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Article(id=1276262847680221323, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276262756814815737, articleNumber=null, orderNo=null, doi=10.3969/j.issn.1000-2561.2024.08.016, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1683216000000, receivedDateStr=2023-05-05, revisedDate=1685462400000, revisedDateStr=2023-05-31, acceptedDate=null, acceptedDateStr=null, onlineDate=1782214374704, onlineDateStr=2026-06-23, pubDate=1724515200000, pubDateStr=2024-08-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1782214374704, onlineIssueDateStr=2026-06-23, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1782214374704, creator=13701087609, updateTime=1782214374704, updator=13701087609, issue=Issue{id=1276262756814815737, tenantId=1146029695717560320, journalId=1235980609244409860, year='2024', volume='45', issue='8', pageStart='1521', pageEnd='1760', issueExtLink='null', onlineDate='null', pubDate='1724515200000', pubDateStr='2024-08-25', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1782214353040, creator='13701087609', updateTime=1782214460949, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1276263209816420382, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276262756814815737, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1276263209816420383, tenantId=1146029695717560320, journalId=1235980609244409860, issueId=1276262756814815737, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=1670, endPage=1676, ext={EN=ArticleExt(id=1276262847936073869, articleId=1276262847680221323, tenantId=1146029695717560320, journalId=1235980609244409860, language=EN, title=Evaluation of the Resistance of Shanlan Upland Rice to Brown Planthopper, columnId=1236292524264968282, journalTitle=Chinese Journal of Tropical Crops, columnName=Plant Protection & Bio-safety, runingTitle=null, highlight=null, articleAbstract=

Shanlan upland rice (Oryza sativa L.) is an unique variety in Hainan, with excellent taste quality traits and rich nutrition. At the same time, it also has excellent stress resistance, such as disease resistance, drought resistance and heat resistance. However, thus far, the resistance of Shanlan upland rice to herbivores remains largely unknown. Thus, in this study, using TN1 and RHT as susceptible and resistant control varieties, respectively, the resistance of three Shanlan upland rice varieties, Shanlanchaihongmi, Shanlanjianshanlannuo and Shanlanyingmigaochan to BPH was evaluated, and their resistance mechanisms were analyzed. The results showed that at the seedling stage, the variety of Shanlanjianshannuo showed resistant to Nuaparvata lugens, brown planthopper BPH, whereas the other two varieties were susceptible to BPH. At the tillering stage, the resistance of the three rice cultivars to BPH was all significantly higher than that of TN1. The mechanism studies revealed that the thickness of the thick-walled tissue in the outer leaf sheaths and the level of cell wall components including lignin, pectin, cellulose and hemicellulose were significantly higher in the three Shanlan upland rice cultivars and the insect-resistant control cultivar RHT than the insect-susceptible control cultivar TN1. Moreover, the basal and BPH-induced levels of JA, JA-Ile, ABA and SA in the three Shanlan upland rice cultivars were significantly higher than those in TN1, and the content of ABA in Shanlanjianshanlannuo plants was the highest in all of the tested varieties. The results demonstrated that the thickness of the thick-walled tissue of the outer leaf sheaths of Shanlan upland rice and the rapid and strong responses of JA, ABA and SA signaling pathways to BPH infestation might be the major reasons why the BPH resistance is stronger in Shanlan upland rice cultivars than TN1.

