过表达组蛋白H3K9me3去甲基化酶对猪克隆胚胎发育的影响

doi:10.16288/j.yczz.22-244

遗传 ›› 2023, Vol. 45 ›› Issue (1): 67-77.doi: 10.16288/j.yczz.22-244

过表达组蛋白H3K9me3去甲基化酶对猪克隆胚胎发育的影响

李亚楠^1,^2,³(), 张贤君^1,^2,³, 张宁^1,^2,³, 梁雅琳^1,^2,³, 张宇星^1,^2,³, 招华兴^1,^2,³, 李紫聪^1,^2,³, 黄思秀^1,^2,³()

1.华南农业大学动物科学学院，国家生猪种业工程技术研究中心，广州 510642
2.畜禽育种国家地方联合工程研究中心，广州 510642
3.广东省农业动物基因组学与分子育种重点实验室，广州 510642

收稿日期:2022-07-18 修回日期:2022-11-08 出版日期:2023-01-20 发布日期:2022-11-28
通讯作者: 黄思秀 E-mail:li321736803@163.com;sxhuang815@scau.edu.cn
作者简介:李亚楠，在读硕士研究生，专业方向：动物遗传育种与繁殖。E-mail: li321736803@163.com
基金资助:
广东省自然科学基金项目(2019B1515210027);广东特支计划本土创新创业团队项目(2019BT02N630);广东省畜禽地方品种保护与开发利用提升工程项目资助

Effects of overexpression of histone H3K9me3 demethylase on development of porcine cloned embryos

Yanan Li^1,^2,³(), Xianjun Zhang^1,^2,³, Ning Zhang^1,^2,³, Yalin Liang^1,^2,³, Yuxing Zhang^1,^2,³, Huaxing Zhao^1,^2,³, Zicong Li^1,^2,³, Sixiu Huang^1,^2,³()

1. National Engineering Research Center for Breeding Swine Industry, College of Animal Science of South China Agricultural University, Guangzhou 510642, China
2. National & Local Joint Engineering Research Center for Breeding Animal Industry, Guangzhou 510642, China
3. Guangdong Provincial Key Laboratory of Agro-animal Genomics and Molecular Breeding, Guangzhou 510642, China

Received:2022-07-18 Revised:2022-11-08 Online:2023-01-20 Published:2022-11-28
Contact: Huang Sixiu E-mail:li321736803@163.com;sxhuang815@scau.edu.cn
Supported by:
the Natural Science Foundation of Guangdong Province(2019B1515210027);Guangdong Special Support Plan Local Innovation and Entrepreneurship Team(2019BT02N630);Guangdong Province Livestock and poultry Local Breed Protection and Development and Utilization Promotion Project]

摘要/Abstract

摘要：

组蛋白异常修饰是克隆胚胎发育的重要制约因素，组蛋白H3K9me3去甲基化酶KDM4家族的过表达可以有效提高克隆胚胎的发育效率。为探究过表达H3K9me3去甲基化酶对猪克隆胚胎发育的影响，本研究在猪克隆胚胎1-细胞期和2-细胞期分别注射KDM4A mRNA和KDM4D mRNA检测胚胎的囊胚率；收集1-细胞期注射KDM4A mRNA和胚胎注射水(对照组)的2-细胞期克隆胚胎检测H3K9me3表达水平；此外，收集1-细胞期注射KDM4A mRNA和胚胎注射水的4-细胞期克隆胚胎进行单细胞转录组测序，并对测序数据进行GO与KEGG富集分析。结果显示：在1-细胞期注射KDM4A mRNA 的猪克隆胚胎囊胚率显著高于对照组(25.32 ± 0.74% vs 14.78 ± 0.87%)，注射KDM4D mRNA对猪克隆胚胎囊胚率无明显作用(16.27 ± 0.77% vs 14.78 ± 0.87%)；在2-细胞期注射KDM4A mRNA和KDM4D mRNA的克隆胚胎囊胚率与对照组相比均无显著差异(32.18 ± 1.67%、30.04 ± 0.91% vs 31.22 ± 1.40%)。在1-细胞期注射KDM4A mRNA的克隆胚胎组蛋白H3K9me3表达水平低于对照组。通过测序，筛选出133个差异表达基因，其中上调基因52个、下调基因81个，GO分析主要富集到与蛋白质定位相关的通路，KEGG分析富集到细胞衰老和急性髓细胞白血病等相关通路。本研究结果表明，过表达组蛋白H3K9me3去甲基化酶KDM4A可以显著提高猪克隆胚胎的发育效率。

