家蚕分子连锁图谱的构建和低分子量热激蛋白基因的克隆与特异表达
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摘要
家蚕是重要的经济昆虫，也是生物学研究的极好材料。家蚕形态遗传学研究
历经80余年，已有300余个形态性状基因定位于传统的遗传连锁图上。近年来，
随着分子生物学的发展，日、美、中等国的学者在家蚕的分子遗传学研究方面已
取得了不少进展。有关的基因克隆，基因定位和分子育种的研究，必将成为家蚕
遗传学的主要研究方向。分子连锁图的构建可以极大地推动家蚕分子生物学的研
究。分子连锁图也是家蚕遗传学基础理论研究与实际应用之间的重要桥梁。它能
对控制品质的数量基因和抗病基因进行分析、定位；能通过分子标记进行选择，
快速制定出富有成效的育种计划；而且我们能够借助分子连锁图提供的指导进行
基因克隆和外源基因导入等方面的研究。本文的主要内容是在建立家蚕RAPD分
子标记筛选方法和技术体系的基础上，构建了我国第一张家蚕的RAPD分子连锁
图；并运用荧光差异显示法（flourescent differential display, FDD），在mRNA水平探
讨肾脏型卵（ki）胚胎致死与正常卵在初期胚胎阶段的基因表达，分离，克隆目的
基因；家蚕ki卵在卵形与胚子两方面都发生异常，是研究胚子发生期中组织分化
与基因表达的良好材料，利用它可阐明基因的表达开关与基因的调控机理，我们
在对ki胚胎致死的FDD分析过程中克隆了2种低分子量热激蛋白基因，即
Bmhsp20.8A与Bmshp20.8B；进而讨论了6种sHSPs基因在家蚕体内的表达，并对
其功能进行分析；同时也对家蚕shsps及其他物种的shsps进行了系统进化分析，
表明shsps以直向同源进化为主，亦存在并向同源进化关系。主要结果如下：
1 分子标记的筛选
1.1RAPD扩增在不同的物种中其反应体系略有不同，经优化选择，确定了适于家
蚕DNA的PCR扩增反应体系及条件。
1.2用477个RAPD引物中的410个引物在两亲本间共得扩增片段5155条，多态
性片段共1496条，占总片段数的29.0％，平均每个引物扩增多态性片段3.6条，
83.6％的RAPD引物在大造和C_(108)间可产生多态性产物。
1.3 家蚕RAPD的遗传方式为显隐遗传模式。80.2％的标记符合3：1的分离比率，
显示孟德尔遗传，发生偏离的标记为19.8％。
MQ
二 家蚕分子连锁图的构建
＊门7个R八卜日引物X69个h 们f小进刊‘5 广仇L师地，W列厂＊h个分 广
标t，然后通过Mapmaker／Exp（Version3．oh）软件构建连锁图，结果如卜：
2．1425个 RAPD随机引物，其中 137个引物（32．3％）在两亲本问的扩增结欲有
5“个多态性位点出现。平均为 4．3 ｛y点／引物。在 5％水平匕经／检测后，有 170
个位点，占总数的80．2％符合3：1的分离比，其中大造255个，乙m215个。
2．2本研究经多次验证发现，只要保持反应条件不变，严格控制反应程序的各个环
节，保证反应体系中的多种试剂来源和浓度一致并不受污染，则可保证其扩地有
较好的稳定性与重复性。
2．3 对470个RAPO多舟叮标记位点用Mapmiik。卜软件进行连锁分析，构建了家蚕
的分于连锁图，结果如下：
2．3．二 将255个来自大造，2 15个米自h。的标匕分为北个连锁群．大造的u个逐
钓Z群命名为a仆群．C．．；。NJ！6个连钓i群命名为 h仆群n介 l业群中响 1（以个什）！．I、连
锁，非连锁的位点有 152个。在 b亚群中有 79个位点连锁，非连锁位点有 126个。
2．3．2 d亚群中各连锁群上标记数目变化范困在2～24个，连锁群的长度变化在
13．SCM～163．6CM，标记的平均图距为12．87CM～20．4CM，平均图距大于10CM ’J’于
20CM的连锁群有18个，大十20CM的连锁群有5个。23个连锁群上位点平均数为
4．5个，连锁群的平均长度为49．7cM。
2．3．3 b亚群中各连锁群上标记数目变化范围在 2～12个，连锁群的长度变化在
6．ZCM～235．ZCM，标记的平均图距为 6．ZCM～ZI．38CM，平均图距小于 10CM的连
锁群有 2个，大于 10。M小于 20州的连锁群有 11个，16个连锁群上位点平均数为
1．9个，还锁群的平均长度为69．（）CM。
2．3．4 在 a亚群中，标记数目与连锁群长度进行相关分析，相关系数巳为 0．9958，
在b亚群中，进行同样的相关分析，相关系数R。为0．9705。这表明了连锁群的长
仪‘J仰匕数H的多少呈小相人。
2．3．5 在 a、b两亚群中，分别对每连锁群的平均遗传图距对每染色体上的札记
数进行回归分析。回归系数分别为－0．6269，0．1269，表明了连锁群的平均图趴与
每染色体卜的标记数无百接关系。
2．3．6。。业价l－It 23个0锁群的总KIA为 1192．SCM，b业仪为 1104．ZCM。y均为
Silkworm(Bombyx mon.) is an important economic insect and an ideal model
organism for the biology study. The morphologic genetics of silkworm has been studied
for more than 80 years, and over 300 genes for morphological traits have been localized
on the traditional genetic linkage map. On the development of molecular biology, a lot
of progress has been made on the molecular genetics of silkworm in many countries
such as Japan, America and China. Those studies related to gene cloning, gene location
and molecular breeding are sure to become the major research field for silkworm
