Abstract：Heterogeneous nuclear ribonucleoprotein K (hnRNP K), a member of hnRNP complex, can interact with nucleic acids and various proteins. The structure of hnRNP K protein mainly contains three K homology (KH) domains that can combine with DNA or RNA, and one K protein interactive region (KI) domain involved in protein interaction. It also contains a nuclear localization signal (NLS) in the N terminal that mediates its transport from the cytoplasm to the nucleus and a nuclear shuttling domain (KNS) in the C terminal that confers the ability to translocate bi-directionally through the nuclear pore complex. In eukaryotes, hnRNP K expresses in many type of cells and tissues, but the expression levels vary during different developmental phases, and it is located predominantly in the nucleus and cytoplasm, also in mitochondria and plasma membrane. In addition, hnRNP K is subject to several post-translational modifications, such as phosphorylation, methylation, sumoylation, ubiquitylatio, acetylation, and so on, which can regulate its interactions with different molecules and influence its functions. The complexity of structure, expression, location and the diversity of post-translational modifications in hnRNP K decide its abundant biological functions, which can participate in transcription, translation, RNA splicing, DNA repair, chromatin remodeling, and also have important roles in spermatogenesis, nervous system and ovary development, organogenesis and carcinogenesis process. Among them, spermatogenesis is tightly regulated and well-studied process consisting of a series of highly specific cellular processes and multiple genes involved in the final formation of the sperm. Many research have shown that hnRNP K may play a key role in spermatogenesis regulation by its special structure, selective splicing and expression pattern. This review focuses on the structure, expression regulation, post-translational modification and biological functions of hnRNP K, and its roles in spermatogenesis. It provides initial value?for?studying?the?biological?functions?of?hnRNP K especially in spermatogenesis.
[1]Abram, C.L. and SA. Courtneidge. Src family tyrosine kinases and growth factor signaling[J].Exp Cell Res, 2000, 254(1):1-13
[2]Adolph, D.N. Flach,KMueller,D. H. Ostareck and A. Ostareck-Lederer. Deciphering the cross talk between hnRNP K and c-Src: the c-Src activation domain in hnRNP K is distinct from a second interaction site[J].Mol Cell Biol, 2007, 27(5):1758-1770
[3]Berkovits, B.D.,LWang,P. Guarnieri and D. J. Wolgemuth. The testis-specific double bromodomain-containing protein BRDT forms a complex with multiple spliceosome components and is required for mRNA splicing and 3'-UTR truncation in round spermatids[J].Nucleic Acids Res, 2012, 40(15):7162-7175
[4]Bomsztyk, K.O. Denisenko and JOstrowski. hnRNP K: one protein multiple processes[J].Bioessays, 2004, 26(6):629-638
[5]Bomsztyk, K.I. Van Seuningen,HSuzuki,O. Denisenko and J. Ostrowski. Diverse molecular interactions of the hnRNP K protein[J].FEBS Lett, 1997, 403(2):113-115