, authors=null, authorsList=Zijie LIU, Yonggen LOU, Baoqian LYU, authorCompany=null, correspAuthors=Baoqian LYU, 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=1276262850582679703, articleId=1276262847680221323, tenantId=1146029695717560320, journalId=1235980609244409860, language=CN, title=山栏稻对褐飞虱抗性评价, columnId=1236292524520820846, journalTitle=热带作物学报, columnName=植物保护与生物安全, runingTitle=null, highlight=null, articleAbstract=

山栏稻（Oryza sativa L.）是海南特有品种，有着优异的食味品质性状，营养丰富，同时还具有抗病、抗旱、耐热等优良的抗逆特性。但目前对山栏稻抗虫方面研究较少。为此，本研究分别以TN1和RHT为感虫和抗虫对照品种，对山栏柴红米、山栏尖山栏糯和山栏硬米高产3个山栏稻品种对褐飞虱的抗性进行评价，并分析其抗性机理。结果表明：在3个山栏稻品种中，山栏尖山栏糯在苗期的抗性表现为抗，另外2个品种均表现为感；在分蘖期，3个山栏稻品种对褐飞虱的抗性均显著高于TN1。抗性机理研究表明：3个山栏稻品种和抗虫对照品种RHT外层叶鞘部位厚壁组织的厚度以及细胞壁组成成分木质素、果胶、纤维素和半纤维素含量均显著高于感虫对照品种TN1。同时，3个山栏稻品种中本底和褐飞虱为害诱导的JA、JA-Ile、ABA和SA含量在各个时间点均显著高于TN1，并且山栏尖山栏糯体内ABA的含量在各个时间均为最高。这些结果表明，山栏稻外层叶鞘部位厚的厚壁组织及其JA、ABA和SA信号途径快速且强烈的响应，可能是山栏稻品种对褐飞虱抗性强于TN1的一个主要原因。

, authors=

* 娄永根（1964—），男，博士，教授，研究方向：昆虫与植物分子互作。

刘子杰（1997—），女，硕士研究生，研究方向：水稻与褐飞虱互作

, authorsList=刘子杰, 娄永根, 吕宝乾, authorCompany=null, correspAuthors=吕宝乾, authorNote=null, correspAuthorsNote=
** 吕宝乾（LYU Baoqian），E-mail：。
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刘子杰（1997—），女，硕士研究生，研究方向：水稻与褐飞虱互作

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刘子杰（1997—），女，硕士研究生，研究方向：水稻与褐飞虱互作

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Different lowercase letters indicate significant difference (P<0.05).

, figureFileSmall=76x+WBHb2tt/uHIzTOoAyw==, figureFileBig=OgsUwTT+XQsJ5kCs3HFIGA==, tableContent=null), ArticleFig(id=1276269311413064150, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=CN, label=图1, caption=分蘖期水稻品种对褐飞虱若虫（A）、卵存活率（B）、雌成虫产卵量（C）的影响

不同小写字母表示差异显著（P<0.05）。

, figureFileSmall=76x+WBHb2tt/uHIzTOoAyw==, figureFileBig=OgsUwTT+XQsJ5kCs3HFIGA==, tableContent=null), ArticleFig(id=1276269311723442647, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=EN, label=Fig. 2, caption=Leaf sheath thick-walled tissue and its thickness of different rice cultivars before BPH infestation

A: Thick-walled tissue of 40-day-old rice leaf sheath paraffin section, scale bar = 20 µm; B: Thickness of thick-walled tissue; Different lowercase letters indicate significant difference (P<0.05).

, figureFileSmall=4LUzZhaCDkHVzoevaIkrSw==, figureFileBig=5SUWO8RRBHuVUiQSCF1CBw==, tableContent=null), ArticleFig(id=1276269311790551512, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=CN, label=图2, caption=BPH为害前各水稻品种叶鞘厚壁组织及其厚度

A：40日龄水稻叶鞘石蜡切片厚壁组织，比例尺=20 µm；B：厚壁组织厚度；不同小写字母表示显著差异（P<0.05）。

, figureFileSmall=4LUzZhaCDkHVzoevaIkrSw==, figureFileBig=5SUWO8RRBHuVUiQSCF1CBw==, tableContent=null), ArticleFig(id=1276269311857660377, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=EN, label=Fig. 3, caption=Content of lignin (A), pectin (B) and cellulose (C) in leaf sheaths of rice cultivars before BPH infestation

Different lowercase letters indicate significant difference (P<0.05).

