Article(id=1277293287593607193, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277293236137890180, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2024.11.010, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1715616000000, receivedDateStr=2024-05-14, revisedDate=1716998400000, revisedDateStr=2024-05-30, acceptedDate=null, acceptedDateStr=null, onlineDate=1782460050724, onlineDateStr=2026-06-26, pubDate=1732464000000, pubDateStr=2024-11-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782460050724, onlineIssueDateStr=2026-06-26, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782460050724, creator=13701087609, updateTime=1782460050724, updator=13701087609, issue=Issue{id=1277293236137890180, tenantId=1146029695717560320, journalId=1235980609244409860, year='2024', volume='45', issue='11', pageStart='2243', pageEnd='2486', issueExtLink='null', onlineDate='null', pubDate='1732464000000', pubDateStr='2024-11-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782460038455, creator='13701087609', updateTime=1782815269280, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1278783182988358204, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277293236137890180, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1278783182988358205, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1277293236137890180, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2320, endPage=2333, ext={EN=ArticleExt(id=1277293287912374299, articleId=1277293287593607193, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Advances in Sugarcane Breeding for Resistance to Sugarcane Mosaic Disease, columnId=1236256430060261740, journalTitle=Chinese Journal of Tropical Crops, columnName=Germplasm Resources, Genetics & Breeding, runingTitle=null, highlight=null, articleAbstract=

Sugarcane mosaic disease is one of the main viral diseases threatening the production of sugarcane. Sugarcane streak mosaic virus (SCSMV), Sorghum mosaic virus (SrMV) and Sugarcane mosaic virus (SCMV) are the main pathogens. The occurrence of sugarcane mosaic disease seriously affects the quality and yield of sugarcane. The most economical and effective way to control the disease is the application of resistant varieties. In this paper, the damage and controlling ways of sugarcane mosaic disease were summarized. The recent research progress on identification of resistance germplasm resources, resistance inheritance, molecular markers, mining and application of the resistance gene, and breeding of disease-resistant varieties were reviewed. Moreover, the problems existing in sugarcane disease resistance breeding were pointed out, and some suggestions were put forward to strengthen the research on sugarcane mosaic disease resistance breeding.

, authors=null, authorsList=Linbo SHEN, Jihong PAN, Huimin WANG, Xiaoyan FENG, Shuzhen ZHANG, Wenzhi WANG, authorCompany=null, correspAuthors=Wenzhi WANG, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1277293288839315485, articleId=1277293287593607193, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=甘蔗抗花叶病育种研究进展, columnId=1236256430219645304, journalTitle=热带作物学报, columnName=种质资源与遗传育种, runingTitle=null, highlight=null, articleAbstract=

甘蔗花叶病是危害甘蔗生产的主要病毒性病害之一,其病原主要包括甘蔗条纹花叶病毒(Sugarcane streak mosaic virus, SCSMV)、高粱花叶病毒(Sorghum mosaic virus, SrMV)和甘蔗花叶病毒(Sugarcane mosaic virus, SCMV)3大主要病原。甘蔗花叶病的发生严重影响甘蔗的品质和产量,防治该病害最经济有效的方法是抗病品种的应用。本文综述了甘蔗花叶病的危害和防治途径、甘蔗种质资源的抗性评价、抗性遗传分析、分子标记、抗性基因的挖掘利用及抗病育种等方面的研究进展,同时指出了甘蔗抗病育种方面存在的问题,并对加强甘蔗抗花叶病育种的研究提出了一些建议。

, authors=

沈林波(1986—),男,硕士,助理研究员,研究方向:甘蔗抗病分子育种。

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* 王文治(WANG Wenzhi),E-mail:
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沈林波(1986—),男,硕士,助理研究员,研究方向:甘蔗抗病分子育种。

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沈林波(1986—),男,硕士,助理研究员,研究方向:甘蔗抗病分子育种。

