Innovative use of forensic STRs for authentication of human cell lines stored over 34 years

Abstract

Aim: In forensic science, the authentication of biological samples is critical for accurate analysis and evidence validation. To evaluate the authenticity and genetic stability of 91 long-term preserved human cell line samples under cryogenic conditions over 34 years, and to explore the applicability of forensic-grade short tandem repeat (STR) profiling for long-term cell line authentication.

Methods: We conducted STR-based authentication using 23 forensic STR markers. STR profiles were analyzed using the Tanabe and Masters algorithms for authentication. The study also examined the effects of prolonged passaging and genetic modification on STR stability, and investigated potential contamination events by comparing profiles with public databases.

Results: All uniquely labeled human cell lines were successfully revived and yielded complete STR profiles, confirming the efficacy of long-term cryopreservation. One male cell line showed the Y-indel in the absence of a Y-allele at Amelogenin, indicating that the Y-indel alone is insufficient for accurate sex determination. Authentication using the Tanabe and Masters algorithms revealed that stringent criteria provided more definitive results than scoring-based methods. Instances of relatedness among unauthenticated cell lines to problematic references in existing databases suggest early-stage contamination rather than recent laboratory errors. STR profile stability persisted through routine cell line passages, and genetically edited cell lines retained profiles identical to their parental counterparts, indicating a need for additional differentiation markers for accurate derivation authentication. No significant differences in the number of altered alleles between normal and tumor cell lines, implying cell type alone did not dictate STR stability.

Conclusion: This study represented one of the most extensive single-laboratory investigations into cell line preservation using forensic-grade tools, providing new reference alleles and valuable insights into cell line authentication. The findings supported the application of forensic STR kits beyond traditional forensic samples, offering a new perspective on their utility in genetic research and laboratory management.

Keywords

Cell line authentication / human cell lines / HeLa contamination / STR profiling / STR status

References

[1] Kaplan J.Cell line characterization and authentication. animal cell culture methods.Methods Cell Biol1998;57:203-16

[2] Corral-Vázquez C,Catalina P.Cell lines authentication and mycoplasma detection as minimun quality control of cell lines in biobanking.Cell Tissue Bank2017;18:271-80

[3] Masters JR.Cell-line authentication: End the scandal of false cell lines.Nature2012;492:186

[4] Nelson-Rees WA,Flandermeyer RR.Cross-contamination of cells in culture.Science1981;212:446-52

[5] Castro F,Fähnrich S,Pawlita M.High-throughput SNP-based authentication of human cell lines.Int J Cancer2013;132:308-14 PMCID:PMC3492511

[6] Lung O,Nebroski M,Buchanan C.High-throughput sequencing for species authentication and contamination detection of 63 cell lines.Sci Rep2021;11:21657 PMCID:PMC8569163

[7] Capes-Davis A,Kline MC.Match criteria for human cell line authentication: where do we draw the line?.Int J Cancer2013;132:2510-9

[8] Almeida JL,Plant AL.Standards for Cell Line Authentication and Beyond.PLoS Biol2016;14:e1002476 PMCID:PMC4907466

[9] Zheng H,Zhang J.Development and validation of a novel SiFaSTR^TM 23-plex system.Electrophoresis2019;40:2644-54

[10] Huang Y,Liu C.Comprehensive landscape of non-CODIS STRs in global populations provides new insights into challenging DNA profiles.Forensic Sci Int Genet2024;70:103010

[11] Chen A,Li J,Chen J.Detecting genetic hypermutability of gastrointestinal tumor by using a forensic STR kit.Front Med2020;14:101-11

[12] Chen A,Qu Y.Opportunity of Next-Generation Sequencing-Based Short Tandem Repeat System for Tumor Source Identification.Front Oncol2022;12:800028 PMCID:PMC8873583

[13] Almeida JL. Authentication of human and mouse cell lines by short tandem repeat (STR) DNA genotype analysis. In: Markossian S, Grossman A, Baskir H, et al., editors. Assay Guidance Manual [Internet]. Bethesda (MD): Eli Lilly & Company and the National Center for Advancing Translational Sciences;2004. Available from: https://www.ncbi.nlm.nih.gov/books/NBK144066/ [Last accessed on 30 June 2025]

