淇河鲫bmp6基因的克隆及表达特性分析

doi:10.7668/hbnxb.20195439

华北农学报 ›› 2025, Vol. 40 ›› Issue (2): 218-229. doi: 10.7668/hbnxb.20195439

• 畜牧·水产·兽医 • 上一篇下一篇

淇河鲫bmp6基因的克隆及表达特性分析

马峻¹^,²^,³, 连凯琪¹^,³, 闪金研¹^,³, 刘玉玲¹^,³, 李晓龙¹^,³, 赵黎明⁴, 李学军²^,, 彭仁海¹^,³^,

¹ 安阳工学院生物与食品工程学院,河南安阳 455000
² 河南师范大学水产学院,河南新乡 453007
³ 河南省海河流域水生生物多样性野外科学观测研究站,河南林州 456576
⁴ 河南省水产技术推广站,河南郑州 450008

收稿日期:2024-09-19 出版日期:2025-05-12
通讯作者:
李学军(1967—),男,河南新乡人,教授,博士,博士生导师,主要从事水产动物遗传育种和水生生物资源保护与开发研究。
彭仁海(1972—),男,河南信阳人,教授,博士,博士生导师,主要从事植物功能基因挖掘与利用研究。
作者简介:
马峻(1999—),男,河南焦作人,在读硕士,主要从事水生动物育种研究。
马峻、连凯琪为同等贡献作者。
基金资助:
国家重点研发计划项目(2023YFD2400300); 河南省科技攻关项目(242102110065); 河南省大宗淡水鱼产业技术体系(HARS-22-16-G1); 安阳市科技攻关项目(2023C01NY013)

Cloning and Expression Analysis of bmp6 Gene in Carassius auratus var.Qihe

MA Jun¹^,²^,³, LIAN Kaiqi¹^,³, SHAN Jinyan¹^,³, LIU Yuling¹^,³, LI Xiaolong¹^,³, ZHAO Liming⁴, LI Xuejun²^,, PENG Renhai¹^,³^,

¹ School of Biological and Food Engineering,Anyang Institute of Technology,Anyang 455000,China
² College of Fisheries,Henan Normal University,Xinxiang 453007,China
³ Haihe River Basin Aquatic Biodiversity Observation and Research Station of Henan Province,Linzhou 456576,China
⁴ Henan Aquatic Technology Promotion Station,Zhengzhou 450008,China

Received:2024-09-19 Published:2025-05-12

摘要/Abstract

摘要：

骨形态发生蛋白6(bmp6)基因对鱼类肌间刺的形成发育具有重要的调控作用。为探究淇河鲫bmp6基因的序列特征和表达特性,通过克隆获得淇河鲫bmp6基因的编码序列(CDS),并进行生物信息学分析;通过实时荧光定量PCR(qRT-PCR)检测,分析bmp6基因在淇河鲫不同组织和发育时期的表达差异。结果显示,淇河鲫bmp6基因由两部分同源基因各3个等位基因组成,分别是bmp6a-1、bmp6a-2、bmp6a-3和bmp6b-1、bmp6b-2、bmp6b-3,其CDS序列全长分别为1 245,1 257 bp,编码414,418个氨基酸。bmp6a和bmp6b均是含有一个信号肽序列且无跨膜结构的不稳定疏水性分泌蛋白,bmp6a主要分布在线粒体、细胞核和细胞质中,而bmp6b主要分布在细胞核与线粒体上。bmp6a和bmp6b二级结构都以无规则卷曲为主,与三级结构预测的结果一致,且都具有1个TGF-β-rel家族保守结构域。氨基酸序列同源性分析表明,淇河鲫bmp6a氨基酸序列与团头鲂bmp6具有较高的相似性,与哺乳动物的相似性较低,而淇河鲫bmp6b氨基酸序列与鲫鱼和银鲫的bmp6b具有较高的相似性。系统发育树分析显示,淇河鲫bmp6a与鲤鱼bmp6a聚于同一进化支,而淇河鲫bmp6b与鲫鱼和银鲫bmp6b聚于同一进化支。qRT-PCR检测结果表明,bmp6a和bmp6b基因在淇河鲫的脑、肌肉、鳃、脾脏、肠、性腺、肝脏和肾脏组织中均有表达,但在肝脏、鳃组织和肾脏中表达量较高;二者在淇河鲫幼鱼不同发育时期具有相似的表达模式,在肌间刺骨化前,bmp6a和bmp6b的表达量最高,之后随着肌间刺的发育形成,bmp6a和bmp6b的表达量降低。综上,通过克隆获得淇河鲫bmp6基因CDS序列,并进行生物信息学分析和qRT-PCR检测,为进一步创制淇河鲫无肌间刺新品系提供了参考依据。