关键词: 猪, 克隆胚胎, H3K9me3, KDM4A, KDM4D

Abstract:

The abnormal modification of histone is an important factor restricting development of porcine cloned embryos. Overexpression of histone H3K9me3 demethylase KDM4 family can effectively improve the developmental efficiency of cloned embryos. In order to explore the effects of overexpression of H3K9me3 demethylase on the development of porcine cloned embryos, KDM4A mRNA and KDM4D mRNA were injected respectively into porcine cloned embryos at the 1-cell stage and 2-cell stage to detect the blastocyst rate; 2-cell stage cloned embryos injected with KDM4A mRNA and embryo injection water (the control group) at the 1-cell stage were collected to detect the expression level of H3K9me3, and 4-cell stage cloned embryos were collected for single cell transcriptome sequencing, then the sequencing data was analyzed with KEGG and GO. The results showed that the blastocyst rate of porcine cloned embryos injected with KDM4A mRNA at 1-cell stage was significantly higher than that of the control group (25.32 ± 0.74% vs 14.78 ± 0.87%), while cloned embryos injected with KDM4D mRNA had a similar blastocyst rate with cloned embryos in control group (16.27 ± 0.77% vs 14.78 ± 0.87%). Porcine cloned embryos injected with KDM4A mRNA and KDM4D mRNA at 2-cell stage had a similar blastocyst rate with cloned embryos in control group (32.18 ± 1.67%, 30.04 ± 0.91% vs 31.22 ± 1.40%). The expression level of H3K9me3 in cloned embryos injected with KDM4A mRNA at 1-cell stage was lower than that in control group. There were 133 differentially expressed genes detected by transcriptome sequencing, including 52 up-regulated genes and 81 down-regulated genes. Pathways enriched by GO analyses were mainly related to protein localization. Pathways enriched by KEGG analyses were related to cellular senescence and acute myeloid leukemia. These results suggest that overexpression of histone H3K9me3 demethylase KDM4A can significantly improve the developmental efficiency of porcine cloned embryos.

Key words: pig, cloning embryo, H3K9me3, KDM4A, KDM4D

李亚楠, 张贤君, 张宁, 梁雅琳, 张宇星, 招华兴, 李紫聪, 黄思秀. 过表达组蛋白H3K9me3去甲基化酶对猪克隆胚胎发育的影响[J]. 遗传, 2023, 45(1): 67-77.

Yanan Li, Xianjun Zhang, Ning Zhang, Yalin Liang, Yuxing Zhang, Huaxing Zhao, Zicong Li, Sixiu Huang. Effects of overexpression of histone H3K9me3 demethylase on development of porcine cloned embryos[J]. Hereditas(Beijing), 2023, 45(1): 67-77.