[14] Masters JR,Daly-Burns B.Short tandem repeat profiling provides an international reference standard for human cell lines.Proc Natl Acad Sci U S A2001;98:8012-7 PMCID:PMC35459

[15] Korch CT,Dirks WG. Human cell line authentication: standardization of short tandem repeat (STR) profiling. ASN-0002 Revised 2021, by American National Standards Institute - American Type Culture Collection Standards Development Organization. Available from: https://www.atcc.org/products/135-ansi-str. [Last accessed on 30 June 2025]

[16] CLASTR. CSSS, Interpretation of results. Available from: https://www.cellosaurus.org/str-search/help.html. [Last accessed on 20 Jun 2025]

[17] Tian L,Xue JL.Four microRNAs promote prostate cell proliferation with regulation of PTEN and its downstream signals in vitro.PLoS One2013;8:e75885 PMCID:PMC3787937

[18] Wang BY,Zhou CH.ΦC31 integrase interacts with TTRAP and inhibitsNFκB activation.Mol Biol Rep2010;37:2809-16

[19] Yang X,Wang Y,Qiu X.Constitutive expression of human coagulating factor IX in HeLa cells by homologous recombination of the promoter.Sci China C Life Sci2001;44:18-24

[20] Lai L,Wang J.Skeletal muscle-specific expression of human blood coagulation factor IX rescues factor IX deficiency mouse by AAV-mediated gene transfer.Sci China C Life Sci1999;42:628-34

[21] Zhou C,Zhu Q,Xue J.Effects of CIITA antisense RNA on the expression of HLA class II molecules.Chin Sci Bull2000;45:2068-71

[22] Chen L,Zou B.Preparation of rAAV/hFlX by HSV/AAV hybrid helper virus and evaluation of its safety.Chin Sci Bull48, 2003, 1369-1374

[23] Dong X,Wang G.Establishment of an AAV reverse infection-based array.PLoS One2010;5:e13479 PMCID:PMC2957432

[24] Lu H,Wang X,Qiu X.Preparation of a recombinant adeno-associated viral vector with a mutation of human factor IX in large scale and its expressionin vitro andin vivo.Chin Sci Bull2001;46:1367-71

[25] Lu DR,Cong S,Qiu XF. Detection and safety of wild-type adenovirus in HSV-tk/NAS system gene therapy for brain tumors. Chin J Virol. 1998;11-16. (in Chinese) Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbfl82koZgsr3yGgO4S9-Rl5duqPi3UBJxok1tja4GZNFsUrq0uc8-fmkk1-7EYknlfP1JmJMxl8wn0sdRh1TYtU64sH-tRGPrgFIxVE7UYJ_DehY4EpS_5-Bfkq5JyNmJ1lFs-0dwViKSclRG33UI7kI0LwOqDyzxI=&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[26] Wang R,Zhou CH,Ji CN.Cdc20 mediates D-box-dependent degradation of Sp100.Biochem Biophys Res Commun2011;415:702-6

[27] Zhou C,Xue J,Chen J.Overexpression of TTRAP inhibits cell growth and induces apoptosis in osteosarcoma cells.BMB Rep2013;46:113-118 PMCID:PMC4133851

[28] Wang J,Wang R,Chen J.Adeno-associated virus Rep78 restricts adenovirus E1B55K-mediated p53 nuclear exportation.Acta Biochim Biophys Sin2013;45:135-40

[29] Ni J,Xue JL.Effects of -epigallocatechin gallate on expressions of hMLH1 and hMSH2.Carcinog Teratog Mutagen2013;25:276-79

[30] Ni J,Cao N,Xue JL. Effects of folate with different oxidative states on genomic DNA methylation in human lymphoblast cell line. Cancer Res Prev Trea. 2014;41:256-9. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbfDq3KdyoHgF7ndvJCe5hp-HCuFyw3heifGw6_zO6QhZLCjK7CufQ67QCaDrwJIR29VhdLcNuq0WHwKuiQkkK-5MNPIdEIhuDnGxqYjDA_IFEOmed-GzkH5BWFVmXNmKoH9tjxgF_uTJEGyjEDyNec-TxWv7KRhIUPQiPEp7WhDqA==&uniplatform=NZKPT&language=CHS [Last accessed on 30 June 2025]

