西藏桑桑牦牛<i>RAN</i>基因克隆、生物信息学及组织表达分析

doi:10.7668/hbnxb.20196146

华北农学报 ›› 2026, Vol. 41 ›› Issue (1): 212-221. doi: 10.7668/hbnxb.20196146

所属专题： 畜牧；热点论文；

• 畜牧·水产·兽医 • 上一篇下一篇

西藏桑桑牦牛RAN基因克隆、生物信息学及组织表达分析

刘毅程¹^,², 洛桑顿珠³, 卓玛次仁⁴, 尼玛加措⁴, 平措占堆³, 马晓明²^,⁵, 梁春年¹^,⁵, 李少斌¹

¹ 甘肃农业大学动物科学技术学院,甘肃兰州 730050
² 中国农业科学院兰州畜牧与兽药研究所,甘肃省牦牛繁育工程重点实验室,甘肃兰州 730050
³ 西藏自治区农牧科学院畜牧兽医研究所,西藏拉萨 850004
⁴ 日喀则市昂仁县农业农村局,西藏日喀则 857001
⁵ 农业农村部青藏高原畜禽遗传育种重点实验室,甘肃兰州 730050

收稿日期:2025-05-19 出版日期:2026-02-28
通讯作者:
李少斌(1983—),男,山西汾阳人,研究员,博士,博士生导师,主要从事现代生物技术与动物生产和适应性进化研究。
梁春年(1973—),男,甘肃武威人,研究员,博士,博士生导师,主要从事动物遗传育种与繁殖研究。
作者简介:
刘毅程(2002—),男,甘肃定西人,硕士,主要从事动物遗传育种与繁殖研究。
基金资助:
西藏自治区区域科技协同创新专项(QYATZX-RKZ2022-03); 西藏自治区重点研发计划项目(XZ202402ZY0012); 现代肉牛牦牛产业技术体系(CARS-37)

Cloning,Bioinformatics and Tissue Expression Analysis of RAN Gene from Sangsang Yak in Tibet

LIU Yicheng¹^,², Lobsang Dunzhu³, Zhuoma Tsering⁴, Nima Gyatso⁴, Phuntsok Zhandui³, MA Xiaoming²^,⁵, LIANG Chunnian¹^,⁵, LI Shaobin¹

¹ College of Animal Science and Technology,Gansu Agricultural University,Lanzhou 730050,China
² Lanzhou Institute of Husbandry and Pharmaceutica Sciences,Chinese Academy of Agricultural Sciences, Gansu Key Laboratory of Yak Breeding Engineering,Lanzhou 730050,China
³ Institute of Animal Husbandry and Veterinary,Tibet Autonomous Regional Academy of Agricultural Sciences,Lhasa 850004,China
⁴ Agriculture and Rural Bureau of Ngamring County,Xigaze City, Xigaze 857001,China
⁵ Key Laboratory of Livestock and Poultry Genetics and Breeding, Qinghai-Tibet Plateau,Ministry of Agriculture and Rural Affairs,Lanzhou 730050,China

Received:2025-05-19 Published:2026-02-28

摘要/Abstract

摘要：

为了探究牦牛Ras相关核蛋白基因(RAN)的结构及生物学功能,解析该基因如何调控牦牛体内细胞增殖和参与蛋白质的合成,以桑桑牦牛肾脏组织cDNA为模板使用RT-PCR技术克隆桑桑牦牛RAN基因CDS区序列,使用多种软件及在线工具对其进行生物信息学分析,并使用qPCR技术检测RAN基因在桑桑牦牛7个组织中的表达情况。结果表明,桑桑牦牛RAN基因CDS区全长651 bp,编码216个氨基酸;通过同源性比对,发现桑桑牦牛与野牦牛和瘤牛之间的亲缘关系最近,相似度达99.2%,与鸡的最远,为86.6%;RAN蛋白分析预测结果显示,该蛋白分子质量24.423 11 ku,理论等电点为7.01,其原子总数为3 449,分子组成为C₁₁₀₉H₁₇₂₅N₂₉₅O₃₁₃S₇。RAN蛋白不存在跨膜结构且无N-糖基化潜在位点,其存在36个磷酸化位点。预测其亲、疏水性及计算不稳定系数,发现该蛋白为稳定型亲水性蛋白。根据亚细胞定位发现,该蛋白存在于牦牛细胞中的高尔基体、线粒体、细胞核和细胞质中;通过预测RAN蛋白结构发现,其高级结构以α-螺旋为主,不包含β-转角。蛋白互作网络结果显示,桑桑牦牛RAN蛋白与RAN结合蛋白1(RANBP1)、RAN结合蛋白2(RANBP2)、RANGTPase激活蛋白1(RANGAP1)等蛋白存在互作关系,其相互间也存在互作关系。RT-qPCR结果显示,桑桑牦牛睾丸组织中RAN基因表达量显著高于其他组织,而在肌肉组织中未检测到表达。综上成功克隆了RAN基因CDS区,并完成其生物信息学分析,同时还研究了该基因在桑桑牦牛组织中的表达情况,发现其在生殖系统的发育、细胞增殖、疾病的预防与控制和参与蛋白质合成等方面发挥重要作用。