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参考文献 38

[1]	Liu Y, Li J, Løvendahl P, Schmidt M, Larsen K, Callesen H. In vitro manipulation techniques of porcine embryos: a meta-analysis related to transfers, pregnancies and piglets. Reprod Fertil Dev, 2015, 27(3): 429-439.
[2]	Yang XQ, Wu ZF, Li ZC. Advances in epigenetic reprogramming of somatic cells nuclear transfer in mammals. Hereditas (Beijing), 2019, 41(12): 1099-1109.
	杨旭琼, 吴珍芳, 李紫聪. 哺乳动物体细胞核移植表观遗传重编程研究进展. 遗传, 2019, 41(12): 1099-1109.
[3]	Huang XW, Cheng XR, Wang N, Zhang YW, Liao C, Jin LH, Lei L. Histone variant H3.3 and its functions in reprogramming. Hereditas (Beijing), 2018, 40(3): 186-196.
	黄星卫, 程香荣, 王楠, 张雨薇, 廖辰, 金连弘, 雷蕾. 组蛋白H3变体H3.3及其在细胞重编程中的作用. 遗传, 2018, 40(3): 186-196.
[4]	Chung YG, Matoba S, Liu YT, Eum JH, Lu F, Jiang W, Lee JE, Sepilian V, Cha KY, Lee DR, Zhang Y. Histone demethylase expression enhances human somatic cell nuclear transfer efficiency and promotes derivation of pluripotent stem cells. Cell Stem Cell, 2015, 17(6): 758-766.
[5]	Matoba S, Liu YT, Lu FL, Iwabuchi KA, Shen L, Inoue A, Zhang Y. Embryonic development following somatic cell nuclear transfer impeded by persisting histone methylation. Cell, 2014, 159(4): 884-895.
[6]	Meng FL, Stamms K, Bennewitz R, Green A, Oback F, Turner P, Wei JW, Oback B. Targeted histone demethylation improves somatic cell reprogramming into cloned blastocysts but not postimplantation bovine concepti. Biol Reprod, 2020, 103(1): 114-125.
[7]	Zhang YM, Wang QQ, Liu KL, Gao EE, Guan H, Hou J. Treatment of donor cells with recombinant KDM4D protein improves preimplantation development of cloned ovine embryos. Cytotechnology, 2018, 70(5): 1469-1477.
[8]	Liu Z, Cai YJ, Wang Y, Nie YH, Zhang CC, Xu YT, Zhang XT, Lu Y, Wang ZY, Poo M, Sun Q. Cloning of macaque monkeys by somatic cell nuclear transfer. Cell, 2018, 172(4): 881-887.
[9]	Janjic A, Wange LE, Bagnoli JW, Geuder J, Nguyen P, Richter D, Vieth B, Vick B, Jeremias I, Ziegenhain C, Hellmann I, Enard W. Prime-seq, efficient and powerful bulk RNA sequencing. Genome Biol, 2022, 23(1): 88.
[10]	Matoba S, Zhang Y. Somatic cell nuclear transfer reprogramming: mechanisms and applications. Cell Stem Cell, 2018, 23(4): 471-485.
[11]	Liu WQ, Liu XY, Wang CF, Gao YW, Gao R, Kou XC, Zhao YH, Li JY, Wu Y, Xiu WC, Wang S, Yin JQ, Liu W, Cai T, Wang H, Zhang Y, Gao SR. Identification of key factors conquering developmental arrest of somatic cell cloned embryos by combining embryo biopsy and single-cell sequencing. Cell Discov, 2016, 2: 16010.
[12]	Wu X, Li G, Ao Z, Shi JS, Cai GY, Liu DW, Wu ZF, Li ZC. Effects of overexpression of H3K9me3 demethylase on the in vitro developmental efficiency of cloned porcine embryos. Guangdong Agric Sci, 2017, 44(10): 96-101.
	吴霄, 李果, 敖政, 石俊松, 蔡更元, 刘德武, 吴珍芳, 李紫聪. 过表达H3K9me3去甲基化酶对猪克隆胚胎体外发育效率的影响. 广东农业科学, 2017, 44(10): 96-101.
[13]	He XY, Tan C, Li ZC, Zhao CF, Shi JS, Zhou R, Wang XW, Jiang GL, Cai GY, Liu DW, Wu ZF. Characterization and comparative analyses of transcriptomes of cloned and in vivo fertilized porcine pre-implantation embryos. Biol Open, 2019, 8(4): bio039917.
[14]	Liu X, Chen L, Wang T, Zhou JL, Li ZK, Bu GW, Zhang JJ, Yin SY, Wu DY, Dou CL, Xu T, He HN, Zhu W, Yu LT, Liu ZT, Zhang X, Chen ZX, Miao YL. TDG is a pig-specific epigenetic regulator with insensitivity to H3K9 and H3K27 demethylation in nuclear transfer embryos. Stem Cell Reports, 2021, 16(11): 2674-2689.
[15]	Sankar A, Kooistra SM, Gonzalez JM, Ohlsson C, Poutanen M, Helin K. Maternal expression of the histone demethylase KDM4A is crucial for pre-implantation development. Development, 2017, 144(18): 3264-3277.
[16]	Lee JE, Chung YG, Eum JH, Lee Y, Lee DR. An efficient SCNT technology for the establishment of personalized and public human pluripotent stem cell banks. BMB Rep, 2016, 49(4): 197-198.
[17]	Ruan DG, Peng JY, Wang XS, Ouyang Z, Zou QJ, Yang Y, Chen FB, Ge WK, Wu H, Liu ZM, Zhao Y, Zhao BT, Zhang QJ, Lai CD, Fan NN, Zhou ZW, Liu QS, Li N, Jin Q, Shi H, Xie JK, Song H, Yang XY, Chen JK, Wang KP, Li XP, Lai LX. XIST derepression in active X chromosome hinders pig somatic cell nuclear transfer. Stem Cell Reports, 2018, 10(2): 494-508.
[18]	Weng XG, Cai MM, Zhang YT, Liu Y, Liu C, Liu ZH. Improvement in the in vitro development of cloned pig embryos after KDM4A overexpression and an H3K9me3 methyltransferase inhibitor treatment. Theriogenology, 2020, 146: 162-170.
[19]	Chen M, Zhu QS, Li C, Kou XC, Zhao YH, Li YH, Xu RM, Yang L, Yang LY, Gu L, Wang H, Liu XY, Jiang CZ, Gao SR. Chromatin architecture reorganization in murine somatic cell nuclear transfer embryos. Nat Commun, 2020, 11(1): 1813.
[20]	Feng Y, Zhao X, Li ZD, Luo C, Ruan ZY, Xu J, Shen PL, Deng YF, Jiang JR, Shi DS, Lu FH. Histone demethylase KDM4D could improve the developmental competence of buffalo (Bubalus bubalis) somatic cell nuclear transfer (SCNT) embryos. Microsc Microanal, 2021, 27(2): 409-419.