genetics. Construction of the molecular linkage map may greatly promote the molecular
biology research in silkworm. Furthermore, it is also an important bridge between basic
and applied research in silkworm. It can carry out the analysis and location of QTLs for
quality or resistance to diseases. It may make an efficient breeding possible through the
selection of molecular markers, and it may also carry out the research of cloning genes
and introduction of foreign genes. The following are the major content of the paper: On
the basis of the establishment of the method of screening RAPD markers, the first
molecular linkage map of silkworm in our country was constructed; FDD(Fluorescent
differential display) was used to study the gene expression of embryonic lethality of ki
and normal egg during the early embryonic stage at mRNA level and to separate the
target gene and clone it. Two small heat shock genes, Bmhsp20.8A and Bmhsp20.8B,
were cloned in the FDD analysis of the embryonic lethality of ki. And the expression of
sHSPs genes was studied in the silkworm, and their functions were analyzed.
Simultaneously systematic evolution of shsps in the silkworm and other species were
analyzed, the results showed that the directly homologous evolution of shsps accounted
for the major, but the simultaneously homologous existed too.
I.Screening for the molecular markers
Random amplified polymorphic DNA, which was known as RAPD, was used to
screen for the DNA markers of silkworm. Using the RAPD, a marker could be
amplified by a single primer (normally 10 bases) at low annealing temperature when
there is a sequence on opposite strand no more than 3000 base pairs in an inverted
orientation. The RAPD method allows a rapid identification of DNA markers and
provides an efficient assay for polymorphism. A large amount of RAPD markers were
obtained using the method to screen the silkworm genome.
1.1 Reaction conditions of RAPD are different in different species. The optimized
RAPD reaction conditions suitable for silkworm were established for silkworm genoniic
DNA in the study.
1.2 With 477 RAPD primers of which each contain 60%?0%GC, the RAPD markers
in C108,Dazao and F1 of Dazao x C108 were screened. Among the 410 of 477 R.APD
primers we identified 5155 amplified DNA fragments. The average number of amplified
DNA fragments was 12.6, varying from 1 band to 23 bands. Among them 1496
amplified DNA bands (29.0% of the total amplified fragments) showed polymorphism
with a mean number fragments of 3.6 per primer. Among the 477 primers 67 of them
produced no bands with the percentage of 14.1, 11 primers which showed no
polymorphism with the percentage of 2.3, and the rest 83.6% could produce bands
showing polymorphism.