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(in Chinese), articleTitle=Construction of ihpRNA expression vector of CP genes of SCYLV and SCMV and genetic transformation in sugarcane, refAbstract=null), Reference(id=1277293320736997643, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[109], rfOrder=178, authorNames=徐景升, 杨永庆, 程光远, 高世武, 郭晋隆, 翟玉山, 彭磊, 邓宇晴, 郑艳茹, 许莉萍, journalName=null, refType=null, unstructuredReference=徐景升, 杨永庆, 程光远, 高世武, 郭晋隆, 翟玉山, 彭磊, 邓宇晴, 郑艳茹, 许莉萍. 利用RNAi沉默SceIF4E1基因培育抗条纹花叶病甘蔗的方法: CN201510570284.2[P]. 2015-09-09., articleTitle=利用RNAi沉默SceIF4E1基因培育抗条纹花叶病甘蔗的方法, refAbstract=null), Reference(id=1277293320799912204, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, doi=null, pmid=null, pmcid=null, year=null, volume=null, issue=null, pageStart=null, pageEnd=null, url=null, language=null, rfNumber=[109], rfOrder=179, authorNames=XU J S, YANG Y Q, CHENG G Y, GAO S W, GUO J L, ZHAI Y S, PENG L, DENG Y Q, ZHENG Y R, XU L P, journalName=null, refType=null, unstructuredReference=XU J S, YANG Y Q, CHENG G Y, GAO S W, GUO J L, ZHAI Y S, PENG L, DENG Y Q, ZHENG Y R, XU L P. The method for gene silencing SceIF4E1 gene by RNAi in sugarcane resistant to stripe sugarcane mosaic disease: CN201510570284.2[P]. 2015-09-09. (in Chinese), articleTitle=The method for gene silencing SceIF4E1 gene by RNAi in sugarcane resistant to stripe sugarcane mosaic disease, refAbstract=null), Reference(id=1277293320892186893, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, doi=null, pmid=null, pmcid=null, year=2018, volume=27, issue=2, pageStart=203, pageEnd=210, url=null, language=null, rfNumber=[110], rfOrder=180, authorNames=ASLAM U, TABASSUM B, NASIR I A, KHAN A, HUSNAIN T, journalName=Transgenic Research, refType=null, unstructuredReference=ASLAM U, TABASSUM B, NASIR I A, KHAN A, HUSNAIN T. A virus-derived short hairpin RNA confers resistance against Sugarcane mosaic virus in transgenic sugarcane[J]. Transgenic Research, 2018, 27(2): 203-210., articleTitle=A virus-derived short hairpin RNA confers resistance against Sugarcane mosaic virus in transgenic sugarcane, refAbstract=null), Reference(id=1277293321110290702, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, doi=null, pmid=null, pmcid=null, year=2021, volume=48, issue=null, pageStart=3047, pageEnd=3054, url=null, language=null, rfNumber=[111], rfOrder=181, authorNames=WIDYANINGRUM S, PUJIASIH D R, SHOLEHA W, HARMOKO R, SUGIHARTO B, journalName=Molecular Biology Reports, refType=null, unstructuredReference=WIDYANINGRUM S, PUJIASIH D R, SHOLEHA W, HARMOKO R, SUGIHARTO B. Induction of resistance to Sugarcane mosaic virus by RNA interference targeting coat protein gene silencing in transgenic sugarcane[J]. Molecular Biology Reports, 2021, 48: 3047-3054., articleTitle=Induction of resistance to Sugarcane mosaic virus by RNA interference targeting coat protein gene silencing in transgenic sugarcane, refAbstract=null)], funds=[Fund(id=1277293300574978133, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, awardId=CARS-17, language=CN, fundingSource=现代农业产业技术体系建设专项资金项目(CARS-17), fundOrder=null, country=null), Fund(id=1277293300654669910, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, awardId=2019RC300, language=CN, fundingSource=海南省基础与应用基础研究计划项目(2019RC300), fundOrder=null, country=null), Fund(id=1277293300730167383, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, awardId=1630052019019, language=CN, fundingSource=中央级公益性科研院所基本科研业务费专项(1630052019019), fundOrder=null, country=null)], companyList=[AuthorCompany(id=1277293289090973727, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, xref=1., ext=[AuthorCompanyExt(id=1277293289099362336, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, companyId=1277293289090973727, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.Institute of Tropical Bioscience and Biotechnology, Chinese Academy of Tropical Agricultural Sciences / Hainan Institute for Tropical Agricultural Resources, Haikou, Hainan 571101, China), AuthorCompanyExt(id=1277293289107750945, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, companyId=1277293289090973727, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=1.