[6]Bonham, K.S. ARitchie,S. M. Dehm,K. Snyder and F. M. Boyd. An alternative,human SRC promoter and its regulation by hepatic nuclear factor-1alpha[J].J Biol Chem, 2000, 275(48):37604-37611
[7]Cao, W.A. Razanau,DFeng,V. G. Lobo and J. Xie. Control of alternative splicing by forskolin through hnRNP K during neuronal differentiation[J].Nucleic Acids Res, 2012, 40(16):8059-8071
[8]Chang, Y.I.,SC. Hsu,G. Y. Chau,C. Y. Huang,J. S. Sung,W. K. Hua and W. J. Lin. Identification of the methylation preference region in heterogeneous nuclear ribonucleoprotein K by protein arginine methyltransferase 1 and its implication in regulating nuclearcytoplasmic distribution[J].Biochem Biophys Res Commun, 2011, 404(3):865-869
[9]Chapman, K.M.,HM. Powell,J. Chaudhary,J. M. Shelton,J. A. Richardson,T. E. Richardson and F. K. Hamra. Linking spermatid ribonucleic acid (RNA) binding protein and retrogene diversity to reproductive success[J].Mol Cell Proteomics, 2013, 12(11):3221-3236
[10]Chaudhury, A.P. Chander and PH. Howe. Heterogeneous nuclear ribonucleoproteins (hnRNPs) in cellular processes: Focus on hnRNP E1's multifunctional regulatory roles[J].RNA, 2010, 16(8):1449-1462
[11]Chen, Y.X. Zhou,NLiu,C. Wang,L. Zhang,W. Mo and G. Hu. Arginine methylation of hnRNP K enhances p53 transcriptional activity[J].FEBS Lett, 2008, 582(12):1761-1765
[12]Choi, H.S.,CK. Hwang,K. Y. Song,P. Y. Law,L. N. Wei and H. H. Loh. Poly(C)-binding proteins as transcriptional regulators of gene expression[J].Biochem Biophys Res Commun, 2009, 380(3):431-436
[13]Ciarlo, M.R. Benelli,OBarbieri,S. Minghelli,P. Barboro,C. Balbi and N. Ferrari. Regulation of neuroendocrine differentiation by AKThnRNPKARbeta-catenin signaling in prostate cancer cells[J].Int J Cancer, 2012, 131(3):582-590
[14]Collier, B.L. Goobar-Larsson,MSokolowski and S. Schwartz. Translational inhibition in vitro of human papillomavirus type 16 L2 mRNA mediated through interaction with heterogenous ribonucleoprotein K and poly(rC)-binding proteins 1 and 2[J].J Biol Chem, 1998, 273(35):22648-22656
[15]Cui, Y.H. Zhu,YZhu,X. Guo,R. Huo,X. Wang,J. Tong,L. Qian,Z. Zhou,Y. Jia,Y. H. Lue,A. S. Hikim,C. Wang,R. S. Swerdloff and J. Sha. Proteomic analysis of testis biopsies in men treated with injectable testosterone undecanoate alone or in combination with oral levonorgestrel as potential male contraceptive[J].J Proteome Res, 2008, 7(9):3984-3993
[16]Du, Q.I. NMelnikova and P. D. Gardner. Differential effects of heterogeneous nuclear ribonucleoprotein K on Sp1- and Sp3-mediated transcriptional activation of a neuronal nicotinic acetylcholine receptor promoter[J].J Biol Chem, 1998, 273(31):19877-19883
[17]Eddy, E.M. Male germ cell gene expression [J][J].Recent Prog Horm Res, 2002, 57(57):103-128
[18]Elliott, D.J. and SN. Grellscheid. Alternative RNA splicing regulation in the testis[J].Reproduction, 2006, 132(6):811-819
[19]Enge, M.W. Bao,EHedstrom,S. P. Jackson,A. Moumen and G. Selivanova. MDM2-dependent downregulation of p21 and hnRNP K provides a switch between apoptosis and growth arrest induced by pharmacologically activated p53[J].Cancer Cell, 2009, 15(3):171-183
