美仁牦牛<i>PDK4</i>基因克隆及组织表达

doi:10.7668/hbnxb.20193924

华北农学报 ›› 2023, Vol. 38 ›› Issue (5): 197-205. doi: 10.7668/hbnxb.20193924

所属专题： 畜牧；

• 畜牧·水产·兽医 • 上一篇下一篇

美仁牦牛PDK4基因克隆及组织表达

马小勇¹^,², 戴荣凤¹^,², 李歆怡¹^,², 杨国武, 扎老³, 路建卫³, 赵雪⁴, 梁春年¹^,²

¹ 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
² 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
³ 合作市佐盖多玛乡畜牧站, 甘肃合作 747000
⁴ 合作市农畜产品质量安全检测检验中心,甘肃合作 747000

收稿日期:2022-12-14 出版日期:2023-10-28
通讯作者:
梁春年(1973-),男,甘肃武威人,研究员,博士,主要从事动物遗传育种与繁殖研究。
作者简介:
马小勇(1999-),男,甘肃清水人,在读硕士,主要从事动物遗传育种与繁殖研究。
基金资助:
国家重点研发计划(2021YFD1600200); 现代肉牛牦牛产业技术体系(CARS-37); 甘肃省基础研究创新群体项目(20JR5RA580); 中国农业科学院创新工程(25-LIHPS-01); 甘肃省科技重大专项(21ZD10NA001); 甘肃省科技重大专项(GZGG-2021-1)

Cloning and Tissue Expression Analysis of PDK4 Gene in Meiren Yak

MA Xiaoyong¹^,², DAI Rongfeng¹^,², LI Xinyi¹^,², YANG Guowu, ZHA Lao³, LU Jianwei³, ZHAO Xue⁴, LIANG Chunnian¹^,²

¹ Lanzhou Institute of Husbandry and Pharmaceutical Sciences of Chinese Academy of Agricultural Sciences, Key Laboratory of Yak Breeding Engineering of Gansu Province,Lanzhou 730050,China
² Key Laboratory of Animal Genetics and Breeding the Tibetan Plateau,Ministry of Agriculture and Rural Affairs,Lanzhou 730050, China
³ Animal Husbandry Station,Zuogaiduoma Township of Hezuo City,Hezuo 747000,China
⁴ Agricultural and Livestock Products Quality and Safety Inspection Center of Hezuo City,Hezuo 747000,China

Received:2022-12-14 Published:2023-10-28

摘要/Abstract

摘要：

为研究美仁牦牛PDK4基因在各组织中的表达特征,探索其与脂肪沉积机制的相关性,采集了美仁牦牛背最长肌、心脏、肝脏、脾脏、肺脏、肾脏、脂肪和睾丸等组织样,并提取了其总RNA,通过RT-PCR、生物信息学软件及qPCR等技术对美仁牦牛PDK4基因进行了扩增、生信分析和表达特征研究。结果表明,美仁牦牛PDK4基因CDS区全长1 224 bp,编码407个氨基酸,蛋白相对分子质量为46.159 14 ku,理论等电点6.60,总平均亲水性-0.169,为亲水性蛋白;系统进化树结果表明,美仁牦牛与野牦牛亲缘关系最近,与鬣狗亲缘关系最远;PDK4蛋白有34处磷酸化位点,无信号肽和跨膜区,主要在线粒体中发挥生物学功能;美仁牦牛PDK4蛋白主要是由α-螺旋和无规则卷曲组成。PDK4基因在7日龄美仁牦牛肝脏和肺脏中的表达量显著高于其他组织(P<0.05),在肌肉、脂肪、脾脏和睾丸中中度表达,在肾和心脏中低表达。随着牦牛年龄增长,PDK4基因在30月龄美仁牦牛脂肪组织中的表达量增加,并显著高于7日龄美仁牦牛脂肪中的表达量(P<0.05)。以上结果表明,PDK4基因可能在美仁牦牛的生长过程中参与脂肪调控,在脂肪沉积过程中发挥重要作用。

关键词: 美仁牦牛, PDK4基因, 克隆, 实时荧光定量PCR

Abstract:

To investigate the expression characteristics of the PDK4 gene in various tissues of Meiren yaks and to explore its correlation with the mechanism of fat deposition,tissue samples were collected from the longest and dorsal muscles,heart,liver,spleen,lung,kidney,fat and testis of Meiren yaks,and total RNA was extracted.The gene was amplified,analyzed and characterized by RT-PCR,bioinformatics software,and qPCR.The results showed that the CDS region of the PDK4 gene was 1 224 bp,407 amino acids,the relative molecular mass of protein was 46.159 14 ku,the theoretical isoelectric point was 6.60,and the total average hydrophilicity was-0.169,which was a hydrophilic protein;the phylogenetic tree results showed that Meiren yak was the closest related to wild yak and the most distant related to hyena;PDK4 protein had 34 phosphorylation sites,no signal peptide and transmembrane region,and mainly plays a biological function in mitochondria;the PDK4 protein of Meiren yak was mainly composed of α-helix and irregular curling.The expression of the PDK4 gene in the liver and lung of 7-day-old Meiren yak was significantly higher than that of other tissues (P<0.05),moderately expressed in muscle,fat,spleen,and testes,and low in the kidney and heart.As the yak aged,the expression of the PDK4 gene in the adipose tissue of 30 months old Meiren yak increased and was significantly higher than that in the fat of 7-day-old Meiren yak(P<0.05).The above results suggest that the PDK4 gene may be involved in fat regulation during the growth process of Meiren yak and play an essential role in the fat deposition process.

Key words: Meiren yak, Pyruvate dehydrogenase kinase 4 gene(PDK4), Cloning, Real-time Fluorescent Quantitative PCR

马小勇, 戴荣凤, 李歆怡, 杨国武, 扎老, 路建卫, 赵雪, 梁春年. 美仁牦牛PDK4基因克隆及组织表达[J]. 华北农学报, 2023, 38(5): 197-205. doi: 10.7668/hbnxb.20193924.

MA Xiaoyong, DAI Rongfeng, LI Xinyi, YANG Guowu, ZHA Lao, LU Jianwei, ZHAO Xue, LIANG Chunnian. Cloning and Tissue Expression Analysis of PDK4 Gene in Meiren Yak[J]. Acta Agriculturae Boreali-Sinica, 2023, 38(5): 197-205. doi: 10.7668/hbnxb.20193924.

http://www.hbnxb.net/CN/Y2023/V38/I5/197

图/表 10

表1 引物序列

Tab.1 Primer sequences

基因 Gene	用途 Purpose	序列(5'-3') Sequence(5'-3')		退火温度/℃ Annealing temperature	产物长度/bp Product length
PDK4	基因克隆	F1	GAGGTGACAGGACACGAACTT	59.0	1 509
		R1	CTGGCACAGACCCACTTTGA
	qPCR	F2	GGCTGCTTCCTGAACGTTTG	58.0	116
		R2	TCCGGTCACATCAGTGTTCG
GAPDH	qPCR	F3	TTGTGATGGGCGTGAACC	58.0	169
		R3	GTCTTCTGGGTGGCAGTGAT

表2 生信分析软件

Tab.2 Bioinformatics analysis software

软件 Software	用途 Application
Blast	序列对比验证
Mega 10.1	构建系统进化树
Open reading frame finder(OFR Finder, http://www.ncbi.nlm.nih.gov/gorf/orfig.cgi)	查看开放阅读框
ProtParam(http://www.expasy.ch/tools/Protparam.html)	蛋白质理化性质分析
ProtScale(https://web.expasy.org/protscale/)	蛋白亲疏水性分析
SignalP 4.1(http://www.cbs.dtu.dk/services/SignalP/)	信号肽预测
TMHMM(http://www.cbs.dtu.dk/services/TMHMM/)	跨膜结构域分析
Psort Ⅱ(https://wolfpsort.hgc.jp/)	蛋白亚细胞定位
NetPhos 3.1 Server(http://www.cbs.dtu.dk/services/NetPhos/)	磷酸化位点预测
Sopma(https://npsa-prabi.ibcp.fr/cgi-bin/npsa_automat.pl?page=npsa_sopma.html)	蛋白质二级结构预测
SWISS-MODEL(http://www.expasy.org/swissmod/SWISS-MODEL.html)	蛋白质三级结构预测
STRING(https://string-db.org/)	蛋白互作网络预测

图1 PDK4基因扩增电泳图 1.PDK4基因扩增产物;M.DL5000 DNA Marker。

Fig.1 Amplified electrophoresis of PDK4 gene 1.Amplification product of PDK4 gene;M.DL5000 DNA Marker.