中国热带农业科学院热带生物技术研究所/海南热带农业资源研究院,海南海口 571101)]), AuthorCompany(id=1277293289179054114, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, xref=2., ext=[AuthorCompanyExt(id=1277293289187442723, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, companyId=1277293289179054114, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.College of Life Sciences, Nanjing Agricultural University, Nanjing, Jiangsu 210095, China), AuthorCompanyExt(id=1277293289200025636, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, companyId=1277293289179054114, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=2.南京农业大学生命科学学院,江苏南京 210095)]), AuthorCompany(id=1277293289304883238, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, xref=3., ext=[AuthorCompanyExt(id=1277293289313271847, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, companyId=1277293289304883238, language=EN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3.College of Agriculture, Nanjing Agricultural University, Nanjing, Jiangsu 210095, China), AuthorCompanyExt(id=1277293289317466152, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, companyId=1277293289304883238, language=CN, country=null, province=null, city=null, postcode=null, companyName=null, departmentName=null, remark=3.南京农业大学农学院,江苏南京 210095)])], figs=[ArticleFig(id=1277293298523963475, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, language=EN, label=Tab. 1, caption=

New hybrid varieties of sugarcance resistant to mosaic disease in China

, figureFileSmall=null, figureFileBig=null, tableContent=
品种名称Variety name抗病等级Disease resistance level亲本Parents选育单位Breeding institution
云蔗05-51中抗崖城90-56 ×ROC 23云南省农业科学院甘蔗研究所[62]
云蔗072178中抗桂糖92-66×Ho CP93-750云南省农业科学院甘蔗研究所[63]
云蔗072800中抗湛蔗92-126×CP88-1762云南省农业科学院甘蔗研究所[64]
云蔗08-1095中抗CP84-1198×科5云南省农业科学院甘蔗研究所[65]
云蔗081609高抗云蔗94-343×粤糖00-236云南省农业科学院甘蔗研究所[66]
云蔗94-375免疫Cp72-1210×崖城73-512云南省农业科学院甘蔗研究所[67]
云蔗99-91高抗新台糖10×崖城84-153云南省农业科学院甘蔗研究所[68]
福农15号高抗CP72-1210×科5福建农林大学甘蔗综合研究所[69]
福农38号高抗粤糖83-257×粤糖83-271福建农林大学甘蔗综合研究所[70]
福农39号中抗粤糖91-976×CP84-1198福建农林大学甘蔗综合研究所[71]
福农41号中抗ROC20×粤糖91-976福建农林大学甘蔗综合研究所[72]
福农81-745粤糖59-65×CP36-105福建农林大学甘蔗综合研究所[73]
福农83-36高抗CP49- 50×福农57-18福建农林大学甘蔗综合研究所[74]
福农91-3623免疫CP72-1210 ×桂73-167福建农林大学甘蔗综合研究所[75]
福农91-4621高抗CP72-1210 ×湛74-141福建农林大学甘蔗综合研究所[76]
福农94-0403中抗CP72-1210×闽糖69-263福建农林大学甘蔗综合研究所[77]
福农95-1702高抗CP72-1210×粤农73-204福建农林大学甘蔗综合研究所[78]
福农98-1103高抗CP72-1210×湛蔗74-141福建农林大学甘蔗综合研究所[79]
桂糖16号粤糖59/65×崖城72/399广西农业科学院甘蔗研究所[80]
桂糖23号高抗ROC 1号×崖城71-374广西农业科学院甘蔗研究所[81]
桂糖24号高抗桂糖71-5×崖城84-153广西农业科学院甘蔗研究所[82]
桂糖25号高抗CP72- 1210×崖城71- 374广西农业科学院甘蔗研究所[83]
桂糖42号高抗ROC 22×桂糖92-66广西农业科学院甘蔗研究所[84]
桂糖54号中抗桂糖00-122×崖城97-47广西农业科学院甘蔗研究所[85]
粤糖00-236高抗粤农73-204×CP72-1210广州甘蔗糖业研究所[86]
粤糖07913高抗HOCP 95-988×粤糖97-76广州甘蔗糖业研究所[87]
粤糖89-240CP72-1210×桂糖73-167广州甘蔗糖业研究所[88]
粤糖95-618免疫ROC10×CP72-1210广州甘蔗糖业研究所[89]
粤糖96-835高抗Co419×新台糖10号广州甘蔗糖业研究所[90]
闽糖88-103高抗闽选703×崖城82-96福建省农业科学院甘蔗研究所[91]
闽糖92-649高抗ROC1号×闽选703福建省农业科学院甘蔗研究所[92]
闽糖95-261高抗(ROC1×CP72-1210)×CP72-1210福建省农业科学院甘蔗研究所[93]
桂柳2号免疫ROC22×HoCP93-746柳城县甘蔗研究中心[94]
赣南02-70高抗桂糖69-435×CP84-1198江西省甘蔗研究所[95]
川蔗27号高抗CP72-1210×崖城90-3四川省植物工程研究院[96]
), ArticleFig(id=1277293298599460948, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1277293287593607193, language=CN, label=表1, caption=