关键词: 桑桑牦牛, RAN基因, 基因克隆, 生物信息学分析, 表达

Abstract:

In order to explore the structure and biological function of Ras related nuclear protein gene(RAN)in yak and resolve how it regulates the proliferation of yak cells and participates in protein synthesis,the CDS region sequence of RAN gene of Sangsang yak was cloned by RT-PCR using Sangsang yak kidney tissue cDNA as template,and bioinformatics analysis was carried out by a variety of software and online tools,the expression of the RAN gene in seven tissues of Sangsang yaks was detected by qPCR technology.The results showed that the CDS region of RAN gene in Sangsang yak was 651 bp in length,encoding 216 amino acids.Through homology comparison,it was found that the genetic relationship between Sangsang yak and wild yak and zebu was the closest,with a similarity of 99.2%,and the farthest from chicken,reaching 86.6%.The prediction results of RAN protein analysis showed that the molecular weight of the protein was 24.423 11 ku,the theoretical isoelectric point was 7.01,the total number of atoms was 3 449,and the molecular composition was C₁₁₀₉H₁₇₂₅N₂₉₅O₃₁₃S₇.RAN protein had no transmembrane structure and no potential sites for N-glycosylation,with 36 phosphorylation sites.The affinity and hydrophobicity were predicted and the instability coefficient was calculated,and it was found that the protein was a stable hydrophilic protein.According to the subcellular localization,the protein was found to be present in the Golgi apparatus,mitochondria,nucleus and cytoplasm in yak cells.Predictions of the RAN protein structure revealed that its higher-order structure consisted mainly of α-helices and did not contain β-turns.Protein interaction network results showed that there was an interaction between the RAN protein and RAN Binding Protein 1(RANBP1),RAN Binding Protein 2(RANBP2),RANGTPase activating protein 1(RANGAP1)and other proteins of Sangsang yak,and there was also an interaction between them.The expression level of the RAN gene in yak testicular tissue was significantly higher than in other tissues,while no expression was detected in muscle tissue.It successfully cloned the CDS region of the RAN gene and completed its bioinformatics analysis,and the expression of this gene in the tissue of Sangsang yak was also studied,and it was found that it played an important role in the development of the reproductive system,cell proliferation,disease prevention and control,and participation in protein synthesis.

Key words: Sangsang yak, RAN gene, Gene cloning, Bioinformatics analysis, Expression

中图分类号:

刘毅程, 洛桑顿珠, 卓玛次仁, 尼玛加措, 平措占堆, 马晓明, 梁春年, 李少斌. 西藏桑桑牦牛RAN基因克隆、生物信息学及组织表达分析[J]. 华北农学报, 2026, 41(1): 212-221. doi: 10.7668/hbnxb.20196146.

LIU Yicheng, Lobsang Dunzhu, Zhuoma Tsering, Nima Gyatso, Phuntsok Zhandui, MA Xiaoming, LIANG Chunnian, LI Shaobin. Cloning,Bioinformatics and Tissue Expression Analysis of RAN Gene from Sangsang Yak in Tibet[J]. Acta Agriculturae Boreali-Sinica, 2026, 41(1): 212-221. doi: 10.7668/hbnxb.20196146.

http://www.hbnxb.net/CN/Y2026/V41/I1/212

图/表 17

表1 RAN基因引物序列信息

Tab.1 Primer sequence information for RAN gene

基因 Genes	引物序列(5'—3') Primer sequences(5'—3')	长度/bp Length	退火温度/℃ Annealing temperature	用途 Application
RAN	F: TTGCGCTTCCGCCATCTTT	875	59.5	PCR
	R: CACTCCAAAGCCCATCCATC
RAN	F: GAAAAGCCCTTCCTCTGGCT	125	59.0	RT-qPCR
	R: TGCTCATACTGTGCTGCCAA
GAPDH	F: GGCATCGTGGAGGGACTTATG	186	59.0	RT-qPCR
	R: CCCAGTGAGCTTCCCGTTGAG