[20]Evans, J.R.,SA. Mitchell,K. A. Spriggs,J. Ostrowski,K. Bomsztyk,D. Ostarek and A. E. Willis. Members of the poly (rC) binding protein family stimulate the activity of the c-myc internal ribosome entry segment in vitro and in vivo[J].Oncogene, 2003, 22(39):8012-8020
[21]Expert-Bezancon, A.J. PLe Caer and J. Marie. Heterogeneous nuclear ribonucleoprotein (hnRNP) K is a component of an intronic splicing enhancer complex that activates the splicing of the alternative exon 6A from chicken beta-tropomyosin pre-mRNA[J].J Biol Chem, 2002, 277(19):16614-16623
[22]Feliers, D.M. JLee,G. Ghosh-Choudhury,K. Bomsztyk and B. S. Kasinath. Heterogeneous nuclear ribonucleoprotein K contributes to angiotensin II stimulation of vascular endothelial growth factor mRNA translation[J].Am J Physiol Renal Physiol, 2007, 293(2):F607-615
[23]Gan, H.T. Cai,XLin,Y. Wu,X. Wang,F. Yang and C. Han. Integrative proteomic and transcriptomic analyses reveal multiple post-transcriptional regulatory mechanisms of mouse spermatogenesis[J].Mol Cell Proteomics, 2013, 12(5):1144-1157
[24]Guo, A.H. Gu,JZhou,D. Mulhern,Y. Wang,K. A. Lee,V. Yang,M. Aguiar,J. Kornhauser,X. Jia,J. Ren,S. A. Beausoleil,J. C. Silva,V. Vemulapalli,M. T. Bedford and M. J. Comb. Immunoaffinity enrichment and mass spectrometry analysis of protein methylation[J].Mol Cell Proteomics, 2014, 13(1):372-387
[25]Habelhah, H.K. Shah,LHuang,A. Ostareck-Lederer,A. L. Burlingame,K. M. Shokat,M. W. Hentze and Z. Ronai. ERK phosphorylation drives cytoplasmic accumulation of hnRNP-K and inhibition of mRNA translation[J].Nat Cell Biol, 2001, 3(3):325-330
[26]Harris, D.Z. Zhang,BChaubey and V. N. Pandey. Identification of cellular factors associated with the 3'-nontranslated region of the hepatitis C virus genome[J].Mol Cell Proteomics, 2006, 5(6):1006-1018
[27]He, Y.and RSmith. Nuclear functions of heterogeneous nuclear ribonucleoproteins AB[J].Cell Mol Life Sci, 2009, 66(7):1239-1256
[28]Hsu, P.P.,SA. Kang,J. Rameseder,Y. Zhang,K. A. Ottina,D. Lim,T. R. Peterson,Y. Choi,N. S. Gray,M. B. Yaffe,J. A. Marto and D. M. Sabatini. The mTOR-regulated phosphoproteome reveals a mechanism of mTORC1-mediated inhibition of growth factor signaling[J].Science, 2011, 332(6035):1317-1322
[29]Hsueh, K.W.,SL. Fu,C. Y. Huang and C. H. Lin. Aurora-A phosphorylates hnRNPK and disrupts its interaction with p53[J].FEBS Lett, 2011, 585(17):2671-2675
[30]Hutchins, E.J. and BG. Szaro. c-Jun N-terminal kinase phosphorylation of heterogeneous nuclear ribonucleoprotein K regulates vertebrate axon outgrowth via a posttranscriptional mechanism[J].J Neurosci, 2013, 33(37):14666-14680
[31]Kamma, H.D. SPortman and G. Dreyfuss. Cell type-specific expression of hnRNP proteins[J].Exp Cell Res, 1995, 221(1):187-196
[32]Kim, J.H.,BHahm,Y. K. Kim,M. Choi and S. K. Jang. Protein-protein interaction among hnRNPs shuttling between nucleus and cytoplasm[J].J Mol Biol, 2000, 298(3):395-405
[33]Kim, W.E. JBennett,E. L. Huttlin,A. Guo,J. Li,A. Possemato,M. E. Sowa,R. Rad,J. Rush,M. J. Comb,J. W. Harper and S. P. Gygi. Systematic and quantitative assessment of the ubiquitin-modified proteome[J].Mol Cell, 2011, 44(2):325-340