我国甘蔗抗花叶病杂交新品种

, figureFileSmall=null, figureFileBig=null, tableContent=
品种名称Variety name抗病等级Disease resistance level亲本Parents选育单位Breeding institution
云蔗05-51中抗崖城90-56 ×ROC 23云南省农业科学院甘蔗研究所[62]
云蔗072178中抗桂糖92-66×Ho CP93-750云南省农业科学院甘蔗研究所[63]
云蔗072800中抗湛蔗92-126×CP88-1762云南省农业科学院甘蔗研究所[64]
云蔗08-1095中抗CP84-1198×科5云南省农业科学院甘蔗研究所[65]
云蔗081609高抗云蔗94-343×粤糖00-236云南省农业科学院甘蔗研究所[66]
云蔗94-375免疫Cp72-1210×崖城73-512云南省农业科学院甘蔗研究所[67]
云蔗99-91高抗新台糖10×崖城84-153云南省农业科学院甘蔗研究所[68]
福农15号高抗CP72-1210×科5福建农林大学甘蔗综合研究所[69]
福农38号高抗粤糖83-257×粤糖83-271福建农林大学甘蔗综合研究所[70]
福农39号中抗粤糖91-976×CP84-1198福建农林大学甘蔗综合研究所[71]
福农41号中抗ROC20×粤糖91-976福建农林大学甘蔗综合研究所[72]
福农81-745粤糖59-65×CP36-105福建农林大学甘蔗综合研究所[73]
福农83-36高抗CP49- 50×福农57-18福建农林大学甘蔗综合研究所[74]
福农91-3623免疫CP72-1210 ×桂73-167福建农林大学甘蔗综合研究所[75]
福农91-4621高抗CP72-1210 ×湛74-141福建农林大学甘蔗综合研究所[76]
福农94-0403中抗CP72-1210×闽糖69-263福建农林大学甘蔗综合研究所[77]
福农95-1702高抗CP72-1210×粤农73-204福建农林大学甘蔗综合研究所[78]
福农98-1103高抗CP72-1210×湛蔗74-141福建农林大学甘蔗综合研究所[79]
桂糖16号粤糖59/65×崖城72/399广西农业科学院甘蔗研究所[80]
桂糖23号高抗ROC 1号×崖城71-374广西农业科学院甘蔗研究所[81]
桂糖24号高抗桂糖71-5×崖城84-153广西农业科学院甘蔗研究所[82]
桂糖25号高抗CP72- 1210×崖城71- 374广西农业科学院甘蔗研究所[83]
桂糖42号高抗ROC 22×桂糖92-66广西农业科学院甘蔗研究所[84]
桂糖54号中抗桂糖00-122×崖城97-47广西农业科学院甘蔗研究所[85]
粤糖00-236高抗粤农73-204×CP72-1210广州甘蔗糖业研究所[86]
粤糖07913高抗HOCP 95-988×粤糖97-76广州甘蔗糖业研究所[87]
粤糖89-240CP72-1210×桂糖73-167广州甘蔗糖业研究所[88]
粤糖95-618免疫ROC10×CP72-1210广州甘蔗糖业研究所[89]
粤糖96-835高抗Co419×新台糖10号广州甘蔗糖业研究所[90]
闽糖88-103高抗闽选703×崖城82-96福建省农业科学院甘蔗研究所[91]
闽糖92-649高抗ROC1号×闽选703福建省农业科学院甘蔗研究所[92]
闽糖95-261高抗(ROC1×CP72-1210)×CP72-1210福建省农业科学院甘蔗研究所[93]
桂柳2号免疫ROC22×HoCP93-746柳城县甘蔗研究中心[94]
赣南02-70高抗桂糖69-435×CP84-1198江西省甘蔗研究所[95]
川蔗27号高抗CP72-1210×崖城90-3四川省植物工程研究院[96]
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甘蔗抗花叶病育种研究进展
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沈林波 1 , 潘继红 1 , 王会敏 1, 2 , 冯小艳 1 , 张树珍 1 , 王文治 1, 3, *
热带作物学报 | 种质资源与遗传育种 2024,45(11): 2320-2333
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热带作物学报 |种质资源与遗传育种 2024 , 45 (11) : 2320 -2333
甘蔗抗花叶病育种研究进展
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沈林波1, 潘继红1, 王会敏1, 2, 冯小艳1, 张树珍1, 王文治1, 3, *
作者信息
  • 1.中国热带农业科学院热带生物技术研究所/海南热带农业资源研究院,海南海口 571101
  • 2.南京农业大学生命科学学院,江苏南京 210095
  • 3.南京农业大学农学院,江苏南京 210095
通讯作者:
* 王文治(WANG Wenzhi),E-mail:
Advances in Sugarcane Breeding for Resistance to Sugarcane Mosaic Disease
Linbo SHEN1, Jihong PAN1, Huimin WANG1, 2, Xiaoyan FENG1, Shuzhen ZHANG1, Wenzhi WANG1, 3, *
Affiliations
  • 1.Institute of Tropical Bioscience and Biotechnology, Chinese Academy of Tropical Agricultural Sciences / Hainan Institute for Tropical Agricultural Resources, Haikou, Hainan 571101, China
  • 2.College of Life Sciences, Nanjing Agricultural University, Nanjing, Jiangsu 210095, China
  • 3.College of Agriculture, Nanjing Agricultural University, Nanjing, Jiangsu 210095, China
出版时间: 2024-11-25 doi: 10.3969/j.issn.1000-2561.2024.11.010
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甘蔗花叶病是危害甘蔗生产的主要病毒性病害之一,其病原主要包括甘蔗条纹花叶病毒(Sugarcane streak mosaic virus, SCSMV)、高粱花叶病毒(Sorghum mosaic virus, SrMV)和甘蔗花叶病毒(Sugarcane mosaic virus, SCMV)3大主要病原。甘蔗花叶病的发生严重影响甘蔗的品质和产量,防治该病害最经济有效的方法是抗病品种的应用。本文综述了甘蔗花叶病的危害和防治途径、甘蔗种质资源的抗性评价、抗性遗传分析、分子标记、抗性基因的挖掘利用及抗病育种等方面的研究进展,同时指出了甘蔗抗病育种方面存在的问题,并对加强甘蔗抗花叶病育种的研究提出了一些建议。

