roots of tall fescue seedlings[J].Mycologia,1995,87(3):289-297.
18 RAFUNAN M H,SAIGA S.Endophytic fungi(Neotyphodium coenophialum)affect the growth and mineral uptake,transport and efficiency ratios in tall fescue(Festuca arundinacea)[J].Plant and Soil,2005,272(1-2):163-171.
19 BLANKE V,RENKER C,WANGER M,et al.Nitrogen supply affects arbuscular mycorrhizal colonization of Artemisia vulgaris in a phosphate polluted field site[J].New Phytologist,2005,166(3):981-992.
20 OMACINI M,SEMMARTIN M,PEREZ L I,et al.Grass-endophytic symbiosis:a neglected aboveground interaction with multiple belowground consequences[J].Applied Soil Ecology,2012,61:273-279.
21马元喜.小麦的根[M].北京:中国农业出版社,1999.
22 TYSLENKO A M.Effect of root number on yield in spring wheat[J].Byulleton Vsesoyuznogo Ordena Lenina Iordena Druzhby Narodov Institute Rastenievodstva Imenin I Vavibva,1981,114:15-17.
23 TIHONOV V E.The role of the number of embryonic roots in spring bread wheats in the semidesert conditions of northern priarale[J].Byulleton Vsesoyuznogo Ordena Lenina Institute Rastenievodstva Imenin I Vavibva,1973,33:3-7.
24薛丽华,段俊杰,王志敏,等.冬小麦调亏灌溉下次生根和初生根对植株生长的影响[J].华北农学报,2010,25(3):87-91.XUE Lihua,DUAN Junjie,WANG Zhimin,et al.The effect of secondary and seminal roots of winter wheat to plant growth under regulated deficit irrigation conditions[J].Acta Agriculturae Boreali-Sinica,2010,25(3):87-91.(in Chinese)
25贾银锁,张锦熙,刘钖山,等.冬小麦不同单位根的功能及分组的研究[J].核农学报,1990,4(1):25-30.JIA Yinsuo,ZHANG Jinxi,LIU Yangshan,et al.Study on the functions of different unit-roots and the grouping of roots in winter wheat[J].Acta Agriculturae Nucleatae Sinica,1990,4(1):25-30.(in Chinese)
26 SUN J Q,GUO L D,ZANG W,et al.Diversity and ecological distribution of endophytic fungi associated with medicinal plants[J].Science in China Series C:Life Sciences,2008,51:751-759.
27 CARON D A,COUNTWAY P D,SAVAI P,et al.Defining DNA-based operational taxonomic units for microbial-eukaryote ecology[J].Applied and environment Microbiology,2009,75(18):5797-5808.
28李中阳,齐学斌,樊向阳,等.生物质炭对冬小麦产量、水分利用效率及根系形态的影响[J].农业工程学报,2015,31(12):119-124.LI Zhongyang,QI Xuebin,FAN Xiangyang,et al.Influences of biochars on growth,yield,water use efficiency and root morphology of winter wheat[J].Transactions of the CSAE,2015,31(12):119-124.(in Chinese)
29张伟明,孟军,王嘉宇,等.生物炭对水稻根系形态与生理特性及产量的影响[J].作物学报,2013,39(8):1445-1451.ZHANG Weiming,MENG Jun,WANG Jiayu,et al.Effect of biochar on root morphological and physiological characteristics and yield in rice[J].Acta Agronomica Sinica,2013,39(8):1445-1451.(in Chinese)
30 ABIVEN S,HUND A,MARTINSEN V,et al.Biochar amendment increases maize root surface areas and branching:a shovelomics study in Zambia[J].Plant Soil,2015,395:45-55.
31李发虎,李明,刘金泉,等.生物炭对温室黄瓜根际土壤真菌丰度和根系生长的影响[J/OL].农业机械学报,2017,48(4):265-270,341.http:∥www.j-csam.org/jcsam/ch/reader/view_abstract.aspx?flag=1&file_no=20170434&journal_id=jcsam.DOI:10.6041/j.issn.1000-1298.2017.04.034.LI Fahu,LI Ming,LIU Jinquan,et al.Effect of biochar on fungal abundance of rhizosphere soil and cucumber root growth in greenhouse[J/OL].Transactions of the Chinese Society for Agricultural Machinery,2017,48(4):265-270,341.(in Chinese)
32李明,胡云,黄修梅,等.生物炭对设施黄瓜根际土壤养分和菌群的影响[J/OL].农业机械学报,2016,47(11):172-178.http:∥www.j-csam.org/jcsam/ch/reader/view_abstract.aspx?flag=1&file_no=20161123&journal_id=jcsam.DOI:10.6041/j.issn.1000-1298.2016.11.023.LI Ming,HU Yun,HUANG Xiumei,et al.Effect of biological carbon on nutrient and bacterial communities of rhizosphere soil of facility cucumber[J/OL].Transactions of the Chinese Society for Agricultural Machinery,2016,47(11):172-178.(in Chinese)