稀有鮈鲫（Gobiocypris rarus）脑垂体结构及POMC基因cDNA序列克隆和分析
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摘要
稀有鮈鲫(Gobiocypris rarus)隶属于鲤形目(Cypriniformes)鲤科(Cyprinidae)鱼丹亚科(Canioninae)鮈鲫属,是中国特有的小型淡水鱼类。
运用常规组织学方法,以Mallory's一步三色法,AF染色法为主,辅于铅苏木精等方法,并以HE法为对照,观察了稀有鮈鲫脑垂体的结构。稀有鮈鲫的脑垂体呈肾形,由神经垂体和腺垂体组成,整个垂体通过垂体柄与下丘脑相连。腺垂体为垂体前外侧部(RPD),垂体中外侧部(PPD)和垂体中间部(PI)三部分。神经垂体及其分支深入腺垂体的各部分,在PI中分布尤为丰富。分布于整个脑垂体中的血管丰富。
克隆各种垂体激素cDNA序列,并设计相应的反义RNA探针。切片原位杂交结果显示,PRL阳性细胞主要分布在RPD区域,在PPD中也有少量零散分布。POMC阳性细胞主要分布于RPD和PI区域,RPD内的POMC阳性细胞形状不规则,呈分散状,偶有几个细胞积聚成团；PI内的细胞沿着神经垂体分支分布,细胞较大,胞质染色深。另外,在PPD中也发现了少量的POMC阳性细胞。GH阳性细胞主要成群分布在PPD中,在RPD中也有极少量分布。细胞圆形,椭圆形或三角形。FSHβ阳性细胞主要分布在RPD、PPD区域,PI中有少量分布。细胞圆形,椭圆形或不规则形,单个或者少数细胞成团分布。LHβ主要是单个或者集群分布在PPD中,在PRD和PI中有零星分布。分布于PPD中的阳性细胞形成一规则弧形,成为与PI的分界线。TSHβ在整个垂体中都有分布,在PPD背面集群分布,在RPD和PI中单个零散分布。GTHα阳性细胞分布范围广,在RPD、PI、PPD中均有分布；以PPD数量最多,且与LHp相似,分布于PPD后缘的阳性细胞形成一弧形,成为PPD与PI的明显界限。SL阳性细胞数量较少,仅在PI后缘腹面有少量阳性细胞成团或分散分布。垂体激素产生细胞在垂体中的分布具有个体差异性。
运用RT-PCR和RACE方法从稀有鮈鲫脑垂体中克隆出了POMC基因的cDNA序列。得到的稀有鮈鲫POMC(rmPOMC)cDNA长846bps,ORF长666bps,编码信号肽(SB，Met1-Ala28),N-端多肽(Gln29-His105),ACTH(Ser108-Met146),α-MSH (Ser108-Gal121),CLIP (Pro126-Met146),β-LPH(Glu149-His221),γ-LPH(Glul49-Ser186),β-MSH (Asp170-Ser186)和β-内啡肽(β-EP,Tyr189-Gln221)。没有发现与γ-MSH(四足类POMC的特点)同源的序列存在。多序列同源性分析发现氨基酸序列与鲤鱼的POMC-Ⅰ和POMC-Ⅱ具有最高的相似性,为92.4％,支持了稀有鮈鲫应属于鮈亚科的观点。系统进化树分析表明稀有鮈鲫的POMC序列属于POMCα。
运用RT-PCR和原位杂交技术分析了rmPOMC基因的垂体和垂体外组织表达情况。阳性细胞主要分布在RPD和PI,也有少数阳性细胞分布在PPD。垂体外组织,包括脑、小肠、性腺、肝胰脏、脾脏和鳃都检测到了rmPOMC mRNA的表达,表明在这些组织中,POMC存在旁分泌作用方式。
Rare minnow (Gobiocypris rarus), belonging to order of Cypriniformes, family of Cyprinidae, subfamily Danioninae, genus Gobiocypris, is a small freshwater fish endemic to China.
The structure of rare minnow pituitary was observed by histology and histochemistry methods. As a kidney, the pituitary of rare minnow was comprised of neurohypophysis and adenohypophysis, attaching to the hypothalamus by a pituitary stalk. Adenohypophysis was divided into three parts, RPD, PPD and PI according to the distinct histochemistrical characteristics. The curved branches of neurohypophysis were inserted in every part of adenohypophysis, and were most plentiful in PI. The blood vessels were rich in all parts of the pituitary.