图2 美仁牦牛与野牦牛PDK4基因碱基序列对比结果

Fig.2 Comparison results of PDK4 gene base sequences in Meiren yak and Bos mutus

图3 不同物种间PDK4氨基酸系统进化树

Fig.3 Phylogenetic tree of PDK4 amino acids among different species

表3 PDK4蛋白中氨基酸含量

Tab.3 Amino acid composition of the PDK4 protein

氨基酸 Amino acid	数量/个 Quantity	含量/% Content	氨基酸 Amino acid	数量/个 Quantity	含量/% Content
丙氨酸 Ala(A)	25	6.10	亮氨酸 Leu(L)	44	10.80
精氨酸 Arg(R)	19	4.70	赖氨酸 Lys(K)	22	5.40
天冬酰胺 Asn(N)	19	4.70	蛋氨酸 Met(M)	13	3.20
天冬氨酸 Asp(D)	21	5.20	苯丙氨酸Phe(F)	19	4.70
半胱氨酸 Cys(C)	6	1.50	脯氨酸 Pro(P)	25	6.10
谷氨酰胺 Gln(Q)	18	4.40	丝氨酸 Ser(S)	33	8.10
谷氨酸 Glu(E)	22	5.40	苏氨酸 Thr(T)	19	4.70
甘氨酸 Gly(G)	18	4.40	色氨酸 Trp(W)	3	0.70
组氨酸 His(H)	11	2.70	酪氨酸 Tyr(Y)	18	4.40
异亮氨酸 Ile(I)	24	5.90	缬氨酸 Val(V)	28	6.90

图4 PDK4蛋白亲/疏水性分析

Fig.4 Hydrophilic and hydrophobic analysis of the PDK4 protein

图5 美仁牦牛PDK4 蛋白信号肽、跨膜结构域及磷酸化位点分析 A.牦牛PDK4 蛋白信号肽预测;B.牦牛PDK4跨膜结构域预测;C.PDK4磷酸化位点分析。

Fig.5 Analysis of signal peptide,transmembrane domain and phosphorylation site of PDK4 protein A.Prediction of yak PDK4 protein signal peptide;B.Prediction of transmembrane domain of PDK4;C.PDK4 phosphorylation site analysis.

图6 美仁牦牛PDK4蛋白二、三级结构及互作网络预测 A.二级结构预测;B.三级结构;C.蛋白互作网络图。

Fig.6 Prediction of the secondary and tertiary structure and interaction network of PDK4 protein in Meiren yak A.The secondary structure prediction;B.The tertiary structure prediction;C.Diagram of protein interaction network.

图7 PDK4基因在各组织中的mRNA表达水平 A.PDK4基因在30月龄美仁牦牛各组织中的相对表达量;B.PDK4基因在7日龄美仁牦牛各组织中的相对表达量;C.30月龄与7日龄美仁牦牛各组织间相对表达量的比较;图 A、B 中,不同小写字母表示差异显著(P<0.05);图 C 中,标记不同大写字母表示组内差异显著(P<0.05)。

Fig.7 mRNA expression levels of PDK4 gene in various tissues A.Relative expression of PDK4 gene in various tissues of 30 months old Meiren yak; B.Relative expression of PDK4 gene in each tissue of 7-day-old Meiren yak; C.Comparison of relative expression levels between tissues of 30 months old and 7-day-old Meiren yak,In Fig.A and B,different lowercase letters indicate significant differences (P<0.05) and in Fig.C,Marking different capital letters indicate significant differences within the group (P<0.05).