[21]	Wang CF, Liu XY, Gao YW, Yang L, Li C, Liu WQ, Chen C, Kou XC, Zhao YH, Chen JY, Wang YX, Le RR, Wang H, Duan T, Zhang Y, Gao SR. Reprogramming of H3K9me3- dependent heterochromatin during mammalian embryo development. Nat Cell Biol, 2018, 20(5): 620-631.
[22]	Cao ZB, Li YS, Chen Z, Wang H, Zhang ML, Zhou NR, Wu RH, Ling YH, Fang FG, Li N, Zhang YH. Genome- wide dynamic profiling of histone methylation during nuclear transfer-mediated porcine somatic cell reprogramming. PLoS One, 2015, 10(12): e0144897.
[23]	Hernández-Rosas F, Hernández-Oliveras A, Flores-Peredo L, Rodríguez G, Zarain-Herzberg Á, Caba M, Santiago-García J. Histone deacetylase inhibitors induce the expression of tumor suppressor genes Per1 and Per2 in human gastric cancer cells. Oncol Lett, 2018, 16(2): 1981-1990.
[24]	Zhao Y, Huang SY, Tan XR, Long LF, He QM, Liang XY, Bai JW, Li QJ, Lin JY, Li YQ, Liu N, Ma J, Chen YP. N6-methyladenosine-modified CBX1 regulates nasopharyngeal carcinoma progression through heterochromatin formation and STAT1 activation. Adv Sci (Weinh), 2022, 30: e2205091.
[25]	Divisato G, Chiariello AM, Esposito A, Zoppoli P, Zambelli F, Elia MA, Pesole G, Incarnato D, Passaro F, Piscitelli S, Oliviero S, Nicodemi M, Parisi S, Russo T. Hmga2 protein loss alters nuclear envelope and 3D chromatin structure. BMC Biol, 2022, 20(1): 171.
[26]	Dong GZ, Zhang R, Hu Q, Martin EM, Qin YF, Lu CC, Xia YK, Wang XR, Du GZ. Prothioconazole induces cell cycle arrest by up-regulation of EIF4EBP1 in extravillous trophoblast cells. Arch Toxicol, 2022, 96(2): 559-570.
[27]	Mahdipour M, Leitoguinho AR, Zacarias Silva RA, van Tol HT, Stout TA, Rodrigues G, Roelen BA. TACC3 is important for correct progression of meiosis in bovine oocytes. PLoS One, 2015, 10(7): e0132591.
[28]	Heinemann-Yerushalmi L, Bentovim L, Felsenthal N, Vinestock RC, Michaeli N, Krief S, Silberman A, Cohen M, Ben-Dor S, Brenner O, Haffner-Krausz R, Itkin M, Malitsky S, Erez A, Zelzer E. BCKDK regulates the TCA cycle through PDC in the absence of PDK family during embryonic development. Dev Cell, 2021, 56(8): 1182-1194.e6.
[29]	Krieg AJ, Mullinax SR, Grimstad F, Marquis K, Constance E, Hong Y, Krieg SA, Roby KF. Histone demethylase KDM4A and KDM4B expression in granulosa cells from women undergoing in vitro fertilization. Assist Reprod Genet, 2018, 35(6): 993-1003.
[30]	Zoabi M, Nadar-Ponniah PT, Khoury-Haddad H, Usaj M, Budowski-Tal I, Haran T, Henn A, Mandel-Gutfreund Y, Ayoub N. RNA-dependent chromatin localization of KDM4D lysine demethylase promotes H3K9me3 demethylation. Nucleic Acids Res, 2014, 42(21): 13026-13038.
[31]	Zhang BY, Long QL, Wu SS, Xu QX, Song SL, Han L, Qian M, Ren XH, Liu HX, Jiang J, Guo JM, Zhang XL, Chang X, Fu Q, Lam EWF, Campisi J, Kirkland JL, Sun Y. KDM4 orchestrates epigenomic remodeling of senescent cells and potentiates the senescence-associated secretory phenotype. Nat Aging, 2021, 1(5): 454-472.
[32]	Massett ME, Monaghan L, Patterson S, Mannion N, Bunschoten RP, Hoose A, Marmiroli S, Liskamp RMJ, Jørgensen HG, Vetrie D, Michie AM, Huang X. A KDM4A-PAF1-mediated epigenomic network is essential for acute myeloid leukemia cell self-renewal and survival. Cell Death Dis, 2021, 12(6): 573.
[33]	Zhao JG, Hao YH, Ross JW, Spate LD, Walters EM, Samuel MS, Rieke A, Murphy CN, Prather RS. Histone deacetylase inhibitors improve in vitro and in vivo developmental competence of somatic cell nuclear transfer porcine embryos. Cell Reprogram, 2010, 12(1): 75-83.
[34]	Kishigami S, Mizutani E, Ohta H, Hikichi T, Thuan NV, Wakayama S, Bui HT, Wakayama T. Significant improvement of mouse cloning technique by treatment with trichostatin a after somatic nuclear transfer. Biochem Biophys Res Commun, 2006, 340(1): 183-189.
[35]	Wang FC, Kou ZH, Zhang Y, Gao SR. Dynamic reprogramming of histone acetylation and methylation in the first cell cycle of cloned mouse embryos. Biol Reprod, 2007, 77(6): 1007-1016.
[36]	Inoue K, Kohda T, Sugimoto M, Sado T, Ogonuki N, Matoba S, Shiura H, Ikeda R, Mochida K, Fujii T, Sawai K, Otte AP, Tian XC, Yang XZ, Ishino F, Abe K, Ogura A. Impeding Xist expression from the active X chromosome improves mouse somatic cell nuclear transfer. Science, 2010, 330(6003): 496-499.
[37]	Li GL, Zhong CL, Ni S, Liu DW, Cai GY, Li ZC, Yang HQ, Wu ZF. Establishment of porcine Xist knockout model using CRISPR/Cas9 system. Hereditas (Beijing), 2016, 38(12): 1081-1089.
	李国玲, 钟翠丽, 倪生, 刘德武, 蔡更元, 李紫聪, 杨化强, 吴珍芳. 利用CRISPR/Cas9系统建立Xist基因敲除猪模型. 遗传, 2016, 38(12): 1081-1089.
[38]	Matoba S, Wang HH, Jiang L, Lu FL, Iwabuchi KA, Wu XJ, Inoue K, Yang L, Press W, Lee JT, Ogura A, Shen L, Zhang Y. Loss of H3K27me 3 imprinting in somatic cell nuclear transfer embryos disrupts post-implantation development. Cell Stem Cell, 2018, 23(3): 343-354.