cRNA probes were designed according to the cloned cDNAs of pituitary hormones. Data shows that:PRL positive cells were mainly distributed in RPD, and few were scattered in PPD. POMC positive cells located in most part of RPD and PI, with patterns of dispersed and compact along the PN branches, respectively. Inaddition, there were some POMC positive cells observed in PPD. GH positive cells were mainly grouped in PPD, few found in RPD, in a shape of round, oval or triangle. FSHβpositive cells located in RPD, PPD and few in PI, in a shape of round, oval or irregular. LHP positive cells distributed in PPD alone or in group, and few cells were also found in RPD and PI. There was a dividing line in PPD and PI, formed by LHβpositive cells. TSHPmainly grouped in the dorsal part of PPD, few scattered in RPD and PI. GTHapositive cells with a large distributing area, detected in every part of the pituitary, mostly in PPD, and formed a distinct limit between PPD and PI like the LHβ positive cells. There were few SL positive cells detected, and these cells were grouped or scattered only in posterior-ventral par of PI. The distributing patterns of hormone producing cells were different from each samples.
A cDNA encoding pro-opiomelanocortin (POMC) gene was cloned from the pituitary gland of the rare minnow (Gobiocypris rarus), a small freshwater fish endemic to China. This was achieved by reverse transcription polymerase chain reaction (RT-PCR) and rapid amplification of cDNA ends (RACE). Data showed that the predicted rare minnow POMC (rmPOMC) cDNA consisted of 846bps coding for the following sequences, flanked by proteolytic cleavage sites:signal peptide (SP, Met1-Ala28), N-terminal peptide (Gln29-His105), ACTH (Ser108-Met146),α-MSH (Ser108-Gal121), CLIP (Pro126-Met146),β-LPH (Glu149-His221),γ-LPH (Glul49-Ser186),β-MSH (Asp 170-Ser186), andβ-endorphin (β-EP, Tyr189-Gln221). Sequence analysis showed no region was homologous toγ-MSH (a tetrapod POMC feature). The amino acid sequence is highly similar to POMC-Ⅰand POMC-Ⅱof the common carp (92.4%), which supporting the viewpoint of rare minnow belongs to subfamily Gobioninae according to homologous alignment. It was POMCαthrough the phylogenetic analysis.
Pituitary and extra-pituitary expression were studied using RT-PCR and in situ hybridization. rmPOMC positive cells were mainly located in the rostral pars distalis (RPD) and pars intermedia (PI). Some rmPOMC positive cells were detected in the proximal pars distalis (PPD) as well, according to in situ hybridization. In the extra-pituitary tissues, positive signals were observed in the brain, intestines, gonads, hepatopancreas, spleen, and gills by RT-PCR analysis, showing that the paracrine pathway was existed in these tissues.
运用常规组织学方法,以Mallory's一步三色法,AF染色法为主,辅于铅苏木精等方法,并以HE法为对照,观察了稀有鮈鲫脑垂体的结构。稀有鮈鲫的脑垂体呈肾形,由神经垂体和腺垂体组成,整个垂体通过垂体柄与下丘脑相连。腺垂体为垂体前外侧部(RPD),垂体中外侧部(PPD)和垂体中间部(PI)三部分。神经垂体及其分支深入腺垂体的各部分,在PI中分布尤为丰富。分布于整个脑垂体中的血管丰富。
克隆各种垂体激素cDNA序列,并设计相应的反义RNA探针。切片原位杂交结果显示,PRL阳性细胞主要分布在RPD区域,在PPD中也有少量零散分布。POMC阳性细胞主要分布于RPD和PI区域,RPD内的POMC阳性细胞形状不规则,呈分散状,偶有几个细胞积聚成团；PI内的细胞沿着神经垂体分支分布,细胞较大,胞质染色深。另外,在PPD中也发现了少量的POMC阳性细胞。GH阳性细胞主要成群分布在PPD中,在RPD中也有极少量分布。细胞圆形,椭圆形或三角形。FSHβ阳性细胞主要分布在RPD、PPD区域,PI中有少量分布。细胞圆形,椭圆形或不规则形,单个或者少数细胞成团分布。LHβ主要是单个或者集群分布在PPD中,在PRD和PI中有零星分布。分布于PPD中的阳性细胞形成一规则弧形,成为与PI的分界线。TSHβ在整个垂体中都有分布,在PPD背面集群分布,在RPD和PI中单个零散分布。GTHα阳性细胞分布范围广,在RPD、PI、PPD中均有分布；以PPD数量最多,且与LHp相似,分布于PPD后缘的阳性细胞形成一弧形,成为PPD与PI的明显界限。SL阳性细胞数量较少,仅在PI后缘腹面有少量阳性细胞成团或分散分布。垂体激素产生细胞在垂体中的分布具有个体差异性。
运用RT-PCR和RACE方法从稀有鮈鲫脑垂体中克隆出了POMC基因的cDNA序列。得到的稀有鮈鲫POMC(rmPOMC)cDNA长846bps,ORF长666bps,编码信号肽(SB，Met1-Ala28),N-端多肽(Gln29-His105),ACTH(Ser108-Met146),α-MSH (Ser108-Gal121),CLIP (Pro126-Met146),β-LPH(Glu149-His221),γ-LPH(Glul49-Ser186),β-MSH (Asp170-Ser186)和β-内啡肽(β-EP,Tyr189-Gln221)。没有发现与γ-MSH(四足类POMC的特点)同源的序列存在。多序列同源性分析发现氨基酸序列与鲤鱼的POMC-Ⅰ和POMC-Ⅱ具有最高的相似性,为92.4％,支持了稀有鮈鲫应属于鮈亚科的观点。系统进化树分析表明稀有鮈鲫的POMC序列属于POMCα。