关键词: 淇河鲫, 骨形态发生蛋白6(bmp6), 基因克隆, 序列分析, 基因表达

Abstract:

Bone morphogenetic protein 6 gene(bmp6) plays an important role in regulating the formation and development of intermuscular bones in fish.In order to explore the sequence characteristics and expression characteristics of bmp6 gene in Carassius auratus var.Qihe,the coding sequence(CDS) of bmp6 gene was cloned and analyzed by bioinformatics.Through Quantitative Real-time PCR (qRT-PCR) detection,the expression differences of bmp6 gene in different tissues and developmental stages were identified.The results showed that the bmp6 gene of C.auratus var.Qihe was composed of two homologous genes,each with three alleles,namely bmp6a-1,bmp6a-2,bmp6a-3 and bmp6b-1,bmp6b-2,bmp6b-3,with full-length CDS of 1 245,1 257 bp,encoding 414 and 418 amino acids,respectively.Both bmp6a and bmp6b were unstable hydrophobic secretory proteins containing a signal peptide sequence and lacking a transmembrane structure.bmp6a was mainly distributed in mitochondria,nucleus and cytoplasm,while bmp6b was mainly distributed in nucleus and mitochondria.The secondary structure of bmp6a and bmp6b was mainly random coil,consistent with the results of tertiary structure prediction,and both contained one TGF-β-rel family conserved domain.The amino acid sequence homology analysis showed that the amino acid sequence of bmp6a had higher similarity with bmp6 of Megalobrama amblycephala,but lower similarity with mammals.The amino acid sequence of bmp6b had higher similarity with that of C.auratus and C.gibelio.Phylogenetic tree analysis showed that the bmp6a of C.auratus var.Qihe and the bmp6a of Cyprinus carpio were clustered in the same clade,while the bmp6b of C.auratus var.Qihe and the bmp6b of C.auratus and C.gibelio were clustered in the same clade.The results of qRT-PCR showed that bmp6a and bmp6b genes were expressed in brain,muscle,gill,spleen,intestine,ovary,liver and kidney tissues of C.auratus var.Qihe,but the expression levels were higher in liver,gill and kidney tissues.The expression patterns of bmp6a and bmp6b were similar in different developmental stages of juvenile C.auratus var.Qihe.The expression levels of bmp6a and bmp6b were the highest before the ossification of intermuscular spines,and then decreased with the development of intermuscular spines.To sum up,the CDS of bmp6 gene of C.auratus var.Qihe was obtained by cloning,and bioinformatics analysis and qRT-PCR detection were carried out,providing a reference for further creating a new strain of C.auratus var.Qihe without intermuscular bones.

Key words: Carassius auratus var.Qihe, Bone morphogenetic protein 6(bmp6), Gene cloning, Sequence analysis, Gene expression

马峻, 连凯琪, 闪金研, 刘玉玲, 李晓龙, 赵黎明, 李学军, 彭仁海. 淇河鲫bmp6基因的克隆及表达特性分析[J]. 华北农学报, 2025, 40(2): 218-229. doi: 10.7668/hbnxb.20195439.