1.3 The genetic pattern of RAPD serves as the basis for localizing genes and
constructing the molecular linkage map. The study shows that RAPD markers have the
pattern of dominance and recess
历经80余年，已有300余个形态性状基因定位于传统的遗传连锁图上。近年来，
随着分子生物学的发展，日、美、中等国的学者在家蚕的分子遗传学研究方面已
取得了不少进展。有关的基因克隆，基因定位和分子育种的研究，必将成为家蚕
遗传学的主要研究方向。分子连锁图的构建可以极大地推动家蚕分子生物学的研
究。分子连锁图也是家蚕遗传学基础理论研究与实际应用之间的重要桥梁。它能
对控制品质的数量基因和抗病基因进行分析、定位；能通过分子标记进行选择，
快速制定出富有成效的育种计划；而且我们能够借助分子连锁图提供的指导进行
基因克隆和外源基因导入等方面的研究。本文的主要内容是在建立家蚕RAPD分
子标记筛选方法和技术体系的基础上，构建了我国第一张家蚕的RAPD分子连锁
图；并运用荧光差异显示法（flourescent differential display, FDD），在mRNA水平探
讨肾脏型卵（ki）胚胎致死与正常卵在初期胚胎阶段的基因表达，分离，克隆目的
基因；家蚕ki卵在卵形与胚子两方面都发生异常，是研究胚子发生期中组织分化
与基因表达的良好材料，利用它可阐明基因的表达开关与基因的调控机理，我们
在对ki胚胎致死的FDD分析过程中克隆了2种低分子量热激蛋白基因，即
Bmhsp20.8A与Bmshp20.8B；进而讨论了6种sHSPs基因在家蚕体内的表达，并对
其功能进行分析；同时也对家蚕shsps及其他物种的shsps进行了系统进化分析，
表明shsps以直向同源进化为主，亦存在并向同源进化关系。主要结果如下：
1 分子标记的筛选
1.1RAPD扩增在不同的物种中其反应体系略有不同，经优化选择，确定了适于家
蚕DNA的PCR扩增反应体系及条件。
1.2用477个RAPD引物中的410个引物在两亲本间共得扩增片段5155条，多态
性片段共1496条，占总片段数的29.0％，平均每个引物扩增多态性片段3.6条，
83.6％的RAPD引物在大造和C_(108)间可产生多态性产物。
1.3 家蚕RAPD的遗传方式为显隐遗传模式。80.2％的标记符合3：1的分离比率，
显示孟德尔遗传，发生偏离的标记为19.8％。
MQ
二 家蚕分子连锁图的构建
＊门7个R八卜日引物X69个h 们f小进刊‘5 广仇L师地，W列厂＊h个分 广
标t，然后通过Mapmaker／Exp（Version3．oh）软件构建连锁图，结果如卜：
2．1425个 RAPD随机引物，其中 137个引物（32．3％）在两亲本问的扩增结欲有
5“个多态性位点出现。平均为 4．3 ｛y点／引物。在 5％水平匕经／检测后，有 170
个位点，占总数的80．2％符合3：1的分离比，其中大造255个，乙m215个。
2．2本研究经多次验证发现，只要保持反应条件不变，严格控制反应程序的各个环
节，保证反应体系中的多种试剂来源和浓度一致并不受污染，则可保证其扩地有
较好的稳定性与重复性。
2．3 对470个RAPO多舟叮标记位点用Mapmiik。卜软件进行连锁分析，构建了家蚕
的分于连锁图，结果如下：
2．3．二 将255个来自大造，2 15个米自h。的标匕分为北个连锁群．大造的u个逐
钓Z群命名为a仆群．C．．；。NJ！6个连钓i群命名为 h仆群n介 l业群中响 1（以个什）！．I、连
锁，非连锁的位点有 152个。在 b亚群中有 79个位点连锁，非连锁位点有 126个。
2．3．2 d亚群中各连锁群上标记数目变化范困在2～24个，连锁群的长度变化在
13．SCM～163．6CM，标记的平均图距为12．87CM～20．4CM，平均图距大于10CM ’J’于
20CM的连锁群有18个，大十20CM的连锁群有5个。23个连锁群上位点平均数为
4．5个，连锁群的平均长度为49．7cM。
2．3．3 b亚群中各连锁群上标记数目变化范围在 2～12个，连锁群的长度变化在
6．ZCM～235．ZCM，标记的平均图距为 6．ZCM～ZI．38CM，平均图距小于 10CM的连
锁群有 2个，大于 10。M小于 20州的连锁群有 11个，16个连锁群上位点平均数为
1．9个，还锁群的平均长度为69．（）CM。
2．3．4 在 a亚群中，标记数目与连锁群长度进行相关分析，相关系数巳为 0．9958，
在b亚群中，进行同样的相关分析，相关系数R。为0．9705。这表明了连锁群的长
仪‘J仰匕数H的多少呈小相人。
2．3．5 在 a、b两亚群中，分别对每连锁群的平均遗传图距对每染色体上的札记
数进行回归分析。回归系数分别为－0．6269，0．1269，表明了连锁群的平均图趴与
每染色体卜的标记数无百接关系。
2．3．6。。业价l－It 23个0锁群的总KIA为 1192．SCM，b业仪为 1104．ZCM。y均为
Silkworm(Bombyx mon.) is an important economic insect and an ideal model
organism for the biology study. The morphologic genetics of silkworm has been studied
for more than 80 years, and over 300 genes for morphological traits have been localized
on the traditional genetic linkage map. On the development of molecular biology, a lot
of progress has been made on the molecular genetics of silkworm in many countries
such as Japan, America and China. Those studies related to gene cloning, gene location
and molecular breeding are sure to become the major research field for silkworm
genetics. Construction of the molecular linkage map may greatly promote the molecular
biology research in silkworm. Furthermore, it is also an important bridge between basic
and applied research in silkworm. It can carry out the analysis and location of QTLs for