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组别	胚胎数(n)	卵裂数	卵裂率(%)	囊胚数	囊胚率(%)	囊胚细胞数
KDM4A	308(8)	228	74.03 ± 0.95	78	25.32 ± 0.74^aA	42.11 ± 2.39
KDM4D	295(8)	217	73.45 ± 0.92	48	16.27 ± 0.77^bB	43.79 ± 2.70
KDM4A+KDM4D	216(7)	155	71.76 ± 0.93	41	18.98 ± 1.01^abAB	38.59 ± 2.8
胚胎注射水(对照)	345(9)	250	72.46 ± 0.65	51	14.78 ± 0.87^bB	40.27 ± 2.22

组别	胚胎数(n)	囊胚数	囊胚率(%)	囊胚细胞数
KDM4A	202(6)	65	32.18 ± 1.67	35.18 ± 1.73^ab
KDM4D	243(7)	73	30.04 ± 0.91	32.47 ± 1.75^b
KDM4A+KDM4D	88(3)	26	29.55 ± 1.95	40.33 ± 4.00^a
胚胎注射水(对照)	221(6)	69	31.22 ± 1.40	35.44 ± 2.05^ab

过表达组蛋白H3K9me3去甲基化酶对猪克隆胚胎发育的影响

Effects of overexpression of histone H3K9me3 demethylase on development of porcine cloned embryos

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[13]	莫健新,王豪强,黄广燕,蔡更元,吴珍芳,张献伟. 微生物源果胶酶在猪PK15细胞中异源表达及其酶学性质分析[J]. 遗传, 2019, 41(8): 736-745.
[14]	周李生, 赵为民, 涂枫, 吴云鹤, 任守文, 方晓敏. 猪乳头性状生理学和遗传学研究进展[J]. 遗传, 2019, 41(5): 384-390.
[15]	黄子莹, 李龙, 李倩倩, 刘向东, 李长春. lncRNA TCONS_00815878对猪骨骼肌卫星细胞分化的影响[J]. 遗传, 2019, 41(12): 1119-1128.