MA Jun, LIAN Kaiqi, SHAN Jinyan, LIU Yuling, LI Xiaolong, ZHAO Liming, LI Xuejun, PENG Renhai. Cloning and Expression Analysis of bmp6 Gene in Carassius auratus var.Qihe[J]. Acta Agriculturae Boreali-Sinica, 2025, 40(2): 218-229. doi: 10.7668/hbnxb.20195439.

http://www.hbnxb.net/CN/Y2025/V40/I2/218

图/表 18

参考文献 31

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物种 Species	GenBank登录号 GenBank accession No.
物种 Species	bmp6a	bmp6b
鲤鱼Cyprinus carpio	BAX00833.1		BAX00834.1
多鳞白甲鱼Onychostoma macrolepis	XP_058621126.1		XP_058621127.1
鲫鱼Carassius auratus	XP_026056593.1		XP_026056594.1
黑鲫Carassius carassius	XP_059380348.1		XP_059380349.1
银鲫Carassius gibelio	XP_052452495.1		XP_052452496.1
露斯塔野鲮Labeo rohita	XP_050954083.1		XP_050954084.1
虎皮鱼Puntigrus tetrazona	XP_043082340.1		XP_043082341.1
滇池金线鲃Sinocyclocheilus grahami	XP_016086723.1		XP_016086724.1
斑马鱼Danio rerio	AAI65212.1
团头鲂 Megalobrama amblycephala	XP_048030213.1
草鱼Ctenopharyngodon idella	XP_051738970.1
波氏吻鰕虎鱼Rhinichthys klamathensis goyatoka	XP_056105783.1
玫瑰高原鳅Triplophysa rosa	XP_057178330.1
尼罗罗非鱼Oreochromis niloticus	XP_025765940.1
鼠Mus musculus	NP_031582.1
鸡Gallus gallus	XP_040520892.1
人Homo sapiens	AAI13019.1

物种 Species	GenBank登录号 GenBank accession No.
物种 Species	bmp6a	bmp6b
鲤鱼Cyprinus carpio	BAX00833.1		BAX00834.1
多鳞白甲鱼Onychostoma macrolepis	XP_058621126.1		XP_058621127.1
鲫鱼Carassius auratus	XP_026056593.1		XP_026056594.1
黑鲫Carassius carassius	XP_059380348.1		XP_059380349.1
银鲫Carassius gibelio	XP_052452495.1		XP_052452496.1
露斯塔野鲮Labeo rohita	XP_050954083.1		XP_050954084.1
虎皮鱼Puntigrus tetrazona	XP_043082340.1		XP_043082341.1
滇池金线鲃Sinocyclocheilus grahami	XP_016086723.1		XP_016086724.1
斑马鱼Danio rerio	AAI65212.1
团头鲂 Megalobrama amblycephala	XP_048030213.1
草鱼Ctenopharyngodon idella	XP_051738970.1
波氏吻鰕虎鱼Rhinichthys klamathensis goyatoka	XP_056105783.1
玫瑰高原鳅Triplophysa rosa	XP_057178330.1
尼罗罗非鱼Oreochromis niloticus	XP_025765940.1
鼠Mus musculus	NP_031582.1
鸡Gallus gallus	XP_040520892.1
人Homo sapiens	AAI13019.1

软件 Software	网址 Website	功能预测 Function prediction
BLAST	https://blast.ncbi.nlm.nih.gov/Blast.cgi	相似性比对
ProtParam	https://web.expasy.org/protparam/	理化性质
ProtScale	https://web.expasy.org/protscale/	亲疏水性
Pfam	http://pfam.xfam.org/	保守结构域
SignalP-4.1	https://services.healthtech.dtu.dk/services/SignalP-4.1/	信号肽分析
TMHMM 2.0	https://services.healthtech.dtu.dk/services/TMHMM-2.0/	跨膜结构域
SOPMA	https://npsa.lyon.inserm.fr/cgi-bin/secpred_sopma.pl	二级结构
SWISS-MODEL	https://swissmodel.expasy.org/interactive/	三级结构

软件 Software	网址 Website	功能预测 Function prediction
BLAST	https://blast.ncbi.nlm.nih.gov/Blast.cgi	相似性比对
ProtParam	https://web.expasy.org/protparam/	理化性质
ProtScale	https://web.expasy.org/protscale/	亲疏水性
Pfam	http://pfam.xfam.org/	保守结构域
SignalP-4.1	https://services.healthtech.dtu.dk/services/SignalP-4.1/	信号肽分析
TMHMM 2.0	https://services.healthtech.dtu.dk/services/TMHMM-2.0/	跨膜结构域
SOPMA	https://npsa.lyon.inserm.fr/cgi-bin/secpred_sopma.pl	二级结构
SWISS-MODEL	https://swissmodel.expasy.org/interactive/	三级结构