, figureFileSmall=UDiwwgXi1Yrl7vSjgaiEqw==, figureFileBig=otLiI59uA3Q7s6ylRuo+kA==, tableContent=null), ArticleFig(id=1276269311933157850, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=CN, label=图3, caption=BPH为害前各水稻品种叶鞘木质素（A）、果胶（B）、纤维素（C）含量

不同小写字母表示显著差异（P<0.05）。

, figureFileSmall=UDiwwgXi1Yrl7vSjgaiEqw==, figureFileBig=otLiI59uA3Q7s6ylRuo+kA==, tableContent=null), ArticleFig(id=1276269313581519323, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=EN, label=Fig. 4, caption=The content of JA, JA-Ile, SA and ABA in leaf sheaths of five rice cultivars at different BPH infestation time

Different lowercase letters indicate significant difference (P<0.05).

, figureFileSmall=zDIxuhn4Cs/YPnoC3vWG8A==, figureFileBig=weuig3ci/glRJ9rGpoiEmw==, tableContent=null), ArticleFig(id=1276269313673794012, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=CN, label=图4, caption=BPH为害不同时间后各水稻品种内JA、JA-Ile、SA、ABA的含量

不同小写字母表示显著差异（P<0.05）。

, figureFileSmall=zDIxuhn4Cs/YPnoC3vWG8A==, figureFileBig=weuig3ci/glRJ9rGpoiEmw==, tableContent=null), ArticleFig(id=1276269313766068701, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=EN, label=Tab. 1, caption=

Resistance level of different rice varieties to N. lugens at seedling stage

, figureFileSmall=null, figureFileBig=null, tableContent=
品种Variety死苗率Mortality/%抗级Resistance grade
TN1100.0±0.0a9 (HS)
RHT3.3±3.3c1 (HR)
山栏柴红米60.0±5.8b7 (S)
山栏尖山栏糯10.0±3.3c3 (R)
山栏硬米高产63.3±8.8b7 (S)
), ArticleFig(id=1276269313828983262, tenantId=1146029695717560320, journalId=1235980609244409860, articleId=1276262847680221323, language=CN, label=表1, caption=

苗期的水稻品种对褐飞虱的抗性水平

, figureFileSmall=null, figureFileBig=null, tableContent=
品种Variety死苗率Mortality/%抗级Resistance grade
TN1100.0±0.0a9 (HS)
RHT3.3±3.3c1 (HR)
山栏柴红米60.0±5.8b7 (S)
山栏尖山栏糯10.0±3.3c3 (R)
山栏硬米高产63.3±8.8b7 (S)
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刘子杰 1, 2 , 娄永根 1, 2 , 吕宝乾 3, 4, **
热带作物学报 | 植物保护与生物安全 2024,45(8): 1670-1676
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热带作物学报 |植物保护与生物安全 2024 , 45 (8) : 1670 -1676
山栏稻对褐飞虱抗性评价
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刘子杰（1997—），女，硕士研究生，研究方向：水稻与褐飞虱互作

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刘子杰（1997—），女，硕士研究生，研究方向：水稻与褐飞虱互作

