麦红吸浆虫滞育的生理生化机制研究
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摘要
麦红吸浆虫Sitodiplosis mosellana(Gehin)是小麦生产中间歇性猖獗成灾的毁灭性害虫。在我国20世纪50和80年代曾两次大面积暴发成灾，损失惨重。近年来，麦红吸浆虫在我国大范围猖獗成灾的势头得到了有效的遏制，但局部地区大发生的现象仍时有发生，并具有扩散蔓延的趋势，对小麦生产的威胁并未解除。多年来的研究表明，滞育是引起其猖獗成灾的主要原因之一，但有关麦红吸浆虫的滞育机理，有许多问题至今仍不清楚。本文在前人研究工作的基础上，采用生物化学、蛋白质组学、放射免疫分析和电镜技术等多学科交叉的前沿方法和手段，继续对麦红吸浆虫滞育机理进行深入细致的研究，取得了一定的新进展，为今后进一步深入研究其滞育机理、成灾规律、预测预报、大面积综合治理奠定了良好的基础。主要研究结果如下：
1．应用考马斯亮蓝G-250染色法、SDS-PAGE及PAGE电泳技术分析了麦红吸浆虫滞育前后及滞育期蛋白质含量和组分的变化。结果表明，不同阶段蛋白质含量存在显著差异，老熟幼虫入土前和滞育解除后均高于入土滞育期。入土滞育期间，同期裸露幼虫与结茧幼虫，以及不同滞育年限幼虫的蛋白质含量差异均不显著；滞育年周期中，无论结茧幼虫还是裸露幼虫，蛋白质含量的变化均可划分为两个阶段，第1阶段为当年6月至12月，蛋白质含量较低，平均为19．2269μg／头和19．5734μg／头；第2阶段为翌年2月底至5月中旬，蛋白质含量相对较高，平均为20．9578μg／头和20．7153μg／头。电泳图谱分析发现，SDS-PAGE 18．3kD和32．0kD蛋白带与4条PAGE蛋白带的表达量，以及SDS-PAGE 18．3kD和19．8kD蛋白带的相对表达量在滞育前后及滞育期发生了一定的变化，但在滞育期不同阶段基本无变化。
2．应用蛋白质双向电泳和质谱分析技术对麦红吸浆虫幼虫滞育前、越夏和越冬滞育期及滞育解除后4个阶段蛋白质组的表达进行了研究。结果表明，在考马斯亮蓝染色的双向电泳pH3-10胶上，PDQuest 7．4软件可识别滞育前幼虫蛋白点约300个，其它时期约275个。以滞育前幼虫为参照，越夏、越冬滞育期及滞育解除后幼虫蛋白表达量差异达2倍以上的点分别有91个、92个和95个。对3个在滞育前高表达或特异表达、2个滞育前低表达或缺失、1个冬季滞育期特异表达、2个夏季和冬季滞育期高表达的8个差异显著的斑点进行质谱测定，均获得肽质量指纹图谱，搜索NCBInr数据库初步鉴定出7个蛋白质，它们分别是阿片黑素促皮质素原，NADH脱氢酶亚基1，F10F2．5，IKK相互作用蛋白同型2，GA10647-PA，紫色CG16784-PB同型B和B0228-6，这些蛋白质在麦红吸浆虫滞育过程中发挥着重要作用。
3．用保护酶活性测试盒分别测定了麦红吸浆虫滞育前后及滞育期过氧化物酶(POD)、超氧化物歧化酶(SOD)和过氧化氢酶(CAT)等3种保护酶的活力。结果表明，幼虫从老熟到进入滞育的初期，3种保护酶的活力均呈下降趋势。滞育年周期中，SOD和CAT活力对环境温度的反应相同，即低温促进其活力升高，高温导致其活力下降；POD活力与环境温度和滞育发育有关；整个滞育期，裸露幼虫和结茧幼虫3种保护酶的活力随季节变化趋势相同，但同期的裸露幼虫活力略高于结茧幼虫；不同滞育年限幼虫3种保护酶的活力差异不显著。滞育解除后，3种保护酶的活力均随着生长发育进程逐渐升高。
4．用代谢酶活性测试盒分别测定了麦红吸浆虫滞育前后和滞育期糖原磷酸化酶(Gpase)、磷酸果糖激酶(PFK)、己糖激酶(HK)和醛缩酶(ALD)等4种代谢酶活力的变化。结果表明，麦红吸浆虫滞育前后代谢酶活力明显不同，滞育后Gpase活力显著提高，糖酵解有关酶PFK、HK和ALD活力降低，以保证更多的糖原转化为海藻糖；滞育解除后Gpase活力降低，PFK、HK和ALD活力升高，以确保昆虫生长发育所需的物质和能量供给。滞育期间，4种代谢酶的活力均与滞育发育有关；同时Gpase和PFK的活力也与环境温度有关，即夏、冬季高于春、秋季；同期不同滞育状态幼虫比较，裸露幼虫PFK、HK和ALD活力总是略高于结茧幼虫，Gpase则相反。同一时期不同滞育年限幼虫4种代谢酶的活力无显著差异。
5．用放射免疫分析法测定了麦红吸浆虫滞育前后及滞育期全虫体内蜕皮甾类激素滴度的变化。结果表明，入土滞育过程中，同期裸露幼虫与结茧幼虫，以及不同滞育年限幼虫的蜕皮甾类激素滴度差异均不显著。在年发生周期中，滞育前后及滞育期蜕皮甾类激素呈现出明显的动态变化，入土进入滞育后显著降低，入土滞育期间，11月5日以前及二次滞育幼虫较低，且变化幅度较小；冬季寒冷季节蜕皮甾类激素最高，为7．0pg／幼虫左右。滞育解除后蜕皮甾类激素滴度显著降低，之后随着幼虫的生长发育逐渐升高，至化蛹前一周左右达到最高峰，为15．22pg／幼虫。
6．应用电镜技术，对麦红吸浆虫麦穗老熟幼虫和滞育圆茧丝腺细胞进行了超微结构观察。结果发现麦穗老熟幼虫和结茧幼虫丝腺胞质中均分布有发达平行排列的粗面内质网、高尔基体和线粒体。但麦穗老熟幼虫胞质中还分布有发达的环形粗面内质网，高尔基体周围电子致密度高，糖原颗粒含量丰富，腺腔中充满了电子致密度高的丝物质；相反，结茧幼虫高尔基体周围电子致密度低，糖原颗粒含量极微，腺腔丝物质含量低。