[34]Kimura, Y.K. Nagata,NSuzuki,R. Yokoyama,Y. Yamanaka,H. Kitamura,H. Hirano and O. Ohara. Characterization of multiple alternative forms of heterogeneous nuclear ribonucleoprotein K by phosphate-affinity electrophoresis[J].Proteomics, 2010, 10(21):3884-3895
[35]Kwiek, N.C.,DF. Thacker,M. B. Datto,H. B. Megosh and T. A. Haystead. PITK,a PP1 targeting subunit that modulates the phosphorylation of the transcriptional regulator hnRNP K[J].Cell Signal, 2006, 18(10):1769-1778
[36]Lau, J.S.,PBaumeister,E. Kim,B. Roy,T. Y. Hsieh,M. Lai and A. S. Lee. Heterogeneous nuclear ribonucleoproteins as regulators of gene expression through interactions with the human thymidine kinase promoter[J].J Cell Biochem, 2000, 79(3):395-406
[37]Laury-Kleintop, L.D.,MTresini and O. Hammond. Compartmentalization of hnRNP-K during cell cycle progression and its interaction with calponin in the cytoplasm[J].J Cell Biochem, 2005, 95(5):1042-1056
[38]Lee, J.J. Hong,EKim,K. Kim,S. W. Kim,H. Krishnamurthy,S. S. Chung,D. J. Wolgemuth and K. Rhee. Developmental stage-specific expression of Rbm suggests its involvement in early phases of spermatogenesis[J].Mol Hum Reprod, 2004, 10(4):259-264
[39]Lee, S.W.,MH. Lee,J. H. Park,S. H. Kang,H. M. Yoo,S. H. Ka,Y. M. Oh,Y. J. Jeon and C. H. Chung. SUMOylation of hnRNP-K is required for p53-mediated cell-cycle arrest in response to DNA damage[J].EMBO J, 2012, 31(23):4441-4452
[40]Leopoldino, A.M.,FCarregaro,C. H. Silva,O. Feitosa,U. M. Mancini,J. M. Freitas and E. H. Tajara. Sequence and transcriptional study of HNRPK pseudogenes,and expression and molecular modeling analysis of hnRNP K isoforms[J].Genome, 2007, 50(5):451-462
[41]Libri, D.A. Piseri and MY. Fiszman. Tissue-specific splicing in vivo of the beta-tropomyosin gene: dependence on an RNA secondary structure[J].Science, 1991, 252(5014):1842-1845
[42]Liu, D.J. MBrockman,B. Dass,L. N. Hutchins,P. Singh,J. R. McCarrey,C. C. MacDonald and J. H. Graber. Systematic variation in mRNA 3'-processing signals during mouse spermatogenesis[J].Nucleic Acids Res, 2007, 35(1):234-246
[43]Liu, Y.C. Gervasi and BG. Szaro. A crucial role for hnRNP K in axon development in Xenopus laevis[J].Development, 2008, 135(18):3125-3135
[44]Liu, Y.and BG. Szaro. hnRNP K post-transcriptionally co-regulates multiple cytoskeletal genes needed for axonogenesis[J].Development, 2011, 138(14):3079-3090
[45]Liu, Y.H. Yu,SK. Deaton and B. G. Szaro. Heterogeneous nuclear ribonucleoprotein K,an RNA-binding protein,is required for optic axon regeneration in Xenopus laevis[J].J Neurosci, 2012, 32(10):3563-3574
[46]Lynch, M.L. Chen,MJ. Ravitz,S. Mehtani,K. Korenblat,M. J. Pazin and E. V. Schmidt. hnRNP K binds a core polypyrimidine element in the eukaryotic translation initiation factor 4E (eIF4E) promoter,and its regulation of eIF4E contributes to neoplastic transformation[J].Mol Cell Biol, 2005, 25(15):6436-6453
[47]Makeyev, A.V. and SA. Liebhaber. The poly(C)-binding proteins: a multiplicity of functions and a search for mechanisms[J].RNA, 2002, 8(3):265-278