[31] Fang YX,Wei W.Development of chimeric gene regulators for cancer-specific gene therapy with both transcriptional and translational targeting.Mol Biotechnol2010;45:71-81

[32] Li Z,Chen F,Meng L.Identification of Key lncRNA-mRNA Pairs and Functional lncRNAs in Breast Cancer by Integrative Analysis of TCGA Data.Front Genet2021;12:709514

[33] Zhu H,Li F,Xue J.Expression of human clotting factor IX mediated by recombinant lentiviral vector in cultured cells and hemophilia B mice.Chin Sci Bull2003;48:2196-200

[34] Lin Y,Wang JJ,Shen Q,Xue JL.Sp100 interacts with phage ΦC31 integrase to inhibit its recombination activity.Acta Biochim Pol2011;58:67-73

[35] Zhou GY,Shen Q,Chen JZ. FHC interacts with bim and protects cell from apoptosis. Prog Biochem Biophys. 2011;38:859-65. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbdyQhxhTYrDqwl9EC4vM4CcXNiG7g_dJucj4hsvk_TNQlVD6FEgjGthmt4qzBdJGdi2Ee0ki_Vsg6KBQA31P2-KcQxM_iCJ7F5bzBZIntu-gT7n_Y7HrBKUM8H7gZv7rwlr2pLo48ulqQjqgp2Y45zZlLoCL4ZmyTKnmv0Oz7uMoA==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[36] Yue YB,Tian L,Chen JZ.Functional differentiation between Rep-mediated site-specific integration and transcriptional repression of the adeno-associated viral p5 promoter.Hum Gene Ther2010;21:728-38

[37] Zhang JQ,Li WJ.Cellular protein TTRAP interacts with HIV-1 integrase to facilitate viral integration.Biochem Biophys Res Commun2009;387:256-60

[38] Xu GL,Wang BY.TTRAP is a novel PML nuclear bodies-associated protein.Biochem Biophys Res Commun2008;375:395-8

[39] Chen JZ,Xu GL.DAXX interacts with phage ΦC31 integrase and inhibits recombination.Nucleic Acids Res2006;34:6298-304

[40] Feng D,Yue Y,Xue J.A 16bp Rep binding element is sufficient for mediating Rep-dependent integration into AAVS1.J Mol Biol2006;358:38-45

[41] Chen JZ,Ji CN,Xie Y.Identification, expression pattern, and subcellular location of human RIP isoforms.DNA Cell Biol2005;24:464-9

[42] Chen JZ,Gu SH.Over-expression of Bim alpha3, a novel isoform of human Bim, result in cell apoptosis.Int J Biochem Cell Biol2004;36:1554-61

[43] Zhang J,Liu L.Different gene expression profiles of AD293 and HEK293 cell lines that show contrasting susceptibility to apoptosis induced by overexpression of Bim L.Acta Biochim Pol2006;53:525-30

[44] Xue YY,Yue YB,Chen JZ.Role and fate of SP100 protein in response to Rep-dependent nonviral integration system.Appl Microbiol Biotechnol2013;97:1141-7

[45] Wang Q,Xing Y,Qiu X.Suicide effect on rat gliomas mediated by recombinant adenovirus thymidine kinase/acyclovir system.Chin Med J1998;111:483-7

[46] Zhang S,Li W.MIR100HG regulates CALD1 gene expression by targeting miR-142-5p to affect the progression of bladder cancer cells in vitro, as revealed by transcriptome sequencing.Front Mol Biosci2021;8:793493