参考文献 32

[1]	Li Z X, Jiang M F. Metabolomic profiles in yak mammary gland tissue during the lactation cycle[J]. PLoS One, 2019, 14(7):e0219220.doi:10.1371/journal.pone.0219220. URL
[2]	陈美, 柴志欣, 武志娟, 王会, 王吉坤, 王嘉博, 钟金城, 信金伟. 牦牛ACAA2基因克隆及组织表达分析[J]. 东北农业大学学报, 2020, 51(12):51-57.doi:10.3969/j.issn.1005-9369.2020.12.006.
	Chen M, Chai Z X, Wu Z J, Wang H, Wang J K, Wang J B, Zhong J C, Xin J W. Cloning and tissue expression analysis of ACAA2 gene in yak[J]. Journal of Northeast Agricultural University, 2020, 51(12):51-57.
[3]	李丹, 王守伟, 臧明伍, 张凯华, 张哲奇, 赵金杨. 我国肉类食品安全风险现状与对策[J]. 肉类研究, 2015, 29(11):34-38.doi:10.15922/j.cnki.rlyj.2015.11.008.
	Li D, Wang S W, Zang M W, Zhang K H, Zhang Z Q, Zhao J Y. Current status and countermeasures of meat safety and risk in China[J]. Meat Research, 2015, 29(11):34-38.
[4]	刘晓畅, 张寿, 孙宝忠, 谢鹏, 徐晨晨, 雷元华, 张松山, 参木友, 鲍宇红. 牦牛肉品质特性研究进展[J]. 肉类研究, 2020, 34(11):78-83.doi:10.7506/rlyj1001-8123-20200917-230.
	Liu X C, Zhang S, Sun B Z, Xie P, Xu C C, Lei Y H, Zhang S S, Shen M Y, Bao Y H. Progress in understanding quality characteristics of yak meat[J]. Meat Research, 2020, 34(11):78-83.
[5]	Holness M J, Sugden M C. Regulation of pyruvate dehydrogenase complex activity by reversible phosphorylation[J]. Biochemical Society Transactions, 2003, 31(6):1143-1151.doi:10.1042/bst0311143. URL
[6]	Michelakis E D, Gurtu V, Webster L, Barnes G, Watson G, Howard L, Cupitt J, Paterson I, Thompson R B, Chow K, O'Regan D P, Zhao L, Wharton J, Kiely D G, Kinnaird A, Boukouris A E, White C, Nagendran J, Freed D H, Wort S J, Gibbs J S R, Wilkins M R. Inhibition of pyruvate dehydrogenase kinase improves pulmonary arterial hypertension in genetically susceptible patients[J]. Science Translational Medicine, 2017, 9(413):eaao4583.doi:10.1126/scitranslmed.aao4583. URL
[7]	Zhang S, Hulver M W, McMillan R P, Cline M A, Gilbert E R. The pivotal role of pyruvate dehydrogenase kinases in metabolic flexibility[J]. Nutrition & Metabolism, 2014, 11(1):10.doi:10.1186/1743-7075-11-10.
[8]	Sugden M C, Kraus A, Harris R A, Holness M J. Fibre-type specific modification of the activity and regulation of skeletal muscle pyruvate dehydrogenase kinase (PDK) by prolonged starvation and refeeding is associated with targeted regulation of PDK isoenzyme 4 expression[J]. The Biochemical Journal, 2000, 346(3):651-657.doi:10.1042/bj3460651. URL
[9]	Sun S J, Liu J, Zhao M S, Han Y Y, Chen P B, Mo Q Q, Wang B B, Chen G, Fang Y, Tian Y, Zhou J F, Ma D, Gao Q L, Wu P. Loss of the novel mitochondrial protein FAM210B promotes metastasis via PDK4-dependent metabolic reprogramming[J]. Cell Death & Disease, 2017, 8(6):e2870.doi:10.1038/cddis.2017.273.
[10]	Xie Y, Wang H, Cheng X F, Wu Y Z, Cao L J, Wu M Q, Xie W, Wang G J, Hao H P. Farnesoid X receptor activation promotes cell proliferation via PDK4-controlled metabolic reprogramming[J]. Scientific Reports, 2016, 6:18751.doi:10.1038/srep18751. pmid: 26728993
[11]	Jeoung N H, Wu P F, Joshi M A, Jaskiewicz J, Bock C B, Depaoli-Roach A A, Harris R A. Role of pyruvate dehydrogenase kinase isoenzyme 4 (PDHK4) in glucose homoeostasis during starvation[J]. The Biochemical Journal, 2006, 397(3):417-425.doi:10.1042/bj20060125. URL