甘蔗  /  甘蔗花叶病  /  病害防治  /  抗花叶病育种

Sugarcane mosaic disease is one of the main viral diseases threatening the production of sugarcane. Sugarcane streak mosaic virus (SCSMV), Sorghum mosaic virus (SrMV) and Sugarcane mosaic virus (SCMV) are the main pathogens. The occurrence of sugarcane mosaic disease seriously affects the quality and yield of sugarcane. The most economical and effective way to control the disease is the application of resistant varieties. In this paper, the damage and controlling ways of sugarcane mosaic disease were summarized. The recent research progress on identification of resistance germplasm resources, resistance inheritance, molecular markers, mining and application of the resistance gene, and breeding of disease-resistant varieties were reviewed. Moreover, the problems existing in sugarcane disease resistance breeding were pointed out, and some suggestions were put forward to strengthen the research on sugarcane mosaic disease resistance breeding.

sugarcane  /  sugarcane mosaic disease  /  disease control  /  sugarcane mosaic disease resistance breeding
沈林波, 潘继红, 王会敏, 冯小艳, 张树珍, 王文治. 甘蔗抗花叶病育种研究进展. 热带作物学报, 2024 , 45 (11) : 2320 -2333 . DOI: 10.3969/j.issn.1000-2561.2024.11.010
Linbo SHEN, Jihong PAN, Huimin WANG, Xiaoyan FENG, Shuzhen ZHANG, Wenzhi WANG. Advances in Sugarcane Breeding for Resistance to Sugarcane Mosaic Disease[J]. Chinese Journal of Tropical Crops, 2024 , 45 (11) : 2320 -2333 . DOI: 10.3969/j.issn.1000-2561.2024.11.010
甘蔗(Saccharum officinarum L.),禾本科甘蔗属,糖含量丰富,以其为原料生产的蔗糖占世界食糖产量的70%,占中国食糖产量的90%[1],是我国最重要的糖料作物,同时在作为生物能源作物方面也具有广阔的应用前景[2]。1892年首次在爪哇发现甘蔗花叶病(sugarcane mosaic disease, SMD)[3],此后该病在全球各大蔗区广泛传播。甘蔗花叶病主要影响甘蔗的光合作用,症状表现为叶部产生与叶脉平行的褪绿条纹,长短不一,布满叶片,其中幼嫩叶片症状最为严重。甘蔗花叶病的发生造成甘蔗的矮化、叶片和茎的坏死以及严重的产量损失[4]。近年来,甘蔗花叶病在我国广西、云南和海南等主要蔗区普遍发生,发病率高达50%~100%,发病严重时甘蔗产量减少30%以上,严重制约了我国甘蔗产业的发展[5-7]。因此,本文对甘蔗花叶病的主要病原及其鉴定检测、传播方式、主要防治途径、甘蔗抗病种质资源鉴定评价,以及抗病品种选育等进行综述,并对未来研究作出展望,以期为选育抗甘蔗花叶病品种提供借鉴。
甘蔗花叶病主要由甘蔗线条花叶病毒(Sugarcane streak mosaic virus, SCSMV)、高粱花叶病毒(Sorghum mosaic virus, SrMV)和甘蔗花叶病毒(Sugarcane mosaic virus, SCMV)3种主要病原物引起[8-9],三者同属于马铃薯Y病毒科(Potyviridae),但SCMV和SrMV是马铃薯Y病毒属(Potyvirus)的成员,而后来发现的SCSMV属于禾本科病毒属(Poacevirus[10]。SCSMV、SrMV和SCMV均为由外壳蛋白(coat protein, CP)包裹着一条正义单链RNA的基因组组成,病毒粒体呈弯曲线状,大小约为10 kb,编码11个功能蛋白:P1、HC-Pro、P3、6K1、CI、6K2、NIa-Vpg、NIa-Pro、NIb、CP和P3N-PIPO[11]。SrMV和SCMV主要感染甘蔗、高粱和玉米[12-13],而SCSMV除了感染甘蔗和高粱外,玉米和苏丹草等禾本科植物也可作为SCSMV的人工寄主,其中对甘蔗的危害最为严重[14-16]
甘蔗花叶病主要由引种、机械摩擦和蚜虫感染等方式进行传播,其中SrMV和SCMV可被玉米蚜(Rhopalsiphum maidis)、桃蚜(Myzus persicae)和黑豆蚜(Aphis craccivora)进行传播,但目前未见SCSMV可被蚜虫传播的报道[17-18]。甘蔗花叶病的防治主要以预防为主。目前,常用的方法主要有2种:一是通过对农具消毒及驱除或杀灭昆虫介体等切断病毒的感染途径,该方法对防治病毒有一定效果,但是费时费力,而且使用化学药剂会造成环境污染[19]。二是通过组织培养脱毒法获得无毒的健康种苗。早在1994年许莉萍等[20]分别利用茎尖组织培养和愈伤组织培养法对甘蔗进行花叶病病原脱毒,结果在供体发病率为100%的情况下,利用茎尖组织培养的后代发病率减少至4.6%,而用愈伤组织培养的后代无一蔗株感病。杨本鹏等[21]通过热处理、温热处理和茎尖分生组织培养,有效去除了甘蔗花叶病病原体,得到了健康的脱毒苗。甘蔗脱毒种苗的应用虽然减轻了病毒在甘蔗中造成的危害,但由于病毒在田间条件下可通过蚜虫等传播,使病毒在再生作物中的危害增加。