[47] Wei W,Fang YX,Guo SR. Biological evaluation on oncolytic herpes simplex virus with PEGylated folate modification. Biotechnol Bull. 2010;201-7. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbeox2SlQvpynyi47ShVMOOIks5U4vaK4fXjoTybTHvCQ0jGlNsQARqfxJiseMCxqr2HnrDkeRsb-LYNKkV6dEXgsU6h0KAzDX63IO7aHHFS3NdbeY6KlReKUz1SY5516mRXx_T85uDfGMgELa2WICKK8hDU8c6px150_30N-Y7lpQ==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[48] Wang H,Wang X et al. Efficient transfer and expression of human clotting factor IX cDNA in neonatal hemophilia B mice mediated by VSV-G pseudotyped retrovi-rus. Chin Sci Bull. 2001;1534-8. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbctf_3kRHESrWU_41LerqhbRNaEkLYHv2AKYryJId3UO7dqERobgw2z2VeEegYDC7-QEZprQaIHtcezHmV2IAp_8BwT8jtmT4d7VHMBesQ6_TGmVP0KZ0Db3IAiLQ-Y4839zg8WMaLkQlZFPmUmpxWtML5CyIlsFzUGMyLrG27Org==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[49] Wang F,Lu DR,Zhang X.Vesicular stomatitis virus G-protein retrovector mediated a herpes simplex virus thymidine kinase gene transduction and expression in the human retinal pigment epithelial cells.Chin J Ophthalmol2003;39:201-5

[50] Li K,Zhou CH,Ji CN. Cdh1 mediate D-box-dependent degradation of Sp100. J Fudan Univ. (Nat Sci.) 2012;51:315-21. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbf1nEqKIsr6SXzUhyEZTVmq63kajmFNNIfgBV2dlAEbCrOHFgGTGKorFH3Bty5aPjZhnjjnhdyMRhQPdoa80fZdXFKG1940IoB7h01QYoKOeDC_ojt8IZ7LoD62UYZSXdDNoALop23Fr1buh_Gbnto69yKT8D5GlBMt5iaarYQKyQ==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[51] Li WJ,Zhang JQ.The 156KELK159 tetrapeptide of HIV-1 integrase is critical for lentiviral gene integration.Mol Biol Rep2012;39:343-9

[52] Huang L,Zhang JQ.Daxx interacts with HIV-1 integrase and inhibits lentiviral gene expression.Biochem Biophys Res Commun2008;373:241-5

[53] Chen H,Huang L,Jia W.Expression of human factor IX gene in murine plasma through lentiviral vector-infected haematopoietic stem cells.Clin Exp Pharmacol Physiol2006;33:1196-201

[54] Yao H,Huang L,Jia W.Isolation and cultivation of murine hematopoietic stem cells and expression of hFIX mediated by recombinant lentiviral vectors in vitro.Front Biol China2006;1:259-62

[55] Chen XG,Gong JL,Xue JL. Efficient delivery of human clotting factor IX after injection of lentiviral vectors in utero. Acta Pharmacol Sin. 2004;25:789-93. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbdNfbJg6UjHCpmqw4Yzcg-Ny2wcqBTxbFmi6ZIWSJTFSQtfKFa-b8xLlI1xL70E_-SHJS5z4XjhlXotVxUzis2lbDpBMXVjw9mfSmgPsiJqGmAlPtkWDzT0lWudquZzpn9tjUz-KLzS6xpcVFlosjbrpVTk4V0KJENCxfd77oKefA==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[56] Chen XG,Yao JH. Gene delivery to mice spermatogenic stem cells by ffectene. Chin Med J. 2004;117:630-2. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbfpD3gcspvIZK2xu2Tag3j0ZPJGqb-YWMso3G7VS2F0km21dXUQgUZbxVzpqjKY53rCtoTafqQolmrzlnrMJ5x2WARVvaAidKd-3WLJgFp9zjXvEXSl3eVZb-Lssjq_lqWmTNCM5EX1wE05t7U4WDo2h1WRMPKeKpwHkK-f65mCxg==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[57] Li Z,Wang J.Evaluation of the interactions of HIV-1 integrase with small ubiquitin-like modifiers and their conjugation enzyme Ubc9.Int J Mol Med2012;30:1053-60