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刘子杰1, 2, 娄永根1, 2, 吕宝乾3, 4, **
作者信息
  • 1.浙江大学昆虫科学研究所，浙江杭州　310058
  • 2.浙江大学海南研究院，海南三亚　572025
  • 3.中国热带农业科学院环境与植物保护研究所，海南海口　571101
  • 4.中国热带农业科学院三亚研究院/海南省南繁生物安全与分子育种重点实验室，海南三亚　572025
通讯作者:
** 吕宝乾（LYU Baoqian），E-mail：。
Evaluation of the Resistance of Shanlan Upland Rice to Brown Planthopper
Zijie LIU1, 2, Yonggen LOU1, 2, Baoqian LYU3, 4, **
Affiliations
  • 1.Institute of Insect Sciences, Zhejiang University, Hangzhou, Zhejiang 310058, China
  • 2.Hainan Institute, Zhejiang University, Sanya, Hainan 572025, China
  • 3.Environment and Plant Protection Institute, Chinese Academy of Tropical Agricultural Sciences, Haikou, Hainan 571101, China
  • 4.Sanya Research Institute, Chinese Academy of Tropical Agricultural Sciences / Hainan Key Laboratory for Biosafety Monitoring and Molecular Breeding in the Off-season Reproduction Regions, Sanya, Hainan 572025, China
出版时间: 2024-08-25 doi: 10.3969/j.issn.1000-2561.2024.08.016
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摘要
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山栏稻（Oryza sativa L.）是海南特有品种，有着优异的食味品质性状，营养丰富，同时还具有抗病、抗旱、耐热等优良的抗逆特性。但目前对山栏稻抗虫方面研究较少。为此，本研究分别以TN1和RHT为感虫和抗虫对照品种，对山栏柴红米、山栏尖山栏糯和山栏硬米高产3个山栏稻品种对褐飞虱的抗性进行评价，并分析其抗性机理。结果表明：在3个山栏稻品种中，山栏尖山栏糯在苗期的抗性表现为抗，另外2个品种均表现为感；在分蘖期，3个山栏稻品种对褐飞虱的抗性均显著高于TN1。抗性机理研究表明：3个山栏稻品种和抗虫对照品种RHT外层叶鞘部位厚壁组织的厚度以及细胞壁组成成分木质素、果胶、纤维素和半纤维素含量均显著高于感虫对照品种TN1。同时，3个山栏稻品种中本底和褐飞虱为害诱导的JA、JA-Ile、ABA和SA含量在各个时间点均显著高于TN1，并且山栏尖山栏糯体内ABA的含量在各个时间均为最高。这些结果表明，山栏稻外层叶鞘部位厚的厚壁组织及其JA、ABA和SA信号途径快速且强烈的响应，可能是山栏稻品种对褐飞虱抗性强于TN1的一个主要原因。

山栏稻  /  褐飞虱  /  抗虫相关植物激素  /  食草性昆虫抗性

Shanlan upland rice (Oryza sativa L.) is an unique variety in Hainan, with excellent taste quality traits and rich nutrition. At the same time, it also has excellent stress resistance, such as disease resistance, drought resistance and heat resistance. However, thus far, the resistance of Shanlan upland rice to herbivores remains largely unknown. Thus, in this study, using TN1 and RHT as susceptible and resistant control varieties, respectively, the resistance of three Shanlan upland rice varieties, Shanlanchaihongmi, Shanlanjianshanlannuo and Shanlanyingmigaochan to BPH was evaluated, and their resistance mechanisms were analyzed. The results showed that at the seedling stage, the variety of Shanlanjianshannuo showed resistant to Nuaparvata lugens, brown planthopper BPH, whereas the other two varieties were susceptible to BPH. At the tillering stage, the resistance of the three rice cultivars to BPH was all significantly higher than that of TN1. The mechanism studies revealed that the thickness of the thick-walled tissue in the outer leaf sheaths and the level of cell wall components including lignin, pectin, cellulose and hemicellulose were significantly higher in the three Shanlan upland rice cultivars and the insect-resistant control cultivar RHT than the insect-susceptible control cultivar TN1. Moreover, the basal and BPH-induced levels of JA, JA-Ile, ABA and SA in the three Shanlan upland rice cultivars were significantly higher than those in TN1, and the content of ABA in Shanlanjianshanlannuo plants was the highest in all of the tested varieties. The results demonstrated that the thickness of the thick-walled tissue of the outer leaf sheaths of Shanlan upland rice and the rapid and strong responses of JA, ABA and SA signaling pathways to BPH infestation might be the major reasons why the BPH resistance is stronger in Shanlan upland rice cultivars than TN1.