随着甘蔗生物技术的发展,培育和利用抗病品种是目前最经济、安全且有效的防控病害途径。近年来,甘蔗育种学家在甘蔗花叶病抗性种质资源鉴定评价、抗性基因的挖掘利用以及抗病品种培育等方面开展了大量的研究,极大促进了甘蔗花叶病抗病育种的发展。
目前,对于甘蔗抗性种质资源的抗性评价,其过程一般为对甘蔗种质进行接种病毒,然后检测种质资源的发病情况,根据发病情况评价甘蔗种质资源的抗性。
甘蔗花叶病的病原人工接种方法主要包括摩擦接种法、浸根接种法和甘蔗生长期切茎接种法等。摩擦接种法通常是指在甘蔗叶面轻轻摩擦,使其表面产生微小创伤,从而使病毒侵染进入甘蔗细胞内。一般情况下,甘蔗花叶病的病毒粒子容易通过汁液接种进行侵染,但在甘蔗叶面摩擦时,由于摩擦力度不易掌握,往往导致叶片表面的伤口不均匀,使接种的病毒分布程度不同,发病情况不稳定,从而影响鉴定结果。摩擦接种法只适用于小批量材料接种,如需要对大量植株进行接种,则可选用浸根接种法,浸根接种法主要用于移栽时的幼苗大量接种。自2008年李文凤等[22]研究了甘蔗生长期切茎接种法这一新技术,鉴于该方法接种后发病均匀、稳定性好,可操作性强等优点,一直沿用至今,是目前甘蔗接种病毒最常用的一种方法。
快速、高效地检测病毒有助于及时阻断病毒的传播。早期甘蔗花叶病的病原检测主要通过生物学检测法、电镜检测法和血清学检测法等,但这些检测方法特异性差、灵敏度弱、操作不方便、容易造成误差[23]。近年来,由于分子生物学的发展,以PCR扩增为原理的检测技术,如逆转录PCR(reverse transcription PCR, RT-PCR)、荧光定量PCR(quantitative real-time PCR, qRT-PCR)、一步法多重RT-PCR(one step multiplex RT-PCR)和环介导等温扩增技术(loop-mediated isothermal amplification, LAMP)等已广泛应用于甘蔗病毒检测[24]。VISWANATHAN等[25]采集了63个甘蔗叶片样本,通过设计SCSMV-CP基因特异性引物进行RT-PCR检测,鉴定出58份甘蔗样本感染了SCSMV。FU等[26]通过建立SCSMV实时荧光定量PCR检测技术,实现了对SCSMV的定性和定量检测,且灵敏度比普通PCR高100倍。FENG等[27]提出了一步五重RT-PCR方法同时检测SCMV、SrMV、SCSMV、SCYLV、SCBV 5种病毒,结果表明,5对引物均能扩增出各自的目标病毒,具有较强的特异性,且检测灵敏度为单倍RT-PCR的10~100倍。CHEN等[28]以一个灵敏的RT-LAMP为基础,设计引物对SCSMV和SCMV外壳蛋白基因的6个不同区域进行扩增,均成功检测到病毒。
甘蔗抗病种质资源,即抗源,是指甘蔗属主要包括栽培种(栽培原种和杂交商业种)、野生种(细茎野生种和大茎野生种)和中间材料(杂交后代)及其近缘属植物在内的所有可利用的遗传资源。甘蔗抗病种质资源是选育优良抗病品种的遗传物质基础,因此,从甘蔗种质资源中找出抗病来源对于甘蔗抗花叶病育种具有重要意义。目前,甘蔗育种学家已在甘蔗野生种和栽培种中筛选出一大批抗病种质资源。
甘蔗不同种质间抗、感病程度各不相同。甘蔗野生种主要有细茎野生种(又称割手密,Saccharum spontaneum L.)和大茎野生种。与栽培种相比,甘蔗野生种遗传多样性丰富,具有很多适应自然逆境的优良性状,如分蘖多、生势强、宿根性好、耐旱、耐寒、抗病虫等,能够适应一系列复杂环境,是抵抗生物胁迫和非生物胁迫的重要基因来源[29]。黄忠兴等[30]通过对国内外不同地理来源的456份割手密资源进行农艺性状表型遗传多样性分析,结果表明割手密对甘蔗花叶病表现高抗。SILVA等[31]为寻找新的抗性种质,对野生甘蔗种质资源进行了抗性筛选,结果表明割手密和大茎野生种具有很好的抗性,其中IS76-155、IJ76-418 red、NG57-50、Ceram red、Badilla、Sac.off 8276、Fiji19、IJ76-313、US57-141-5、Krakatau、IN84-58、IN84-88、IN84-82、Gandacheni、Chin可能是抗花叶病的良好种质。LI等[32]对70份野生甘蔗种质资源(包括37份割手密和33份斑茅)进行抗SrMV鉴定评价,39份野生种(55.7%)表现为从一级高抗到三级中抗,具有优良的抗SrMV基因,其中云南82-1、云南83-157和四川79-I-15等7份割手密表现为一级高抗。海南甘蔗育种场以Badila、Lethers和Fiji等热带种与不同类型的割手密进行高贵化育种,获得含有割手密血缘后代的崖城系列材料,利用这批材料育成粤糖64-395等多个优良抗花叶病品种[33]
近年来,国内外甘蔗育种学家致力于甘蔗抗花叶病资源的筛选与创制,已筛选出一批优良的抗花叶病育种材料,并且大部分已应用于育种中。周仲驹等[34]利用摩擦法对9个供试品种进行人工接种SCMV-A株系,其中桂糖11和闽糖70/611表现为高抗。李文凤等[35]对32份优良甘蔗品种人工接种强致病性SrMV-HH1,筛选到云蔗89-351、云瑞99-155和赣蔗95-108等11份免疫品种和1份高抗品种Q170。付瑜华等[36]通过人工接种和RT-PCR技术对黔糖5号的抗病性进行了系统鉴定与评价,结果表明黔糖5号免疫SCSMV,高抗SrMV和SCMV。2017年美国佛罗里达州甘蔗品种委员会(Florida Sugarcane League)发布了产糖量较高的甘蔗杂交品种CP 06-2425、CP 06-2495、CP 06-2964、CP 06-3103和CP 07-1313,这些品种对SCMV-E株系具有一定抗性[37]。LI等[38]通过对71个优良甘蔗新品种同时接种SCSMV-JP1和SrMV-HH,筛选出粤甘34号、云蔗06-80、云蔗03-258等5个优良双抗品种。韦金菊等[39]对引进的41份甘蔗材料进行病害检测,其中CPCL97-2730、CPCL02-6848和CPCL02-1295等8个美国甘蔗种质与VN66-03、VN09-213和VN66-06等4个越南甘蔗种质对花叶病表现为一定的抗性。LI等[40]通过对41份甘蔗优良创新种质及其亲本人工接种SrMV和SCSMV对其进行双重抗性鉴定评价,综合分析得出,对SCSMV和SrMV均具有一级高抗性的种质资源有6份(云09-604、云09-607、云09-619、云09-633、云09-656、云滇95-19)。杨荣仲等[41]对102份国内外甘蔗亲本对花叶病的抗性进行鉴定评价,结果表明,ROC26、LC03-1137和GT02-761等8份亲本表现高抗,LC03-296、GT03-557和GT03-713等47份亲本表现抗病。通过我国甘蔗育种学家的不懈努力,大量甘蔗抗花叶病种质资源得到了精准挖掘与评价,这些抗性材料的获得为甘蔗花叶病抗病基因的挖掘和抗病品种的培育奠定了基础。