[58] Wang JJ,Li W,Chen JZ.Daxx interacts with phage ΦBT1 integrase and inhibits its recombination.Prog Biochem Biophys2013;40:37-42

[59] Fang Y,Shen Q,Tian L.MicroRNA-7 inhibits tumor growth and metastasis by targeting the phosphoinositide 3-kinase/Akt pathway in hepatocellular carcinoma.Hepatology2012;55:1852-62

[60] Chen L,Li G,Xue J.Novel rAAV production system with low contamination of helper virus.Chin Sci Bull2003;48:472-5

[61] Gao X,Shi D,Qiu X.A study of gene transfer and expression of human clotting factor IX in hemophilia B mice mediated by mini-adenoviral vector.Sci China C Life Sci2003;46:631-40

[62] Gao XB,Wang F,Qiu XF.In vitro expression of human clotting factor IX minigene mediated by mini adenoviral vector.Chin J Virol2000;294-8.Available from: https://europepmc.org/article/cba/341696. [Last accessed on 30 June 2025]

[63] Zhao Y,Miao N.Use of miRNA sequencing to reveal Hub miRNAs and the effect of miR-582-3p/SMAD2 in the progression of hepatocellular carcinoma.Front Genet2022;13:819553 PMCID:PMC8977860

[64] Zhu P,Yu J.High-throughput sequencing approach for the identification of lncRNA biomarkers in hepatocellular carcinoma and revealing the effect of ZFAS1/miR-150-5p on hepatocellular carcinoma progression.PeerJ2023;11:e14891

[65] Hsueh JL.Clinical protocol of human gene transfer for hemophilia B.Hum Gene Ther1992;3:543-52

[66] Dai YF,Xue JL. High efficient transfer and expression of human clotting factor IX cDNA in cultured human primary skin fibroblasts from hemophilia B patient by retroviral vectors. Sci China B. 1992;35:183-93. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbc49TnNHmygfQ-0kMSwAw-Vm8l7rPzp9EzH0k8-VE9RCt9CgwThZTiTsFNXn4XqcrF5GF04PAE0ARIkxPRu_WOb50JXDI01vtK3FZ9N4r2S7JmGiHHXL1CdjWzgArVhdu4Jd4FPs_X6jyFmBGn1QaK-GI1dkJ9kPlr0SNpfU_G2sQ==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[67] Zhou JM,Qiu XF,Akira Y.Expression of human factor IX cDNA in mice by implants of genetically modified skin fibroblasts from a hemophilia B patient.Sci China B1993;36:1082-92

[68] Wang HW,Xue JL. Expression of human clotting factor IX with EBV shuttle vector. Chin Sci Bull. 1995;1829-33. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbeNi9W5FuF6GkEv1S8C3wWV7IA6mRubAL67_i1cub43knocA2yJfoD6KJtGOQctxuOIRuTPNwUuJ8MogmBuEKyiI45wzEQgUN1PmGpNqp_Ll-zGGzbjcYxjOyx5O38a5PxilBAU6PQexb7bcVSoUijV9KHNORWLvH5Iy5kKbdWA_g==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[69] Wang JS,Chen ZX,Xue JL. Construction of a polycistronic retroviral vector for multidrug resistance genes and transfection of human umbilical cord blood CD34+ cells. Natl Med J China. 2000;77-79. (in Chinese) Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbfdMMmeCA3PFNLmuvfhcFD0vcbIvBa_sTbbSzR-ehsGVVWPxKfJxdW2fTP1-F_FbmixyhovN_CIrrKYT4wnSrMc1adI6MOF-oY2NZapMfY-9VUnOCj1mULfVPNxYzD5rpff2tOuPvkJoHsJTyMBHH10D4bAlIRVx80C_TOVmQFLLQ==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[70] Lu DR,Zheng B,Qiu XF. Regulation of in vitro expression of human coagulation factor IX cDNA. Prog Nat Sci. 1998;43-48. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbcoh0njtXqbn4MdqnUMdbaEyKBmW07gZ0398eL9P6t5a_Ee710ndFwpH0WA847E1hYtOThwiNZWfjxWSaHU0b5190nbIe5Y7gFHg_iHZStgealAgEhUUGx5Eg9fpQsXFry7tPtbgCPkNDIqBby3lT-5eocC26RCrrmupBrRAuKCsQ==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[71] Lu DR,Zheng B.Stage I clinical trial of gene therapy for hemophilia B.Sci China B1993;36:1342-51