Shanlan upland rice  /  brown planthopper  /  defense-related phytohormone  /  herbivore resistance
刘子杰, 娄永根, 吕宝乾. 山栏稻对褐飞虱抗性评价. 热带作物学报, 2024 , 45 (8) : 1670 -1676 . DOI: 10.3969/j.issn.1000-2561.2024.08.016
Zijie LIU, Yonggen LOU, Baoqian LYU. Evaluation of the Resistance of Shanlan Upland Rice to Brown Planthopper[J]. Chinese Journal of Tropical Crops, 2024 , 45 (8) : 1670 -1676 . DOI: 10.3969/j.issn.1000-2561.2024.08.016
正文
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水稻作为我国十分重要的粮食作物，对其安全生产的保障举足轻重。然而褐飞虱[Nuaparvata lugens（Stål）]是一种具有迁飞和爆发性的昆虫[1]，通过取食、产卵和传播病毒给水稻生产带来巨大的经济损失[2]。现如今防治褐飞虱最环保、有效、经济的方法就是对抗虫品种的合理利用[3-4]。因此，研究水稻抗虫机理、挖掘新的水稻抗虫基因、培育新的水稻抗虫品种对我国水稻生产具有重要意义[5]。
已有研究表明，水稻内源的茉莉酸（jasmonic acid，JA）、水杨酸（salicylic acid，SA）、脱落酸（abscisic acid，ABA）等植物激素在调控水稻对褐飞虱的抗性中发挥着重要作用[6]。在水稻中，JA信号通路损伤突变体（OsAOC或OsMYC2敲除突变体）体内次生代谢物，如酚胺类、黄酮类和挥发物等物质显著降低，最终导致褐飞虱的卵孵化率显著上升，水稻抗性减弱[7]。研究发现，稻纵卷叶螟（Cnaphalocrocis medinalis）取食水稻叶片能够诱导SA产生，且外施SA提高了水稻TrypPIs的含量；抗虫水稻品种bph6受到褐飞虱为害后能积累更高的SA水平，表明SA参与调控水稻对不同类型害虫为害的应答[8-9]。研究表明，外施ABA的水稻在褐飞虱为害后，体内β-1,3-葡聚糖酶活性降低，胼胝质合成酶活性升高，诱导产生的胼胝质影响褐飞虱卵黄原蛋白的转录和褐飞虱的产卵行为，最终提高水稻对褐飞虱的抗性；最近研究发现，ABA水解酶OsABA8ox3也参与该过程的调控[10-12]。此外，水稻对褐飞虱的抗性也与水稻厚壁组织的厚度有关[13]。厚壁组织位于表皮下，是维管束附近的结构支持组织，距离水稻叶鞘维管束只有1到2层细胞。厚壁组织包含多层细胞，坚硬的次生壁形成1个厚厚的区域，以保护和机械支撑内部组织。这表明它可能在褐飞虱口针到达韧皮部取食的过程中发挥作用。
山栏稻是适合海南中西部干旱地区种植的特有稻种资源。作为珍贵种质资源的山栏稻，具有许多优良的抗逆特性，比如抗病、抗旱、耐热等。但是抗虫方面的信息报道较少[14-15]。山栏稻抗虫性评价是水稻抗虫的种质资源获得的重要环节，可进一步应用于水稻育种，这对改良、利用和保护山栏稻种质资源具有重要意义。
1　材料与方法
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1.1　材料
1.1.1　供试虫源