甘蔗是一种无性繁殖作物,具有高度杂合性的异源多倍体或非整倍体,遗传背景十分复杂。2005年薛志平[19]通过研究表明甘蔗抗花叶病遗传是受微效多基因控制的数量性状。近年来,有关甘蔗对花叶病的抗性遗传研究仍存在不足,甘蔗抗花叶病的遗传机制尚未明确,但可以借鉴玉米和高粱的甘蔗花叶病抗性遗传研究。CONDE等[42]研究表明,高粱对SCMV的抗性受单基因控制,抗性为显性,并且Krish基因对澳大利亚和一些国外甘蔗花叶病毒株系均具有一定抗性。KOVACS等[43]通过用SCMV-MB和SCMV-A机械接种11个玉米自交系及其杂交种,综合考虑了病株数和病征发展速度,结果表明多数自交系的病毒抗性遗传在很大程度上或一定程度上是显性的。WU等[44]研究发现2个显性互补基因共同控制玉米对SCMV的抗性,并利用微卫星标记将这2个基因分别定位在3号和6号染色体上。
近年来,陆续筛选和鉴定出甘蔗花叶病的QTL及分子标记,早期主要以AFLP分子标记和SSR分子标记为主。MEDEIROS等[45]通过cDNA-AFLP分析,研究了2个抗病性较强的甘蔗品种在受到SCMV病毒株系攻击后其转录谱的变化,结果表明,在测序的23个TDF中,有10个TDF(抗病品种独有)与植物对病原体的防御机制有关。目前,有关甘蔗抗花叶病的分子标记研究还存在不足,大多为玉米对花叶病抗性的分子标记,如DUBLE等[46]鉴定了位于6号染色体上的主基因Scmv1和位于3号染色体上的主基因Scmv2,通过靶向分离分析(targeted bulked segregant analysis, tBSA)在2个染色体区域进一步富集了SSR和AFLP标记,将14个SSR标记和6个AFLP标记定位到Scmv1区域,11个SSR标记和18个AFLP标记位于Scmv2区域。DUBLE等[47]以2个欧洲玉米自交系D32(抗性)×D145(易感)的219个F2后代为材料,定位并鉴定影响SCMV抗性的QTL位点,共鉴定出5个SCMV抗性QTL,分别位于1、3、5、6和10号染色体上。LÜ等[48]通过编译现有的81个QTL位点,构建了玉米抗甘蔗花叶病毒(SCMV)的QTL综合图谱,在3号染色体的“共识QTL”中鉴定出4个抗性候选基因。
植物病毒基因组过于简单,如果不与寄主因子相互作用,就无法在寄主植物上建立系统性感染[49]。近年来,随着分子生物学和基因组学的发展,甘蔗育种学家揭示了甘蔗花叶病病原与寄主互作的分子机制。董萌[50]通过研究表明,SCMV基因组连接蛋白VPg与真核细胞翻译起始因子4E(eIF4E)的互作在病毒成功侵染甘蔗的过程中起关键作用。朱海龙等[51]从甘蔗中筛选到与SCSMV-P3互作的基因ScRbcL,研究表明二者的互作可能是SCSMV侵染甘蔗导致花叶病症状发生的分子基础。ZHAI等[52]研究表明,ScELC基因表达的蛋白与SCSMV-VPg相互作用,并且其互作是甘蔗花叶病病原侵染甘蔗所必需的。袁军涛等[53]从甘蔗品种ROC22中克隆到ScPCaP1基因,研究表明该基因可能在SCSMV、SrMV和SCMV侵染寄主的过程中协助病毒粒子在植物胞间连丝中运动。ZHANG等[54]通过Y2H分析发现,与SCMV-6K2相互作用的RNS4、ULP5和VAMP727等7个甘蔗蛋白可能参与病毒的胁迫和防御反应。张海等[55]从热带种Badila中克隆了ScCRT1基因,研究表明该基因编码的蛋白与SCMV-6K2蛋白互作,并在SCMV侵染过程中起着积极作用。
由于甘蔗是异源多倍体作物,具有庞大的基因组,其后代性状遗传非常复杂,所以较少甘蔗抗性基因挖掘和利用的研究。凌辉[56]通过对甘蔗响应SrMV侵染的转录组分析表明,Pan1、ScSGS3、ScX1、WRKY、WRKY53MAP3K等基因是甘蔗中潜在的甘蔗抗SrMV侵染相关基因。PENG等[57]研究表明,甘蔗病程相关蛋白ScPR 10在SrMV的防御中起着积极作用,参与SrMV胁迫的防御反应,在抗病甘蔗品种中可被SrMV诱导表达。SILVA等[58]通过研究表明,抗SCMV甘蔗品种中天冬氨酸蛋白酶和VQ蛋白(VQ protein)基因获得显著上调,在甘蔗花叶病遗传抗性中发挥作用,并表明胞嘧啶甲基化的变化可能调控基因对甘蔗花叶病的抗性。杨宗桃等[59]从甘蔗品种ROC22中分离、鉴定了与SCMV-6K2互作的类泛素蛋白ScUBL5,研究表明ScUBL5可能介导甘蔗对SCMV的抗性,在甘蔗抗花叶病育种上具有潜在的应用价值。这些甘蔗寄主与病毒互作关系的分析以及抗性基因的挖掘为甘蔗抗花叶病育种提供了依据,今后可通过常规杂交、抗性基因转化、感病基因沉默等方法提高甘蔗品种对SCSMV、SrMV和SCMV的抗性。
甘蔗杂交育种是通过对不同亲本进行有性杂交获得杂交种子再进行培育、选择以育成新品种的方法。甘蔗育种学家很早就致力于通过该方法培育甘蔗新品种。20世纪前期甘蔗育种学家相继在爪哇和巴巴多斯发现的甘蔗热带种和甘蔗野生种割手密自然杂交的实生苗是甘蔗杂交育种的一次“飞跃式”进展[60]。澳大利亚育种学家BERDING等[61]通过以热带种为母本与缅甸割手密无性系Mandalay杂交,通过高贵化育种培育出对甘蔗花叶病具有较强抗性的Q系列品种。20世纪中后期,我国学者开始重视通过杂交育种获得具有甘蔗花叶病抗性的后代,在甘蔗育种学家的努力下,育成了对甘蔗花叶病不同抗性的甘蔗新品种(表1)。
由于甘蔗是高度杂合的异源多倍体或非整倍体无性繁殖作物,遗传背景复杂且传统育种具有长期性,使得通过常规手段去改变现有甘蔗品种的抗性比较困难。分子育种可直接从基因层面改变甘蔗品种的抗性,为甘蔗抗病育种开辟了新途径。目前,常见的分子育种方法主要包括蛋白质介导的抗性育种和利用RNA干扰培育抗病毒的转基因植物。