表2 桑桑牦牛RAN蛋白生物信息学分析软件

Tab.2 Sangsang yak RAN protein bioinformatics analysis software

软件 Software	网站 Website	功能 Function
ORF Finder	https://www. ncbi. nlm. nih.gov/orffinder/	开放阅读框分析
BLAST	https://blast.ncbi.nlm.nih.gov/Blast.cgi	同源性比对分析
ProtParam	https://web.expasy.org/protparam	蛋白质理化性质分析
ProtScale	https://web.expasy.org/protscale	蛋白质亲疏水性分析
SignalP 5.0	https: //services.healthtech.dtu.dk/services/SignalP-4.1/	蛋白质信号肽分析
TMHMM 2.0	https://services.healthtech.dtu.dk/service.php?TMHMM-2.0	蛋白跨膜结构预测
NetPhos 3.1	https://services. healthtech.dtu.dk/service.php?NetPhos-3.1	蛋白磷酸化位点预测
NetNGlyc	https://services.healthtech.dtu.dk/service.php?NetNGlyc-1.0	蛋白糖基化位点预测
STRING 11.5	https://string-db.org	蛋白互作网络预测
SOPMA	https://npsa-pbil.ibcp.fr/cgi-bin/npsa_automat.pl?page=npsa_sopma.html	蛋白质二级结构预测
SWISS-MODEL	https://swissmodel. expasy. org/interactive	蛋白质三级结构预测
PSORT Ⅱ	https://www. genscript. com/psort. html	蛋白亚细胞定位预测

图1 桑桑牦牛RAN基因PCR扩增结果 1—3.桑桑牦牛RAN基因PCR扩增的产物;M.DL2000 DNA Marker。

Fig.1 PCR amplification results of RAN gene in Sangsang yak 1—3.PCR amplification products of RAN gene of Sangsang yak;M.DL2000 DNA Marker.

表3 桑桑牦牛RAN基因的开放阅读框

Tab.3 Open reading frame of the RAN gene of Sangsang yak

标签 Label	链 Strand	阅读框 Frame	起始位置/bp Start	终止位置/bp Stop	长度(核苷酸 \| 氨基酸) Length(bp\|aa)
ORF3	+	3	39	689	651 \| 216
ORF1	+	1	337	543	207 \| 68
ORF4	-	1	197	>3	195 \| 64
ORF5	-	2	604	500	105 \| 34
ORF2	+	2	227	310	84 \| 27

图2 桑桑牦牛RAN基因的氨基酸序列

Fig.2 Amino acid sequence of RAN gene in Sangsang yak

图3 11种物种RAN核苷酸序列同源性比对

Fig.3 Sequence homology comparison of RAN nucleotides in 11 species

图4 不同物种RAN基因系统进化树

Fig.4 Phylogenetic tree of RAN gene in different species

表4 桑桑牦牛RAN蛋白氨基酸组成成分

Tab.4 Amino acid composition of RAN protein from Sangsang yak

氨基酸种类 Amino acid	数量/个 Amount	含量/% Content	氨基酸种类 Amino acid	数量/个 Amount	含量/% Content
丙氨酸Ala	17	7.9	亮氨酸Leu	18	8.3
精氨酸Arg	9	4.2	赖氨酸Lys	18	8.3
天冬酰胺Asn	10	4.6	甲硫氨酸Met	4	1.9
天冬氨酸Asp	15	6.9	苯丙氨酸Phe	11	5.1
半胱氨酸Cys	3	1.4	脯氨酸Pro	12	5.6
谷氨酰胺Gln	9	4.2	丝氨酸Ser	4	1.9
谷氨酸Glu	12	5.6	苏氨酸Thr	11	5.1
甘氨酸Gly	14	6.5	色氨酸Trp	3	1.4
组氨酸His	6	2.8	苏氨酸Tyr	8	3.7
异亮氨酸Ile	10	4.6	缬氨酸Val	22	10.2

图5 蛋白亲、疏水性预测

Fig.5 Prediction of protein affinity and hydrophobicity

图6 蛋白跨膜结构预测

Fig.6 Prediction of transmembrane structure

图7 蛋白信号肽预测

Fig.7 Protein signal peptide prediction

图8 蛋白潜在磷酸化位点预测

Fig.8 Prediction of potential phosphate sites

图9 蛋白糖基化位点预测

Fig.9 Protein glycosylation site prediction

图10 蛋白二级结构预测 A.蛋白二级结构位点图:h.α-螺旋,c.无规则卷曲,e.延伸链;B.蛋白质二级结构峰图;C.蛋白质二级结构竖条纹图。

Fig.10 Protein secondary structure prediction A.Protein secondary structure site diagram:h.α-helix,c.Random coil,e.Extended strand;B.Protein secondary structure peak diagram;C.Protein secondary structure vertical fringe diagram.