[12]	Kase E T, Wensaas A J, Aas V, Højlund K, Levin K, Thoresen G H, Beck-Nielsen H, Rustan A C, Gaster M. Skeletal muscle lipid accumulation in type 2 diabetes may involve the liver X receptor pathway[J]. Diabetes, 2005, 54(4):1108-1115.doi:10.2337/diabetes.54.4.1108. pmid: 15793250
[13]	Caton P W, Holness M J, Bishop-Bailey D, Sugden M C. PPARα-LXR as a novel metabolostatic signalling axis in skeletal muscle that acts to optimize substrate selection in response to nutrient status[J]. The Biochemical Journal, 2011, 437(3):521-530.doi:10.1042/BJ20110702. URL
[14]	Tao R, Xiong X, Harris R A, White M F, Dong X C. Genetic inactivation of pyruvate dehydrogenase kinases improves hepatic insulin resistance induced diabetes[J]. PLoS One, 2013, 8(8):e71997.doi:10.1371/journal.pone.0071997. URL
[15]	Cadoudal T, Distel E, Durant S, Fouque F, Blouin J M, Collinet M, Bortoli S, Forest C, Benelli C. Pyruvate dehydrogenase kinase 4:regulation by thiazolidinediones and implication in glyceroneogenesis in adipose tissue[J]. Diabetes, 2008, 57(9):2272-2279.doi:10.2337/db08-0477. pmid: 18519799
[16]	马小勇, 马海梅, 戴荣凤, 马晓明, 吴晓云, 喇永福, 褚敏, 郭宪, 阎萍, 梁春年. 牦牛GEM基因克隆及生物信息学分析[J]. 中国草食动物科学, 2022, 6(6):25-31.doi:10.3969/j.issn.2095-3887.2022.06.005.
	Ma X Y, Ma H M, Dai R F, Ma X M, Wu X Y, La Y F, Chu M, Guo X, Yan P, Liang C N. Cloning and bioinformatics analysis of GEM gene in yak[J]. China Herbivore Science, 2022, 6(6):25-31.
[17]	顾亚荣, 马兰花, 王福彬, 张永峰, 阎萍, 潘和平. 牦牛FTO基因克隆及表达分析[J]. 农业生物技术学报, 2022, 30(4):51.doi:10.3969/j.issn.1674-7968.2022.04.005.
	Gu Y R, Ma L H, Wang F B, Zhang Y F, Yan P, Pan H P. Cloning and expression analysis of FTO gene in yak (Bos grunniens)[J]. Journal of Agricultural Biotechnology, 2022, 30(4):51.
[18]	马彦茹, 于永生, 曹阳, 秦立红, 赵玉民, 吴健. 中国草原红牛 PDK4基因多态性及其与肉质性状的关联分析[J]. 中国畜牧兽医, 2022, 49(6):2186-2194.doi:10.16431/j.cnki.1671-7236.2022.06.019.
	Ma Y R, Yu Y S, Cao Y, Qin L H, Zhao Y M, Wu J. Polymorphism of PDK4 gene and its association with meat quality traits in Chinese steppe red cattle[J]. China Animal Husbandry & Veterinary Medicine, 2022, 49(6):2186-2194.
[19]	Popov K M, Hawes J W, Harris R A. Mitochondrial alpha-ketoacid dehydrogenase kinases:a new family of protein kinases[J]. Advances in Second Messenger and Phosphoprotein Research, 1997, 31:105-111. pmid: 9344245
[20]	Moon H, Lim H S. Synthesis and screening of small-molecule α-helix mimetic libraries targeting protein-protein interactions[J]. Current Opinion in Chemical Biology, 2015, 24:38-47.doi:10.1016/j.cbpa.2014.10.023. pmid: 25461722
[21]	Kim J H, Lee J, Oh B, Kimm K, Koh I. Prediction of phosphorylation sites using SVMs[J]. Bioinformatics, 2004, 20(17):3179-3184.doi:10.1093/bioinformatics/bth382. pmid: 15231530
[22]	Li Z, Peng Y X, Li J X, Chen Z J, Chen F, Tu J, Lin S B, Wang H S. N⁶-methyladenosine regulates glycolysis of cancer cells through PDK4[J]. Biomolecules, 2020, 11(1):2578.doi:10.1038/s41467-020-16306-5.