quality or resistance to diseases. It may make an efficient breeding possible through the
selection of molecular markers, and it may also carry out the research of cloning genes
and introduction of foreign genes. The following are the major content of the paper: On
the basis of the establishment of the method of screening RAPD markers, the first
molecular linkage map of silkworm in our country was constructed; FDD(Fluorescent
differential display) was used to study the gene expression of embryonic lethality of ki
and normal egg during the early embryonic stage at mRNA level and to separate the
target gene and clone it. Two small heat shock genes, Bmhsp20.8A and Bmhsp20.8B,
were cloned in the FDD analysis of the embryonic lethality of ki. And the expression of
sHSPs genes was studied in the silkworm, and their functions were analyzed.
Simultaneously systematic evolution of shsps in the silkworm and other species were
analyzed, the results showed that the directly homologous evolution of shsps accounted
for the major, but the simultaneously homologous existed too.
I.Screening for the molecular markers
Random amplified polymorphic DNA, which was known as RAPD, was used to
screen for the DNA markers of silkworm. Using the RAPD, a marker could be
amplified by a single primer (normally 10 bases) at low annealing temperature when
there is a sequence on opposite strand no more than 3000 base pairs in an inverted
orientation. The RAPD method allows a rapid identification of DNA markers and
provides an efficient assay for polymorphism. A large amount of RAPD markers were
obtained using the method to screen the silkworm genome.
1.1 Reaction conditions of RAPD are different in different species. The optimized
RAPD reaction conditions suitable for silkworm were established for silkworm genoniic
DNA in the study.
1.2 With 477 RAPD primers of which each contain 60%?0%GC, the RAPD markers
in C108,Dazao and F1 of Dazao x C108 were screened. Among the 410 of 477 R.APD
primers we identified 5155 amplified DNA fragments. The average number of amplified
DNA fragments was 12.6, varying from 1 band to 23 bands. Among them 1496
amplified DNA bands (29.0% of the total amplified fragments) showed polymorphism
with a mean number fragments of 3.6 per primer. Among the 477 primers 67 of them
produced no bands with the percentage of 14.1, 11 primers which showed no
polymorphism with the percentage of 2.3, and the rest 83.6% could produce bands
showing polymorphism.
1.3 The genetic pattern of RAPD serves as the basis for localizing genes and
constructing the molecular linkage map. The study shows that RAPD markers have the
pattern of dominance and recess
引文
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2．陈洪，朱立煌，徐吉臣等．(1995)．RAPD标记构建水稻分子连锁图．植物学报，37(9)：677～684
3．陈常庆，杨香娇．(1989)．DNA体外扩增技术的突破——聚合酶连反应(PCR)的发展与应用，生物科学信息，1(3)：14～17
4．陈受宜，朱立煌等．(1991．水稻抗盐突变体的分子生物学鉴定，植物学报，33(8)，569～573
5．董海涛，何祖华，吴玉良等．(1998)．水稻抗稻瘟病近等基因系mRNA差别显示分析．农业生物技术学报，6(3)：223-228
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