The wheat blossom midge,Sitodiplosis mosellana(Gehin),is one of the intermittentlydisastrous insect pests injuring wheat.Two times of broad and disastrous damages wererecorded in China history in 1950's and 1980's respectively.In about ten years recently,thedisaster caused by the wheat blossom midge in a broad area was controlled effectiviely,butthe local damage was still often found in some wheat-growing regions.A trend extending itsdistribution area is obvious.Therefore,the potential threat of the wheat blossom midge to thewheat production did not disappear.It is said that diapause is one of key factors causingrampancy endangerment with intermittent and regional characteristics.However,manyproblems about the diapause mechanisms in wheat blossom midge remain uncertain.In thispaper,diapause mechanism of wheat blossom midge was conducted by means of methods andtechniques of biochemistry,proteomics,electron microscopy and radioimmunoassay on thebasis of previous studies.The present research will provide a foundation for further studies ondiapause regulation mechanism,forecasting,control of the wheat blossom midge.The mainresults were as follows.
1.The comassie brilliant blue dying,SDS-PAGE and PAGE techniques were used toanalyze protein contents and compositions in pre-diapause,diapause and post-diapause larvaeof the wheat blossom midge.The protein content in different diapause stages wassignificantly different,which in pre-diapause and post-diapause larvae were higher than thosein diapause larvae.However,no obvious difference of protein content was observed between1st and 2nd year of diapause larvae,as well as between cocooned and non-cocooned larvae inthe same diapause periods.Based on the protein contents,the larval diapause in the wheatblossom midge larvae could be divided into two stages during diapause cycle:stage 1 is fromJune to December,protein average content 19.2269μg/individual in cocooned larvae and19.5734μg/individual in non-cocooned larvae.The stage 2 from late February to mid-May ofnext year,average 20.9578 and 20.7153μg/individual respectively,both higher than stage 1.According to the electrophoresis diagrams,expressions levels of two protein bands(18.3 and32.0 kD)from SDS-PAGE,and four protein bands from PAGE,as well as relativeexpressions amounts of protein bands 18.3 and 19.8 kD from SDS-PAGE obviously changed at various periods of pre-diapause,diapause and post-diapause,but their expressions weresimilar in different stages of diapause period.