[23]	Song X X, Liu J, Kuang F M, Chen X, Zeh H J 3rd, Kang R, Kroemer G, Xie Y C, Tang D L. PDK4 dictates metabolic resistance to ferroptosis by suppressing pyruvate oxidation and fatty acid synthesis[J]. Cell Reports, 2021, 34(8):108767.doi:10.1016/j.celrep.2021.108767. URL
[24]	Zhao Y L, Tran M, Wang L, Shin D J, Wu J G. PDK4-deficiency reprograms intrahepatic glucose and lipid metabolism to facilitate liver regeneration in mice[J]. Hepatology Communications, 2020, 4(4):504-517.doi:10.1002/hep4.1484. pmid: 32258946
[25]	Sugden M C, Holness M J. Recent advances in mechanisms regulating glucose oxidation at the level of the pyruvate dehydrogenase complex by PDKs[J]. American Journal of Physiology Endocrinology and Metabolism, 2003, 284(5):E855-E862.doi:10.1152/ajpendo.00526.2002. URL
[26]	蔡沛燃, 邹菊红, 胡艳, 张叁保, 宋颖, 蒋钦杨. 努比亚山羊PDK4基因克隆、序列分析及表达水平研究[J]. 黑龙江畜牧兽医, 2022(9):55-60,141.doi:10.13881/j.cnki.hljxmsy.2021.07.0235.
	Cai P R, Zou J H, Hu Y, Zhang S B, Song Y, Jiang Q Y. Study on cloning,sequence analysis and expression level of PDK4 gene in Nubian goats[J]. Heilongjiang Animal Science and Veterinary Medicine, 2022(9):55-60,141.
[27]	孙瑞萍, 王峰, 晁哲, 刘海隆, 邢漫萍, 刘圈炜, 黄丽丽, 郑心力, 魏立民. 屯昌猪PDK4基因克隆及其组织表达分析[J]. 浙江农业学报, 2020, 32(6):978-985.doi:10.3969/j.issn.1004-1524.2020.06.05.
	Sun R P, Wang F, Chao Z, Liu H L, Xing M P, Liu Q W, Huang L L, Zheng X L, Wei L M. Cloning and tissue expression analysis of PDK4 gene in Tunchang pig[J]. Acta Agriculturae Zhejiangensis, 2020, 32(6):978-985.
[28]	杨洋, 许厚强, 陈伟, 周迪, 徐敏, 张青青, 赵焕平, 孙成娟, 王圆圆, 张鸣, 杨涛. 贵州从江香猪PDK4、FGF10基因的克隆及组织表达分析[J]. 中国畜牧兽医, 2017, 44(12):3401-3409.doi:10.16431/j.cnki.1671-7236.2017.12.002.
	Yang Y, Xu H Q, Chen W, Zhou D, Xu M, Zhang Q Q, Zhao H P, Sun C J, Wang Y Y, Zhang M, Yang T. Cloning and tissue expression analysis of PDK4 and FGF10 genes in Guizhou Congjiang Xiang pig[J]. China Animal Husbandry & Veterinary Medicine, 2017, 44(12):3401-3409.
[29]	孟宪然. 绒山羊背最长肌比较转录组分析及肉品质候选基因筛选[D]. 呼和浩特: 内蒙古农业大学, 2015.
	Meng X R. Comparative transcriptome analysis of longissimus dorsi muscle in Cashmere goats and screening of candidate genes for meat quality[D]. Hohhot: Inner Mongolia Agricultural University, 2015.
[30]	奚子英, 何小梅, 陈刚, 周加义, 钟翠丽, 李永霞, 张榕婧, 聂庆华, 刘满清. 鸡PDK4基因的克隆及组织差异表达研究[J]. 中国家禽, 2015, 37(15):6-10.doi:10.16372/j.issn.1004-6364.2015.15.002.
	Xi Z Y, He X M, Chen G, Zhou J Y, Zhong C L, Li Y X, Zhang R J, Nie Q H, Liu M Q. Cloning and tissue differential expression analysis of chicken PDK4 gene[J]. China Poultry, 2015, 37(15):6-10.
[31]	Lan J, Lei M G, Zhang Y B, Wang J H, Feng X T, Xu D Q, Gui J F, Xiong Y Z. Characterization of the porcine differentially expressed PDK4 gene and association with meat quality[J]. Molecular Biology Reports, 2009, 36(7):2003-2010.doi:10.1007/s11033-008-9411-4.
[32]	Nii M, Hayashi T, Tani F, Niki A, Mori N, Fujishima-Kanaya N, Komatsu M, Aikawa K, Awata T, Mikawa S. Quantitative trait loci mapping for fatty acid composition traits in perirenal and back fat using a Japanese wild boar×Large White intercross[J]. Animal Genetics, 2006, 37(4):342-347.doi:10.1111/j.1365-2052.2006.01485.x.