图11 蛋白三级结构预测

Fig.11 Protein tertiary structure prediction

图12 蛋白互作网络分析图

Fig.12 Protein interaction network analysis diagram

图13 RAN基因在桑桑牦牛不同组织中mRNA 相对表达量不同小写字母表示各组织间差异显著(P<0.05)。

Fig.13 Relative mRNA expression levels of RAN gene in different tissues of Sangsang yak Different lowercase letters indicated significant differences among tissues(P<0.05).

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[23]	张娟香, 马小勇, 路建卫, 仁青吉, 罗胜伟, 肖光峰, 王有鹤, 马晓明, 喇永福, 郭宪, 褚敏, 阎萍, 梁春年. 牦牛FTL基因克隆、生物信息学及表达分析[J]. 华北农学报, 2023, 38(6):192-200.doi:10.7668/hbnxb.20194239.
	Zhang J X, Ma X Y, Lu J W, Ren Q J, Luo S W, Xiao G F, Wang Y H, Ma X M, La Y F, Guo X, Chu M, Yan P, Liang C N. Cloning,bioinformatics analysis and expression analysis of FTL gene in yak[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(6):192-200.
[24]	杨明. 应用XL-MS研究Ran、RanBP1和RCC1蛋白间的相互作用界面[D]. 呼和浩特: 内蒙古大学, 2023.doi:10.27224/d.cnki.gnmdu.2023.000523.
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[25]	梁海霞. Ran GTPase在嗜热四膜虫大核无丝分裂和程序性核死亡过程的功能研究[D]. 太原: 山西大学, 2013.
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[26]	Zhu C, Zhu J R, Duan Q Y, Jiang Y, Yin H, He Y L, Li F, An X P. Exploration of the lactation function of protein phosphorylation sites in goat mammary tissues by phosphoproteome analysis[J]. BMC Genomics, 2021, 22(1):703.doi:10.1186/s12864-021-07993-5. pmid: 34583635
[27]	Choudhury S, Mansi, Muthusamy S K, Padaria J C, Dalal M. Genome-wide identification of Ran GTPase family genes from wheat(T.aestivum)and their expression profile during developmental stages and abiotic stress conditions[J]. Functional & Integrative Genomics, 2021, 21(2):239-250.doi:10.1007/s10142-021-00773-0.
[28]	任宣奇. RCC1协同PALB2通过组蛋白乙酰化调控DNA损伤修复分子机制的研究[D]. 上海: 上海师范大学, 2021.doi:10.27312/d.cnki.gshsu.2021.000513.
	Ren X Q. Study on the molecular mechanism of RCC1 and PALB2 regulating DNA damage repair through histone acetylation[D]. Shanghai: Shanghai Normal University, 2021.
[29]	张晓娥. XPO1共价抑制剂的虚拟筛选及皮肤损伤治疗的研究[D]. 大连: 大连理工大学, 2015.
	Zhang X E. Virtual screening of XPO1 covalent inhibitors and study on treatment of skin injury[D]. Dalian: Dalian University of Technology, 2015.
[30]	Liu X M, Liu J Y, Xiao W, Zeng Q H, Bo H, Zhu Y X, Gong L, He D, Xing X W, Li R H, Zhou M, Xiong W, Zhou Y H, Zhou J D, Li X H, Guo F, Xu C X, Chen X, Wang X Y, Wang F, Wang Q, Cao K. SIRT1 regulates N6-methyladenosine RNA modification in hepatocarcinogenesis by inducing RANBP2-dependent FTO SUMOylation[J]. Hepatology, 2020, 72(6):2029-2050.doi:10.1002/hep.31222. URL
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[37]	王纪良. 湖羊生长相关基因ATF4、RAC1和RAN的分子鉴定、组织表达及多态性分析[D]. 昆明: 云南农业大学, 2023.doi:10.27458/d.cnki.gynyu.2023.000331.
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西藏桑桑牦牛RAN基因克隆、生物信息学及组织表达分析

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