[48]Messias, A.C.,CHarnisch,A. Ostareck-Lederer,M. Sattler and D. H. Ostareck. The DICE-binding activity of KH domain 3 of hnRNP K is affected by c-Src-mediated tyrosine phosphorylation[J].J Mol Biol, 2006, 361(3):470-481
[49]Meyerowitz, J., S. J. Parker, L. J. Vella, D. Ng, K. A. Price, J. R. Liddell, A. Caragounis, Q. X. Li, C. L. Masters, T. Nonaka, M. Hasegawa, M. A. Bogoyevitch, K. M. Kanninen, P. J. Crouch and A. R. White. C-Jun N-terminal kinase controls TDP-43 accumulation in stress granules induced by oxidative stress [J].[J].Mol Neurodegener, 2011, 6(6):57-57
[50]Michelotti, E.F.,GA. Michelotti,A. I. Aronsohn and D. Levens. Heterogeneous nuclear ribonucleoprotein K is a transcription factor[J].Mol Cell Biol, 1996, 16(5):2350-2360
[51]Mikula, M.A. Dzwonek,JKarczmarski,T. Rubel,M. Dadlez,L. S. Wyrwicz,K. Bomsztyk and J. Ostrowski. Landscape of the hnRNP K protein-protein interactome[J].Proteomics, 2006, 6(8):2395-2406
[52]Mikula, M.P. Gaj,KDzwonek,T. Rubel,J. Karczmarski,A. Paziewska,A. Dzwonek,P. Bragoszewski,M. Dadlez and J. Ostrowski. Comprehensive analysis of the palindromic motif TCTCGCGAGA: a regulatory element of the HNRNPK promoter[J].DNA Res, 2010, 17(4):245-260
[53]Moumen, A.C. Magill,KL. Dry and S. P. Jackson. ATM-dependent phosphorylation of heterogeneous nuclear ribonucleoprotein K promotes p53 transcriptional activation in response to DNA damage[J].Cell Cycle, 2013, 12(4):698-704
[54]Moumen, A.P. Masterson,MJ. O'Connor and S. P. Jackson. hnRNP K: an HDM2 target and transcriptional coactivator of p53 in response to DNA damage[J].Cell, 2005, 123(6):1065-1078
[55]Notari, M.P. Neviani,RSanthanam,B. W. Blaser,J. S. Chang,A. Galietta,A. E. Willis,D. C. Roy,M. A. Caligiuri,G. Marcucci and D. Perrotti. A MAPKHNRPK pathway controls BCRABL oncogenic potential by regulating MYC mRNA translation[J].Blood, 2006, 107(6):2507-2516
[56]Ostareck-Lederer, A.and DH. Ostareck. Control of mRNA translation and stability in haematopoietic cells: the function of hnRNPs K and E1E2[J].Biol Cell, 2004, 96(6):407-411
[57]Ostareck-Lederer, A.and DH. Ostareck. Precision mechanics with multifunctional tools: how hnRNP K and hnRNPs E1E2 contribute to post-transcriptional control of gene expression in hematopoiesis[J].Curr Protein Pept Sci, 2012, 13(4):391-400
[58]Ostareck-Lederer, A.D. HOstareck,C. Cans,G. Neubauer,K. Bomsztyk,G. Superti-Furga and M. W. Hentze. c-Src-mediated phosphorylation of hnRNP K drives translational activation of specifically silenced mRNAs[J].Mol Cell Biol, 2002, 22(13):4535-4543
[59]Ostareck-Lederer, A.D. HOstareck,K. P. Rucknagel,A. Schierhorn,B. Moritz,S. Huttelmaier,N. Flach,L. Handoko and E. Wahle. Asymmetric arginine dimethylation of heterogeneous nuclear ribonucleoprotein K by protein-arginine methyltransferase 1 inhibits its interaction with c-Src[J].J Biol Chem, 2006, 281(16):11115-11125
[60]Ostareck-Lederer, A.D. HOstareck,N. Standart and B. J. Thiele. Translation of 15-lipoxygenase mRNA is inhibited by a protein that binds to a repeated sequence in the 3' untranslated region[J].EMBO J, 1994, 13(6):1476-1481
[61]Ostareck, D.H.,AOstareck-Lederer,I. N. Shatsky and M. W. Hentze. Lipoxygenase mRNA silencing in erythroid differentiation: The 3'UTR regulatory complex controls 60S ribosomal subunit joining[J].Cell, 2001, 104(2):281-290