用于试验的褐飞虱[Nuaparvata lugens（Stål），rice brown planthopper，BPH]种群来自海南三亚稻田，在人工气候室条件下[温度（26±2）℃，光照12 h，湿度70%~80%]，用催芽7 d左右的TN1水稻幼苗饲喂并进行种群繁殖。本试验的褐飞虱均为室内培养30代以上，且试验前通过苗期抗性鉴定试验对褐飞虱致害力进行评价。进行生测测定试验前，选取数量比为2∶1的初羽化BPH雌成虫和雄成虫至新笼中，用TN1水稻苗（催苗后约7 d左右）继续饲养，让其自由交配和怀卵，3~4 d后挑选大小一致BPH怀卵雌成虫用于相应的生物测定试验。
1.1.2　供试水稻
在本研究中，所用水稻品种分别为感虫对照品种Taichung native 1（TN1）、抗虫对照品种Rathu Heenati（RHT）和3个山栏稻品种（山栏柴红米、山栏尖山栏糯和山栏硬米高产）。前2个品种由中国水稻研究所提供种子，后面3个山栏稻品种为海南当地品种，由海南琼中村民提供。将水稻种子放于培养皿中，用清水浸泡24 h使种子发芽，其后注意清洗，并保持种子表面呈湿润且非浸泡状态，待苗长高至4~6 cm后转移至30 L塑料筐中并置于温室[温度（28±2）℃，光照14 L∶10 D，湿度50%~60%]，期间每10 d左右更换培养液1次。待生长30 d后，选取生长状态较好、长势一致的水稻，摘除黄叶，根据不同试验需要提前3~4 d以单株形式转移至不透光的塑料杯（直径8 cm，高10 cm）中，用相同的营养液培养，恢复生长3~4 d后用于后续试验。
1.2　方法
1.2.1　苗期抗性试验
采用标准苗期集团筛选法（SSST法）[16]。将每个品种的水稻种子（包括感虫对照TN1和抗虫对照RHT）播种于育秧盘中，每品种10穴，设3次重复。各品种随机排列。试验于室外进行，采用自然光照。种子发芽1周后去掉未发芽的种子或者生长不佳的稻苗，每穴只留1株健康稻苗。到2叶1心时期，平均每株水稻苗接入褐飞虱1~2龄若虫5~6头。当TN1受害率达70%时，每天记录各品种稻苗受害情况；当TN1全部枯死时，评定各品种抗性等级：死苗率低于1.0%为免疫，死苗率1.1%~10.0%为1级（高抗）、10.1%~30.0%为3级（抗）、30.1%~50.0%为5级（中抗）、50.1%~70.0%为7级（感）、高于70.1%为9级（高感）[17]。
1.2.2　分蘖期抗性试验
接虫时，将不同水稻品种分组。每株稻苗茎秆下部罩上特制的玻璃筒（直径4 cm，高8 cm，筒壁均匀分布48个直径0.8 mm的小孔），然后接入初孵若虫20头，玻璃管的上端用海绵封口。每个处理设置10个重复。每天记录若虫存活情况，直到第12天。根据结果计算若虫存活率。
按上述分组每株水稻苗接入10头褐飞虱待产卵雌成虫，每个水稻品种重复10次。24 h后去除褐飞虱成虫。每天观察记录各水稻苗上孵化的若虫数，至无若虫孵化时，剪取水稻苗于显微镜下计数未孵化卵的数量，计算卵孵化率。
按上述分组每株稻苗茎秆下部罩上特制的玻璃筒（直径4 cm，高8 cm，筒壁均匀分布48个直径0.8 mm的小孔），筒内放置1头初羽化雌成虫和1头初羽化雄成虫。8 d后，将虫移除，每个处理设置10个重复，使用电子显微镜检查每株水稻上的产卵量。
1.2.3　厚壁组织厚度测量
切取水稻苗叶鞘，长约2 cm，固定在FAA溶液中，然后进行脱水、包埋、切片、番红固绿、全片扫描。用NDP.view 2软件对水稻苗最外两层叶鞘厚壁组织厚度进行分析。
1.2.4　细胞壁组成成分测定
取水稻苗最外两层水稻叶鞘（长约2 cm）进行细胞壁组分的测量。
果胶含量测定：采用比色法测定。参照苏州科铭生物技术有限公司总果胶试剂盒提供的实验方法。
木质素含量测定：取样品300 mg，加入甲醇反复震荡洗涤，剩余组织残渣烘干后用于木质素的提取。具体提取方法参考XU等[18]的方法，使用分光光度计检测，设置3个生物学重复。