2.The proteomic approach was adopted to investigate proteins extracted from larvae ofwheat blossom midge at defferent developmental stages including pre-diapause,over-summering diapause,over-wintering diapause and post-diapause.On 2-DE gels stained bycoomassie brilliant blue,about 300 protein spots were detected in pre-diapause,and 275protein spots in each of other periods.There were 91,92 and 95 protein spots that showedmore than 2-fold changes in abundance in larvae at over-summering diapause,over-winteringdiapause and post-diapause in comparison with pre-diapause larvae.Eight protein spots,which were with most significant differences in larvae at different diapause periods,wereanalyzed by Matrix-assisted laser desorption ionization time of flight mass spectrometry(MALDI-TOF-MS).Among these eight proteins,three were up-regulated or unique duringpre-diapause,two were down-regulated or absent during pre-diapause,one was unique duringover-wintering diapause period,and the other two were up-regulated during over-summeringand over-wintering diapause stages.Seven of the eight protein spots were successfullyidentified from peptide mass fingerprints on the basis of NCBInr database.They wereproopiomelanocortin,NADH dehydrogenase subunit 1,F10F2.5,IKK interacting proteinisoform 2,GA10647-PA,purple CG16784-PB isoform B and B0228.6,respectively.Theseproteins play important roles in wheat blossom midge.
3.The activities of protective enzymes peroxidase(POD),superoxide dismutase(SOD)and catalase(CAT)in pre-diapause,diapause and post-diapause larvae of wheat blossommidge were determined by using the protective enzyme testing kits.The results indicated thatactivities of the three protective enzymes showed a decreasing trend from pre-diapause toearly diapause.In one-year cycle,SOD and CAT activities of diapause larvae had the sameresponses to environment temperature,i.e.,increased at lower temperature but decreased athigher temperature,while POD activity was related to both environmental temperature anddevelopment situation of the larvae.A similar seasonal variation trend was observed in thethree protective enzymes of both no-cocooned and cocooned larvae in one-year cycle.Thethree protective enzyme activities of no-cocooned larvae were higher than those of cocoonedlarvae at the same diapause stage.No significant difference of the three protective enzymeactivities was found between 1st and 2nd year diapause larvae.The activities of the threeprotective enzymes increased gradually with the development of post-diapause larvae.
4.The activities of metabolic enzymes Hexokinase(HK),Aldolase(ALD),Glycogenphosphorylase(Gpase)and Phosphofructokinase(PFK)in pre-diapause,diapause andpost-diapause larvae of wheat blossom midge were measured using testing kits.Flexible metabolic rearrangement was observed in diapause processing of S.mosellana.Gpase activitywas strengthened significantly,and activities of glycolysis enzymes(HK,ALK,PFK)wereweakened significantly after entering diapause,which ensured more carbon flowing intotrehalose synthesis.On the contrary,Gpase activity decreased,and activities of glycolysisenzymes increased after diapause termination,which ensured to form glycogen from trehaloseto supply energy for development.In one-year diapause cycle,activities of four enzymes wereall related with development situation of diapause larvae.Gpase and PFK were alsoinfluenced by environmental temperature.At the same diapause stage,activities of HK,ALKand PFK of no-cocooned larvae were higher than those of the cocooned larvae.On thecontrary,Gpase activities of no-cocooned larvae were lower than those of the cocooned larvae.No obvious difference of four metabolic enzyme activities was found between 1st and 2ndyear diapause larvae.
5.The ecdysteroid titers in pre-diapause,diapause and post-diapause larvae of wheatblossom midge were determined by radioimmunoassay.No obvious difference of ecdysteroidtiters was found between 1 st and 2nd year of diapause larvae,as well as between no-cocoonedand cocooned larvae in the same diapause stages.In one-year cycle,the ecdysteroid titersshowed obviously a dynamic changes,which decreased significantly after diapausing,andthose of over-wintering larvae were highest(about 7.0pg/larva)during diapausing,whilethose of other season were low and had a minor difference.The ecdysteroid titers reducedsignificantly after diapause termination,and increased gradually with the development andreached peak level(15.22 pg/larva)about one week before pupation.