[62]Ostareck, D.H.,AOstareck-Lederer,M. Wilm,B. J. Thiele,M. Mann and M. W. Hentze. mRNA silencing in erythroid differentiation: hnRNP K and hnRNP E1 regulate 15-lipoxygenase translation from the 3' end[J].Cell, 1997, 89(4):597-606
[63]Ostrowski, J.and KBomsztyk. Nuclear shift of hnRNP K protein in neoplasms and other states of enhanced cell proliferation[J].Br J Cancer, 2003, 89(8):1493-1501
[64]Ostrowski, J.K. Klimek-Tomczak,LS. Wyrwicz,M. Mikula,D. S. Schullery and K. Bomsztyk. Heterogeneous nuclear ribonucleoprotein K enhances insulin-induced expression of mitochondrial UCP2 protein[J].J Biol Chem, 2004, 279(52):54599-54609
[65]Ostrowski, J.D. SSchullery,O. N. Denisenko,Y. Higaki,J. Watts,R. Aebersold,L. Stempka,M. Gschwendt and K. Bomsztyk. Role of tyrosine phosphorylation in the regulation of the interaction of heterogenous nuclear ribonucleoprotein K protein with its protein and RNA partners[J].J Biol Chem, 2000, 275(5):3619-3628
[66]Ostrowski, J.L. Wyrwicz,LRychlewski and K. Bomsztyk. Heterogeneous nuclear ribonucleoprotein K protein associates with multiple mitochondrial transcripts within the organelle[J].J Biol Chem, 2002, 277(8):6303-6310
[67]Pelisch, F.B. Pozzi,GRisso,M. J. Munoz and A. Srebrow. DNA damage-induced heterogeneous nuclear ribonucleoprotein K sumoylation regulates p53 transcriptional activation[J].J Biol Chem, 2012, 287(36):30789-30799
[68]Ritchie, S.F. MBoyd,J. Wong and K. Bonham. Transcription of the human c-Src promoter is dependent on Sp1,a novel pyrimidine binding factor SPy,and can be inhibited by triplex-forming oligonucleotides[J].J Biol Chem, 2000, 275(2):847-854
[69]Ritchie, S.A.,MK. Pasha,D. J. Batten,R. K. Sharma,D. J. Olson,A. R. Ross and K. Bonham. Identification of the SRC pyrimidine-binding protein (SPy) as hnRNP K: implications in the regulation of SRC1A transcription[J].Nucleic Acids Res, 2003, 31(5):1502-1513
[70]Say, E.H. GTay,Z. S. Zhao,Y. Baskaran,R. Li,L. Lim and E. Manser. A functional requirement for PAK1 binding to the KH(2) domain of the fragile X protein-related FXR1[J].Mol Cell, 2010, 38(2):236-249
[71]Schullery, D.S.,JOstrowski,O. N. Denisenko,L. Stempka,M. Shnyreva,H. Suzuki,M. Gschwendt and K. Bomsztyk. Regulated interaction of protein kinase Cdelta with the heterogeneous nuclear ribonucleoprotein K protein[J].J Biol Chem, 1999, 274(21):15101-15109
[72]Shang, E.H. DNickerson,D. Wen,X. Wang and D. J. Wolgemuth. The first bromodomain of Brdt,a testis-specific member of the BET sub-family of double-bromodomain-containing proteins,is essential for male germ cell differentiation[J].Development, 2007, 134(19):3507-3515
[73]Shnyreva, M.D. SSchullery,H. Suzuki,Y. Higaki and K. Bomsztyk. Interaction of two multifunctional proteins. Heterogeneous nuclear ribonucleoprotein K and Y-box-binding protein[J].J Biol Chem, 2000, 275(20):15498-15503
[74]Skalweit, A.A. Doller,AHuth,T. Kahne,P. B. Persson and B. J. Thiele. Posttranscriptional control of renin synthesis: identification of proteins interacting with renin mRNA 3'-untranslated region[J].Circ Res, 2003, 92(4):419-427