[72] Wang P,Qiu XF. Efficient expression of human factor IX cDNA in human cells under the control of the human cytomegalovirus promoter. Chin Sci Bull. 1991;36:1021-3. (in Chinese) Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbd4eBgXnVpqMKWKZoQTt4DLbRwJp_5k9IDOY1ZSjU3fp0VVqzMTYIbF7amGWgBHj-Uf2MWPxnrWvtduKnaRCHFJ5iPEoaD01eSE9bSbM36bCWoJ_SFUVuRXMHmPaoT0W7-QasoI9135yqHeNbH86IP1Aj4Lj5nDinibd3pxjvvdhA==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[73] Zhou JM,Hu YP,Xue JL. Construction of recombinant plasmid containing human factor Ⅸ cDNA and its expression by transfected cells in mice. Chin Sci Bull. 1993;956-60. Available from: https://kns.cnki.net/kcms2/article/abstract?v=UThtwiquHbeE5zJ5jATPlJl-HWrUPg0jWykVU0Ig5AqMjWC6UkfMyxQX8tQRIJm7Nnf70H42OiQCzySbiBfU6YsWGuKEknRC4rQhd2kj2-tsyIhDLmyiZQyU2Luo--ONpNheubkXysptvey1THO_zwJryE_VjEmZNXGpSvvcOU2Gv8buYt4_MA==&uniplatform=NZKPT&language=CHS. [Last accessed on 30 June 2025]

[74] Hu Y,Xue J.Regulation of human clotting factor IX cDNA expression in transgenic mice.Sci China B1995;38:825-33

[75] Souren NY,Heck S.Cell line authentication: a necessity for reproducible biomedical research.EMBO J2022;41:e111307

[76] Freedman LP,Ethier SP,Neve RM.Reproducibility: changing the policies and culture of cell line authentication.Nat Methods2015;12:493-7

[77] Takayama T.A novel mutation at the AMEL primer binding region on the Y chromosome in AMELY negative male.Int J Legal Med2022;136:519-26

[78] and Let Live: The remarkable story of HEK293 Cells.Hum Gene Ther2020;31:485-7

[79] Hayflick L.The limited in vitro lifetime of human diploid cell strains.Exp Cell Res1965;37:614-36

[80] Macleod RA,Matsuo Y,Milch H.Widespread intraspecies cross-contamination of human tumor cell lines arising at source.Int J Cancer1999;83:555-63

[81] Lucey BP,Hutchins GM.Henrietta lacks, hela cells, and cell culture contamination.Arch Pathol Lab Med2009;133:1463-7

[82] Lin LC,Ramamoorthi M.Cross-contamination of the human salivary gland HSG cell line with HeLa cells: a STR analysis study.Oral Dis2018;24:1477-83

[83] Huang Y,Zheng C.Investigation of Cross-contamination and misidentification of 278 widely used tumor cell lines.PLoS One2017;12:e0170384 PMCID:PMC5249119

[84] Wahls WP,Moore PD.Hypervariable minisatellite DNA is a hotspot for homologous recombination in human cells.Cell1990;60:95-103

[85] Liu Y,Wu Q.Short tandem repeats of human genome are intrinsically unstable in cultured cells in vivo.Gene2023;877:147539

[86] Press MO,Queitsch C.The overdue promise of short tandem repeat variation for heritability.Trends Genet2014;30:504-12 PMCID:PMC4254273

[87] Gymrek M,Guilmatre A.Abundant contribution of short tandem repeats to gene expression variation in humans.Nat Genet2016;48:22-9

[88] Rubocki R,Shepard K,Shepherd S.Loss of heterozygosity detected in a short tandem repeat (STR) locus commonly used for human DNA identification.J Forensic Sci2000;45:1087-9