6.By means of electron microscopy,ultrastructure of silk glands in S.mosellana larvaecollected from wheat ear and cocooned larvae from soil were observed.The results showedthat parallelly rough endoplasmic reticulum,golgi apparatus and mitochondrion were all verydeveloped in silk glands cell of larvae both from wheat ear and soil.Moreover,annularlyrough endoplasmic reticulum,which was very complex,was found in silk gland cells oflarvae from wheat ear.Glycogenosome,secretory granules around golgi apparatus,and silkmaterial of glandular cavity were richer in silk glands cell of larvae from wheat ear than thoseof cocooned larvae.
1．应用考马斯亮蓝G-250染色法、SDS-PAGE及PAGE电泳技术分析了麦红吸浆虫滞育前后及滞育期蛋白质含量和组分的变化。结果表明，不同阶段蛋白质含量存在显著差异，老熟幼虫入土前和滞育解除后均高于入土滞育期。入土滞育期间，同期裸露幼虫与结茧幼虫，以及不同滞育年限幼虫的蛋白质含量差异均不显著；滞育年周期中，无论结茧幼虫还是裸露幼虫，蛋白质含量的变化均可划分为两个阶段，第1阶段为当年6月至12月，蛋白质含量较低，平均为19．2269μg／头和19．5734μg／头；第2阶段为翌年2月底至5月中旬，蛋白质含量相对较高，平均为20．9578μg／头和20．7153μg／头。电泳图谱分析发现，SDS-PAGE 18．3kD和32．0kD蛋白带与4条PAGE蛋白带的表达量，以及SDS-PAGE 18．3kD和19．8kD蛋白带的相对表达量在滞育前后及滞育期发生了一定的变化，但在滞育期不同阶段基本无变化。
2．应用蛋白质双向电泳和质谱分析技术对麦红吸浆虫幼虫滞育前、越夏和越冬滞育期及滞育解除后4个阶段蛋白质组的表达进行了研究。结果表明，在考马斯亮蓝染色的双向电泳pH3-10胶上，PDQuest 7．4软件可识别滞育前幼虫蛋白点约300个，其它时期约275个。以滞育前幼虫为参照，越夏、越冬滞育期及滞育解除后幼虫蛋白表达量差异达2倍以上的点分别有91个、92个和95个。对3个在滞育前高表达或特异表达、2个滞育前低表达或缺失、1个冬季滞育期特异表达、2个夏季和冬季滞育期高表达的8个差异显著的斑点进行质谱测定，均获得肽质量指纹图谱，搜索NCBInr数据库初步鉴定出7个蛋白质，它们分别是阿片黑素促皮质素原，NADH脱氢酶亚基1，F10F2．5，IKK相互作用蛋白同型2，GA10647-PA，紫色CG16784-PB同型B和B0228-6，这些蛋白质在麦红吸浆虫滞育过程中发挥着重要作用。
3．用保护酶活性测试盒分别测定了麦红吸浆虫滞育前后及滞育期过氧化物酶(POD)、超氧化物歧化酶(SOD)和过氧化氢酶(CAT)等3种保护酶的活力。结果表明，幼虫从老熟到进入滞育的初期，3种保护酶的活力均呈下降趋势。滞育年周期中，SOD和CAT活力对环境温度的反应相同，即低温促进其活力升高，高温导致其活力下降；POD活力与环境温度和滞育发育有关；整个滞育期，裸露幼虫和结茧幼虫3种保护酶的活力随季节变化趋势相同，但同期的裸露幼虫活力略高于结茧幼虫；不同滞育年限幼虫3种保护酶的活力差异不显著。滞育解除后，3种保护酶的活力均随着生长发育进程逐渐升高。
4．用代谢酶活性测试盒分别测定了麦红吸浆虫滞育前后和滞育期糖原磷酸化酶(Gpase)、磷酸果糖激酶(PFK)、己糖激酶(HK)和醛缩酶(ALD)等4种代谢酶活力的变化。结果表明，麦红吸浆虫滞育前后代谢酶活力明显不同，滞育后Gpase活力显著提高，糖酵解有关酶PFK、HK和ALD活力降低，以保证更多的糖原转化为海藻糖；滞育解除后Gpase活力降低，PFK、HK和ALD活力升高，以确保昆虫生长发育所需的物质和能量供给。滞育期间，4种代谢酶的活力均与滞育发育有关；同时Gpase和PFK的活力也与环境温度有关，即夏、冬季高于春、秋季；同期不同滞育状态幼虫比较，裸露幼虫PFK、HK和ALD活力总是略高于结茧幼虫，Gpase则相反。同一时期不同滞育年限幼虫4种代谢酶的活力无显著差异。
5．用放射免疫分析法测定了麦红吸浆虫滞育前后及滞育期全虫体内蜕皮甾类激素滴度的变化。结果表明，入土滞育过程中，同期裸露幼虫与结茧幼虫，以及不同滞育年限幼虫的蜕皮甾类激素滴度差异均不显著。在年发生周期中，滞育前后及滞育期蜕皮甾类激素呈现出明显的动态变化，入土进入滞育后显著降低，入土滞育期间，11月5日以前及二次滞育幼虫较低，且变化幅度较小；冬季寒冷季节蜕皮甾类激素最高，为7．0pg／幼虫左右。滞育解除后蜕皮甾类激素滴度显著降低，之后随着幼虫的生长发育逐渐升高，至化蛹前一周左右达到最高峰，为15．22pg／幼虫。
6．应用电镜技术，对麦红吸浆虫麦穗老熟幼虫和滞育圆茧丝腺细胞进行了超微结构观察。结果发现麦穗老熟幼虫和结茧幼虫丝腺胞质中均分布有发达平行排列的粗面内质网、高尔基体和线粒体。但麦穗老熟幼虫胞质中还分布有发达的环形粗面内质网，高尔基体周围电子致密度高，糖原颗粒含量丰富，腺腔中充满了电子致密度高的丝物质；相反，结茧幼虫高尔基体周围电子致密度低，糖原颗粒含量极微，腺腔丝物质含量低。