运用RT-PCR和原位杂交技术分析了rmPOMC基因的垂体和垂体外组织表达情况。阳性细胞主要分布在RPD和PI,也有少数阳性细胞分布在PPD。垂体外组织,包括脑、小肠、性腺、肝胰脏、脾脏和鳃都检测到了rmPOMC mRNA的表达,表明在这些组织中,POMC存在旁分泌作用方式。
Rare minnow (Gobiocypris rarus), belonging to order of Cypriniformes, family of Cyprinidae, subfamily Danioninae, genus Gobiocypris, is a small freshwater fish endemic to China.
The structure of rare minnow pituitary was observed by histology and histochemistry methods. As a kidney, the pituitary of rare minnow was comprised of neurohypophysis and adenohypophysis, attaching to the hypothalamus by a pituitary stalk. Adenohypophysis was divided into three parts, RPD, PPD and PI according to the distinct histochemistrical characteristics. The curved branches of neurohypophysis were inserted in every part of adenohypophysis, and were most plentiful in PI. The blood vessels were rich in all parts of the pituitary.
cRNA probes were designed according to the cloned cDNAs of pituitary hormones. Data shows that:PRL positive cells were mainly distributed in RPD, and few were scattered in PPD. POMC positive cells located in most part of RPD and PI, with patterns of dispersed and compact along the PN branches, respectively. Inaddition, there were some POMC positive cells observed in PPD. GH positive cells were mainly grouped in PPD, few found in RPD, in a shape of round, oval or triangle. FSHβpositive cells located in RPD, PPD and few in PI, in a shape of round, oval or irregular. LHP positive cells distributed in PPD alone or in group, and few cells were also found in RPD and PI. There was a dividing line in PPD and PI, formed by LHβpositive cells. TSHPmainly grouped in the dorsal part of PPD, few scattered in RPD and PI. GTHapositive cells with a large distributing area, detected in every part of the pituitary, mostly in PPD, and formed a distinct limit between PPD and PI like the LHβ positive cells. There were few SL positive cells detected, and these cells were grouped or scattered only in posterior-ventral par of PI. The distributing patterns of hormone producing cells were different from each samples.
A cDNA encoding pro-opiomelanocortin (POMC) gene was cloned from the pituitary gland of the rare minnow (Gobiocypris rarus), a small freshwater fish endemic to China. This was achieved by reverse transcription polymerase chain reaction (RT-PCR) and rapid amplification of cDNA ends (RACE). Data showed that the predicted rare minnow POMC (rmPOMC) cDNA consisted of 846bps coding for the following sequences, flanked by proteolytic cleavage sites:signal peptide (SP, Met1-Ala28), N-terminal peptide (Gln29-His105), ACTH (Ser108-Met146),α-MSH (Ser108-Gal121), CLIP (Pro126-Met146),β-LPH (Glu149-His221),γ-LPH (Glul49-Ser186),β-MSH (Asp 170-Ser186), andβ-endorphin (β-EP, Tyr189-Gln221). Sequence analysis showed no region was homologous toγ-MSH (a tetrapod POMC feature). The amino acid sequence is highly similar to POMC-Ⅰand POMC-Ⅱof the common carp (92.4%), which supporting the viewpoint of rare minnow belongs to subfamily Gobioninae according to homologous alignment. It was POMCαthrough the phylogenetic analysis.
Pituitary and extra-pituitary expression were studied using RT-PCR and in situ hybridization. rmPOMC positive cells were mainly located in the rostral pars distalis (RPD) and pars intermedia (PI). Some rmPOMC positive cells were detected in the proximal pars distalis (PPD) as well, according to in situ hybridization. In the extra-pituitary tissues, positive signals were observed in the brain, intestines, gonads, hepatopancreas, spleen, and gills by RT-PCR analysis, showing that the paracrine pathway was existed in these tissues.
引文
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