纤维素含量测定：首先制备细胞壁物质，获得的细胞壁物质进行纤维素的提取和测定。参照苏州科铭生物技术有限公司纤维素试剂盒提供的实验方法。
1.2.5　激素含量测定
水稻生长25~30 d后，单株移至塑料杯中，每株水稻接15头BPH怀卵雌成虫，BPH怀卵雌成虫处理0、8、24 h后，用剪刀剪取BPH产卵、为害部位的水稻茎秆，迅速浸入液氮中，–80 ℃保存。称取液氮研磨后的样品0.15 g左右，每个处理设置3个生物学重复。JA、JA-Ile、SA、ABA激素含量的提取及测定方法参照HETTENHAUSEN等[19]的方法。
1.3　数据处理
采用Excel软件进行数据整理，根据试验设计和统计分析原理，利用SPSS 16.0（Chicago，Illinois State，USA）软件完成数据统计分析。本试验数据均先采用单因素方差分析（one-way ANOVA），再采用Tukey’s HSD post- hoc tests进行检验。
2　结果与分析
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2.1　不同水稻品种苗期抗褐飞虱鉴定结果
参试的5个水稻品种，经苗期抗性鉴定，RHT和山栏尖山栏糯在苗期抗性较好，分别为高抗和抗；山栏柴红米和山栏硬米高产则表现为感（表1）。
2.2　不同水稻品种成株期对褐飞虱生长发育繁殖的影响
褐飞虱初孵若虫在5个品种上的存活率存在显著差异，其中在TN1上的存活率最高，为73.5%；在抗虫对照品种RHT上的存活率最低，为35.0%；而在山栏柴红米、山栏尖山栏糯、山栏硬米高产上的存活率介于上述二者之间，分别为44%、59%、46.5%（图1A）。褐飞虱雌成虫在TN1上的卵孵化率最高，为73.17%，在山栏柴红米、山栏尖山栏糯、山栏硬米高产上的卵孵化率分别为66.69%、65.52%、67.22%，在RHT上的卵孵化率最低，为61.09%；在TN1上的卵孵化率显著高于3种山栏稻和RHT（图1B）。褐飞虱雌成虫在TN1上的产卵量（244.3粒）显著高于其在3种山栏稻品种上产卵量；在山栏硬米高产上的产卵量为197.5粒，在山栏柴红米的产卵量为172.6粒，在山栏尖山栏糯的产卵量为176.7粒，在RHT的产卵量为131.3粒（图1C）。
2.3　不同水稻品种厚壁组织厚度
对5个水稻品种叶鞘进行石蜡切片，结果显示RHT以及3个山栏稻品种最外两层叶鞘厚壁组织的厚度均显著厚于TN1（图2）。
2.4　不同水稻品种细胞壁组成成分含量
对水稻叶鞘外层细胞壁组成成分木质素、果胶、纤维素进行测定，结果表明RHT、山栏尖山栏糯及山栏硬米高产体内木质素含量均显著高于TN1（图3A）；山栏尖山栏糯体内果胶含量显著高于其他4种水稻品种（图3B）；RHT与3种山栏稻体内纤维素含量均显著高于TN1（图3C）。
2.5　褐飞虱为害对水稻体内激素含量的影响
研究结果表明，3个山栏稻品种中本底的JA和JA-Ile含量均显著高于TN1和RHT，其中尤以山栏硬米高产品种中JA含量最高；褐飞虱为害8~24 h时5个水稻品种体内JA含量整体呈先上升后下降的趋势，并且总体上3个山栏稻品种和RHT中的JA和JA-Ile含量均显著高于TN1（图4A、图4B）。RHT品种和3个山栏稻品种中的SA和ABA含量，无论是本底的还是褐飞虱为害8 h或24 h诱导的，均显著高于品种TN1（图4C、图4D）。
3　讨论
收起
本研究通过开展3种山栏稻对褐飞虱抗性评价，发现相比于TN1，山栏稻对BPH的生长与繁殖有一定抑制作用，说明山栏稻对褐飞虱具有一定的抗性。其中，山栏尖山栏糯在3种山栏稻中抗性最好，抗性级别最高。