[75]Stanton, P.G.,PSluka,C. F. Foo,A. N. Stephens,A. I. Smith,R. I. McLachlan and L. O'Donnell. Proteomic changes in rat spermatogenesis in response to in vivo androgen manipulation; impact on meiotic cells[J].PLoS One, 2012, 7(7):e41718-e41718
[76]Takimoto, M.T. Tomonaga,MMatunis,M. Avigan,H. Krutzsch,G. Dreyfuss and D. Levens. Specific binding of heterogeneous ribonucleoprotein particle protein K to the human c-myc promoter,in vitro[J].J Biol Chem, 1993, 268(24):18249-18258
[77]Tomonaga, T.and DLevens. Heterogeneous nuclear ribonucleoprotein K is a DNA-binding transactivator[J].J Biol Chem, 1995, 270(9):4875-4881
[78]Torvund-Jensen, J., J. Steengaard, L. Reimer, L. B. Fihl and L. S. Laursen. Transport and translation of MBP mRNA is differentially regulated by distinct hnRNP proteins [J][J].J Cell Sci, 2014, 127(7):1550-1564
[79]Valverde, R.L. Edwards and LRegan. Structure and function of KH domains[J].FEBS J, 2008, 275(11):2712-2726
[80]Van Seuningen, I.J. Ostrowski,XR. Bustelo,P. R. Sleath and K. Bomsztyk. The K protein domain that recruits the interleukin 1-responsive K protein kinase lies adjacent to a cluster of c-Src and Vav SH3-binding sites. Implications that K protein acts as a docking platform[J].J Biol Chem, 1995, 270(45):26976-26985
[81]van Venrooij, W.J. and DB. Janssen. HnRNP particles[J].Mol Biol Rep, 1978, 4(1):3-8
[82]Verpoest, W.and HTournaye. ICSI: hype or hazard?[J].Hum Fertil (Camb), 2006, 9(2):81-92
[83]Wagner, S.A.,PBeli,B. T. Weinert,M. L. Nielsen,J. Cox,M. Mann and C. Choudhary. A proteome-wide,quantitative survey of in vivo ubiquitylation sites reveals widespread regulatory roles[J].Mol Cell Proteomics, 2011, 10(10):M111013284-M111013284
[84]Wang, N.P. Zhang,XGuo,Z. Zhou and J. Sha. Hnrnpk,a protein differentially expressed in immature rat ovarian development,is required for normal primordial follicle assembly and development[J].Endocrinology, 2011, 152(3):1024-1035
[85]Wyrwicz, L.S.,PGaj,M. Hoffmann,L. Rychlewski and J. Ostrowski. A common cis-element in promoters of protein synthesis and cell cycle genes[J].Acta Biochim Pol, 2007, 54(1):89-98
[86]Yano, M.H. JOkano and H. Okano. Involvement of Hu and heterogeneous nuclear ribonucleoprotein K in neuronal differentiation through p21 mRNA post-transcriptional regulation[J].J Biol Chem, 2005, 280(13):12690-12699
[87]Yoga, Y.M.,DA. Traore,M. Sidiqi,C. Szeto,N. R. Pendini,A. Barker,P. J. Leedman,J. A. Wilce and M. C. Wilce. Contribution of the first K-homology domain of poly(C)-binding protein 1 to its affinity and specificity for C-rich oligonucleotides[J].Nucleic Acids Res, 2012, 40(11):5101-5114
[88]Yu, Y.C. Zhao,ZLv,W. Chen,M. Tong,X. Guo,L. Wang,J. Liu,Z. Zhou,H. Zhu,Q. Zhou and J. Sha. Microinjection manipulation resulted in the increased apoptosis of spermatocytes in testes from intracytoplasmic sperm injection (ICSI) derived mice[J].PLoS One, 2011, 6(7):e22172-e22172
[89]Zhou, M.A. Xu,PK. Tam,K. S. Lam,B. Huang,Y. Liang,I. K. Lee,D. Wu and Y. Wang. Upregulation of UCP2 by adiponectin: the involvement of mitochondrial superoxide and hnRNP K[J].PLoS One, 2012, 7(2):e32349-e32349