理化性质 Physical and chemical properties	bmp6a	bmp6b
分子质量/ku Molecular weight	47.543	47.666
等电点 pI	7.22	6.36
正电荷残基数/个Positively charged residues	46	46
负电荷残基数/个Negatively charged residues	46	51
不稳定指数Instability index	49.08	54.69
脂肪系数Aliphatic index	77.95	75.84
稳定性Stability	不稳定	不稳定

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淇河鲫bmp6基因的克隆及表达特性分析

Cloning and Expression Analysis of bmp6 Gene in Carassius auratus var.Qihe

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基因 Gene	引物序列(5'—3') Primer sequence(5'—3')	产物长度/bp Product size	退火温度/℃ Annealing temperature	用途 Application
bmp6a	F:ATGACAAGCGTCTGGTTTGC R:TCAGTGGCAGCCGCATGCGC	1 245	64	克隆
bmp6b	F:ATGACAAGCGTCTGGTTTGC R:TCAGTGGCAGCCACATGCACG	1 257	64	克隆
bmp6a	Q-F:GCACCCGAGGGCTATGCAGC Q-R:GAGCACGGAGATTGCATGCAG	174	60	qRT-PCR
bmp6b	Q-F:CTTCACTAGAGTTTCTGCATGAC Q-R:TGGAATCTGGGACAGGTTGA	128	60	qRT-PCR
β-actin	F:TGGAAGGTGGACAGGGAAGC R:ACCATCTACCCCGGTATTGC	149	60	qRT-PCR

bmp6a			bmp6b
氨基酸 Amino acid	数量/个 Number	比例/% Percentage	氨基酸 Amino acid	数量/个 Number	比例/% Percentage
Ala (A)	26	6.3	Ala (A)	31	7.4
Arg (R)	30	7.2	Arg (R)	29	6.9
Asn (N)	27	6.5	Asn (N)	23	5.5
Asp (D)	17	4.1	Asp (D)	19	4.5
Cys (C)	9	2.2	Cys (C)	8	1.9
Gln (Q)	16	3.9	Gln (Q)	15	3.6
Glu (E)	29	7.0	Glu (E)	32	7.7
Gly (G)	19	4.6	Gly (G)	19	4.5
His (H)	14	3.4	His (H)	16	3.8
Ile (I)	14	3.4	Ile (I)	14	3.3
Leu (L)	42	10.1	Leu (L)	40	9.6
Lys (K)	16	3.9	Lys (K)	17	4.1
Met (M)	10	2.4	Met (M)	12	2.9
Phe (F)	19	4.6	Phe (F)	21	5.0
Pro (P)	19	4.6	Pro (P)	19	4.5
Ser (S)	35	8.5	Ser (S)	40	9.6
Thr (T)	22	5.3	Thr (T)	18	4.3
Trp (W)	8	1.9	Trp (W)	8	1.9
Tyr (Y)	15	3.6	Tyr (Y)	11	2.6
Val (V)	27	6.5	Val (V)	26	6.2

bmp6a亚细胞定位 Subcellular localization of bmp6a	可能性/% Probability	bmp6b亚细胞定位 Subcellular localization of bmp6b	可能性/% Probability
线粒体Mitochondrial	30.4	线粒体Mitochondrial	26.1
细胞核Nuclear	17.4	细胞核Nuclear	52.2
细胞质Cytoplasmic	17.4	细胞质Cytoplasmic	4.3
细胞外Extracellular	13.0	细胞外Extracellular	17.4
内质网 Endoplasmic reticulum	13.0
囊泡Vesicles	8.7

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淇河鲫bmp6基因的克隆及表达特性分析

Cloning and Expression Analysis of bmp6 Gene in Carassius auratus var.Qihe

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