The wheat blossom midge,Sitodiplosis mosellana(Gehin),is one of the intermittentlydisastrous insect pests injuring wheat.Two times of broad and disastrous damages wererecorded in China history in 1950's and 1980's respectively.In about ten years recently,thedisaster caused by the wheat blossom midge in a broad area was controlled effectiviely,butthe local damage was still often found in some wheat-growing regions.A trend extending itsdistribution area is obvious.Therefore,the potential threat of the wheat blossom midge to thewheat production did not disappear.It is said that diapause is one of key factors causingrampancy endangerment with intermittent and regional characteristics.However,manyproblems about the diapause mechanisms in wheat blossom midge remain uncertain.In thispaper,diapause mechanism of wheat blossom midge was conducted by means of methods andtechniques of biochemistry,proteomics,electron microscopy and radioimmunoassay on thebasis of previous studies.The present research will provide a foundation for further studies ondiapause regulation mechanism,forecasting,control of the wheat blossom midge.The mainresults were as follows.
1.The comassie brilliant blue dying,SDS-PAGE and PAGE techniques were used toanalyze protein contents and compositions in pre-diapause,diapause and post-diapause larvaeof the wheat blossom midge.The protein content in different diapause stages wassignificantly different,which in pre-diapause and post-diapause larvae were higher than thosein diapause larvae.However,no obvious difference of protein content was observed between1st and 2nd year of diapause larvae,as well as between cocooned and non-cocooned larvae inthe same diapause periods.Based on the protein contents,the larval diapause in the wheatblossom midge larvae could be divided into two stages during diapause cycle:stage 1 is fromJune to December,protein average content 19.2269μg/individual in cocooned larvae and19.5734μg/individual in non-cocooned larvae.The stage 2 from late February to mid-May ofnext year,average 20.9578 and 20.7153μg/individual respectively,both higher than stage 1.According to the electrophoresis diagrams,expressions levels of two protein bands(18.3 and32.0 kD)from SDS-PAGE,and four protein bands from PAGE,as well as relativeexpressions amounts of protein bands 18.3 and 19.8 kD from SDS-PAGE obviously changed at various periods of pre-diapause,diapause and post-diapause,but their expressions weresimilar in different stages of diapause period.