选择文件类型/文献管理软件名称

选择包含的内容

美仁牦牛PDK4基因克隆及组织表达

Cloning and Tissue Expression Analysis of PDK4 Gene in Meiren Yak

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

图/表 10

参考文献 32

相关文章 15

编辑推荐

Metrics

本文评价

[1]	赖章龙, 阮超岭, 肖伟, 刘彦斌, 俞兆曦, 刘凯, 赛清云, 田永华, 张锋, 杨瑞兰, 柳婷, 杨立强, 王玉涛, 连总强. *兰州鲇NPY基因的克隆和表达分析*[J]. 华北农学报, 2023, 38(5): 227-238.
[2]	黄渺, 罗悠悠, 倪芮, 肇瑾. *观赏辣椒ARP基因克隆及表达分析*[J]. 华北农学报, 2023, 38(5): 69-76.
[3]	刘晓龙, 廖婧芃, 钟歆, 段惜淼, 胡永轩, 刘嘉诚, 刘泽凯, 杨洪涛. 不同生育期高温胁迫下水稻OsRboh家族基因的表达模式[J]. 华北农学报, 2023, 38(5): 1-9.
[4]	王茂存, 曹嘉伟, 周贺, 贾明轩, 魏淑珍, 巩校东, 刘玉卫, 谷守芹, 董金皋. *玉米大斑病菌转录因子基因StbZIP9的克隆、表达分析及其下游靶基因的筛选*[J]. 华北农学报, 2023, 38(5): 158-165.
[5]	杨国武, 王佟, 马小勇, 扎老, 路建卫, 赵雪, 阎萍, 梁春年. 美仁牦牛PHB基因克隆及组织表达[J]. 华北农学报, 2023, 38(5): 218-226.
[6]	聂利珍, 张志成, 谢锐, 常悦, 张琼琳, 韩平安, 羿静. *彩色马铃薯StANS1a基因的克隆及表达分析*[J]. 华北农学报, 2023, 38(4): 38-46.
[7]	周倩怡, 黄思杰, 田洁. 大蒜2个中性/碱性转化酶基因AsNI的克隆与表达分析[J]. 华北农学报, 2023, 38(4): 54-64.
[8]	陈阳, 王琦, 高岩松, 尤学, 熊毅, 金一锋. *草地早熟禾蛋白激酶CIPK32基因克隆及非生物胁迫响应分析*[J]. 华北农学报, 2023, 38(4): 65-73.
[9]	赵胜男, 高美欣, 于朝杭, 聂凯悦, 王浩然, 孟杰, 朱虹, 李帅. *大豆GmFLK基因的克隆及特性分析*[J]. 华北农学报, 2023, 38(3): 26-34.
[10]	张沛沛, 陈涛, 景凡丽, 刘媛, 马靖福, 田甜, 王鹏, 杨德龙. *小麦TaPSKR1基因的克隆与表达分析*[J]. 华北农学报, 2023, 38(3): 1-9.
[11]	王川川, 母童, 李德生, 张迪, 李蕾蕾, 张娟, 顾亚玲. *荷斯坦奶牛FABP4基因结构与功能分析*[J]. 华北农学报, 2023, 38(3): 204-212.
[12]	王鹏, 田哲娟, 赵雪芳, 康忱, 吴志明, 李亚栋, 黄冀楠. *番茄SlCaM3、SlCaM4和SlCaM5基因的克隆及低温胁迫下的表达分析*[J]. 华北农学报, 2023, 38(2): 75-84.
[13]	王莹, 王晓宇, 孙德慧, 霍红雁, 刘海臣, 徐惠, 张继星. 蓖麻钙依赖蛋白激酶29基因克隆与表达分析[J]. 华北农学报, 2023, 38(2): 85-92.
[14]	范依琳, 赵丹, 马妍, 岳永起, 冯欣欣, 张姬越, 字向东, 熊显荣, 付伟, 熊燕. *牦牛ANXA5基因的克隆、序列分析及表达研究*[J]. 华北农学报, 2023, 38(2): 224-231.
[15]	周小南, 师丹丹, 丁燕玲, 张岩峰, 赵志艳, 康晓龙. *秦川牛CDS1基因克隆、分子特征及组织表达分析*[J]. 华北农学报, 2023, 38(1): 218-225.

模态框（Modal）标题

选择文件类型/文献管理软件名称

选择包含的内容

美仁牦牛PDK4基因克隆及组织表达

Cloning and Tissue Expression Analysis of PDK4 Gene in Meiren Yak

RichHTML

PDF

可视化

被引次数

摘要/Abstract

引用本文

使用本文

图/表 10

参考文献 32

相关文章 15

编辑推荐

Metrics

本文评价