BPH是一种具有毁灭性的水稻害虫，BPH的口针穿过厚壁组织，在韧皮部建立为害位点。前人研究结果表明厚壁组织增厚是阻止褐飞虱到达韧皮部进行为害的物理障碍，从而增加水稻对BPH的抗性[13]。本研究结果显示，RHT及3种山栏稻厚壁组织厚度在褐飞虱为害前著高于TN1，并且细胞壁组成成分的含量均显著高于TN1，厚壁组织的增厚与细胞壁组成成分密切相关，而厚壁组织的增厚又进一步增加水稻对褐飞虱的抗性。这些研究结果表明，加强通往韧皮部路径上的细胞壁可能是抵抗植物韧皮部昆虫为害的常见机制。
植物激素在植物抵御昆虫为害中发挥重要作用。JA信号通路在调控水稻抗BPH中发挥着重要作用。为了应对BPH的攻击，JA在水稻叶鞘中迅速积累，随后激活下游JA介导的防御反应，如防御性次生代谢产物的生物合成。而JA-Ile是JA的活性物质，前人研究表明，植物在受到外界胁迫后会促进体内JA-Ile的合成，进而诱导植物形成防御系统，诱导防御相关基因表达以保护植株免受外界胁迫损伤[20]。目前研究发现，缺乏ABA会降低植物对植食性动物的抗性，如有研究表明甜菜夜蛾更倾向于取食ABA含量低的番茄，抗虫能力越差ABA含量越低[21-23]。除此之外，研究表明，ABA还能提高水稻对褐飞虱的抗性，增强水稻对褐飞虱抗性的重要原因之一是诱导了胼胝质的合成，形成一道可有效抵御褐飞虱侵害的屏障[24]，从而提高水稻的抗虫性。SA普遍存在于植物体内，并且在抗盐胁迫、抗冷、抗病和抗虫等方面扮演着重要的角色。本研究结果表明，3个山栏稻品种和RHT中褐飞虱为害诱导的JA、JA-Ile、ABA和SA含量均显著高于TN1品种，表明这些植物激素可能均参与山栏稻对褐飞虱的抗性的调控。综上所述，山栏稻对褐飞虱存在一定抗性，其中山栏尖山栏糯的抗性表现最好。山栏稻外层叶鞘部位的厚壁组织及其JA、ABA和SA信号途径快速且强烈的响应，可能是山栏稻品种对褐飞虱抗性强于TN1的一个主要原因。
基金
收起
  • 海南省国际科技合作研发项目(GHYF2022002)
  • 海南省重大科技计划项目(ZDKJ202002; ZDKJ201901)
文献
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2024年第45卷第8期
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文章信息
doi: 10.3969/j.issn.1000-2561.2024.08.016
  • 接收时间：2023-05-05
  • 首发时间：2026-06-23
  • 出版时间：2024-08-25
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  • 收稿日期：2023-05-05
  • 修回日期：2023-05-31
基金
海南省国际科技合作研发项目(GHYF2022002)
海南省重大科技计划项目(ZDKJ202002; ZDKJ201901)
作者信息
    1.浙江大学昆虫科学研究所，浙江杭州　310058
    2.浙江大学海南研究院，海南三亚　572025
    3.中国热带农业科学院环境与植物保护研究所，海南海口　571101
    4.中国热带农业科学院三亚研究院/海南省南繁生物安全与分子育种重点实验室，海南三亚　572025

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** 吕宝乾（LYU Baoqian），E-mail：。
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
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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