2.The proteomic approach was adopted to investigate proteins extracted from larvae ofwheat blossom midge at defferent developmental stages including pre-diapause,over-summering diapause,over-wintering diapause and post-diapause.On 2-DE gels stained bycoomassie brilliant blue,about 300 protein spots were detected in pre-diapause,and 275protein spots in each of other periods.There were 91,92 and 95 protein spots that showedmore than 2-fold changes in abundance in larvae at over-summering diapause,over-winteringdiapause and post-diapause in comparison with pre-diapause larvae.Eight protein spots,which were with most significant differences in larvae at different diapause periods,wereanalyzed by Matrix-assisted laser desorption ionization time of flight mass spectrometry(MALDI-TOF-MS).Among these eight proteins,three were up-regulated or unique duringpre-diapause,two were down-regulated or absent during pre-diapause,one was unique duringover-wintering diapause period,and the other two were up-regulated during over-summeringand over-wintering diapause stages.Seven of the eight protein spots were successfullyidentified from peptide mass fingerprints on the basis of NCBInr database.They wereproopiomelanocortin,NADH dehydrogenase subunit 1,F10F2.5,IKK interacting proteinisoform 2,GA10647-PA,purple CG16784-PB isoform B and B0228.6,respectively.Theseproteins play important roles in wheat blossom midge.
3.The activities of protective enzymes peroxidase(POD),superoxide dismutase(SOD)and catalase(CAT)in pre-diapause,diapause and post-diapause larvae of wheat blossommidge were determined by using the protective enzyme testing kits.The results indicated thatactivities of the three protective enzymes showed a decreasing trend from pre-diapause toearly diapause.In one-year cycle,SOD and CAT activities of diapause larvae had the sameresponses to environment temperature,i.e.,increased at lower temperature but decreased athigher temperature,while POD activity was related to both environmental temperature anddevelopment situation of the larvae.A similar seasonal variation trend was observed in thethree protective enzymes of both no-cocooned and cocooned larvae in one-year cycle.Thethree protective enzyme activities of no-cocooned larvae were higher than those of cocoonedlarvae at the same diapause stage.No significant difference of the three protective enzymeactivities was found between 1st and 2nd year diapause larvae.The activities of the threeprotective enzymes increased gradually with the development of post-diapause larvae.
4.The activities of metabolic enzymes Hexokinase(HK),Aldolase(ALD),Glycogenphosphorylase(Gpase)and Phosphofructokinase(PFK)in pre-diapause,diapause andpost-diapause larvae of wheat blossom midge were measured using testing kits.Flexible metabolic rearrangement was observed in diapause processing of S.mosellana.Gpase activitywas strengthened significantly,and activities of glycolysis enzymes(HK,ALK,PFK)wereweakened significantly after entering diapause,which ensured more carbon flowing intotrehalose synthesis.On the contrary,Gpase activity decreased,and activities of glycolysisenzymes increased after diapause termination,which ensured to form glycogen from trehaloseto supply energy for development.In one-year diapause cycle,activities of four enzymes wereall related with development situation of diapause larvae.Gpase and PFK were alsoinfluenced by environmental temperature.At the same diapause stage,activities of HK,ALKand PFK of no-cocooned larvae were higher than those of the cocooned larvae.On thecontrary,Gpase activities of no-cocooned larvae were lower than those of the cocooned larvae.No obvious difference of four metabolic enzyme activities was found between 1st and 2ndyear diapause larvae.
5.The ecdysteroid titers in pre-diapause,diapause and post-diapause larvae of wheatblossom midge were determined by radioimmunoassay.No obvious difference of ecdysteroidtiters was found between 1 st and 2nd year of diapause larvae,as well as between no-cocoonedand cocooned larvae in the same diapause stages.In one-year cycle,the ecdysteroid titersshowed obviously a dynamic changes,which decreased significantly after diapausing,andthose of over-wintering larvae were highest(about 7.0pg/larva)during diapausing,whilethose of other season were low and had a minor difference.The ecdysteroid titers reducedsignificantly after diapause termination,and increased gradually with the development andreached peak level(15.22 pg/larva)about one week before pupation.
6.By means of electron microscopy,ultrastructure of silk glands in S.mosellana larvaecollected from wheat ear and cocooned larvae from soil were observed.The results showedthat parallelly rough endoplasmic reticulum,golgi apparatus and mitochondrion were all verydeveloped in silk glands cell of larvae both from wheat ear and soil.Moreover,annularlyrough endoplasmic reticulum,which was very complex,was found in silk gland cells oflarvae from wheat ear.Glycogenosome,secretory granules around golgi apparatus,and silkmaterial of glandular cavity were richer in silk glands cell of larvae from wheat ear than thoseof cocooned larvae.
引文
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