已有研究表明,利用病毒本身的蛋白基因转化植物体后,病毒表达的蛋白积累在寄主细胞中,阻碍了病毒的复制,使转化植物对该病毒产生抗性,病毒蛋白基因的表达量决定了植物体的抗性程度[97]。自1986年POWELL等首次将烟草花叶病毒(Tobacco mosaic virus, TMV)CP基因导入烟草获得抗TMV植株,该策略开始广泛应用于其他植物的抗病育种中[98]。目前,这种病毒抗性育种技术已相当成熟,在甘蔗花叶病的抗性育种中得到广泛应用。INGELBRECHT等[99]和BUTTERFIELD等[100]SrMV-CP基因转入甘蔗中,获得的转基因甘蔗在大田中对SrMV表现出抗性。APRIASTI等[101]SCMV-CP cDNA插入双载体pRI101-ON中,然后将其导入甘蔗,结果表明经病毒人工感染后,CP基因的全序列破坏了病毒的组装和包装,从而产生对SCMV的抗性。GILBERT等[102]将含有SCMV-CP基因的质粒导入甘蔗品种CP80-1827和CP84-1198中,通过对转基因甘蔗进行田间抗病性评价,发现转基因品系CP80-1827的花叶病发生率显著降低。福建农林大学甘蔗研究所利用受体品种Badila转入SCMV-CP基因培育抗花叶病甘蔗品种,获得转SCMV-CP基因品种福果系列,使甘蔗花叶病发病率大大降低[103]。杨川毓等[104]对转SrMV-P1基因甘蔗无性系TF50、TF37、TF63、TF53、TF67和TF64的抗病性进行了分析,结果表明转SrMV-P1基因甘蔗对花叶病的抗病性显著提高。BUTTERFIELD等[100]通过研究证明了甘蔗转基因植株的遗传稳定性,表明转基因甘蔗亲本可用于甘蔗育种。
RNA干扰(RNA interference, RNAi)是目前培育抗病毒转基因植物的常用方法。通过RNA介导的序列特异性相互作用抑制病毒基因表达被认为是控制植物病毒感染的有效途径。肖淑惠等[105]利用RNAi技术构建了能同时诱导SrMV、SCMV和SCYLV沉默的多价基因,并导入甘蔗胚性愈伤组织,获得了具有广谱抗性的转基因甘蔗。董萌[50]研究发现,利用RNAi技术沉默与Potyvirus互作的真核细胞翻译起始因子eIF4eIF(iso)4E基因可以阻断病毒的复制,转基因植物不仅可以正常生长而且对多种病毒产生抗性。徐景升等[106]从热带种Badila中克隆到甘蔗转录起始因子基因SceIF4E1,并以其作为RNAi的靶序列,经遗传转化后获得抗病性广、抗性持久的抗SCSMV转基因甘蔗。GUO等[107]SrMV-CP基因高度保守的序列为沉默靶点构建RNAi表达载体,将其导入到甘蔗品种福农15号的愈伤组织中,经鉴定表明转基因株系对SrMV具有一定抗性。陈利平等[108]将SrMV和ScYLV的双价RNAi结构构建干扰表达载体,并将其导入甘蔗品种福农15号,获得双抗SrMV和SCYLV的转基因甘蔗。徐景升等[109]克隆了与P3N-PIPO互作的甘蔗基因ScPCaP,并将其作为RNAi干扰靶序列,通过遗传转化获得了同时抗SCSMV、SrMV和SCMV的转基因植株。ASLAM等[110]选择SCMV-CP基因的保守区作为目的基因构建RNAi载体,将RNAi载体导入2个甘蔗品种SPF-234和NSG-311,获得抗SCMV的转基因甘蔗植株。WIDYANINGRUM等[111]SCMV-CP基因插入pGreen-0179质粒中,利用RNAi技术靶向沉默CP基因,获得转基因甘蔗,通过人工接种病毒后观察花叶症状和免疫印迹分析,发现转基因株系不受SCMV侵染。
甘蔗是我国重要的糖料作物之一,在糖料供给方面发挥重要作用。甘蔗花叶病的发生严重影响甘蔗的品质和产量,花叶病的防治已成为甘蔗生产过程中的主要任务之一。在分子育种时代,运用分子标记技术可以有效定位抗性基因、聚合多个抗性基因从而缩短育种年限,分子标记技术是甘蔗育种的一个重要方向。甘蔗育种学家应致力于全面阐明甘蔗抗花叶病的遗传机制,利用分子生物学技术培育高抗花叶病的甘蔗新品种。
目前,有关甘蔗花叶病的抗病机理和分子育种研究还存在不足。如,甘蔗花叶病的抗病机理仍不清楚;甘蔗抗花叶病的分子标记技术仍未在品种选育中发挥真正作用;传统的杂交育种仍是甘蔗抗花叶病育种的主要方式,大多数育成的抗病品种还依赖栽培种之间的杂交选育,很多野生种质资源如割手密中含有丰富的抗逆性基因,对多种病害高抗甚至免疫,但由于不同种质间的倍性差异和杂交亲和力低等因素,甘蔗野生种质资源的利用还非常有限。因此,目前甘蔗抗花叶病育种应着重以下几个方面:(1)结合甘蔗栽培种和野生种的全基因组测序结果,通过转录组学分析挖掘花叶病抗性关键基因,深入研究甘蔗抗花叶病的分子机理,阐明甘蔗抗花叶病的遗传机制;(2)充分挖掘甘蔗抗病种质资源的遗传多样性,利用分子标记技术聚合多个抗性基因进行综合性能良好和高抗花叶病甘蔗新品种的选育;(3)利用常规杂交育种、抗性基因转化、RNAi基因沉默等方法,将抗病基因导入甘蔗栽培种或沉默甘蔗种质中与病毒互作的基因,使甘蔗种质对病毒产生一定抗性。
  • 现代农业产业技术体系建设专项资金项目(CARS-17)
  • 海南省基础与应用基础研究计划项目(2019RC300)
  • 中央级公益性科研院所基本科研业务费专项(1630052019019)
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2024年第45卷第11期
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doi: 10.3969/j.issn.1000-2561.2024.11.010
  • 接收时间:2024-05-14
  • 首发时间:2026-06-26
  • 出版时间:2024-11-25
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  • 收稿日期:2024-05-14
  • 修回日期:2024-05-30
基金
现代农业产业技术体系建设专项资金项目(CARS-17)
海南省基础与应用基础研究计划项目(2019RC300)
中央级公益性科研院所基本科研业务费专项(1630052019019)
作者信息
    1.中国热带农业科学院热带生物技术研究所/海南热带农业资源研究院,海南海口 571101
    2.南京农业大学生命科学学院,江苏南京 210095
    3.南京农业大学农学院,江苏南京 210095

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* 王文治(WANG Wenzhi),E-mail:
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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

Genus
种数
Number of
species
占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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