Klukovich R, Nilsson E, Sadler-Riggleman I, Beck D, Xie Y, Yan W*, Skinner MK* (2019) Environmental toxicant induced epigenetic transgenerational inheritance of prostate pathology and stromal-epithelial cell epigenome and transcriptome alterations: ancestral origins of prostate disease.Scientific Reports 18;9(1):2209. doi: 10.1038/s41598-019-38741-1

Yuan S, Liu Y, Peng H, Tang C, Hennig GW, Wang Z, Yu T, Klukovich R, Zhang Y, Zheng H, Hess RA, Xu C, Wu J, Yan W (2018) Motile cilia in the male reproductive system require miR-34/miR-449 for development and function to prevent sperm traffic jam by generating luminal turbulence.Proceedings of the National Academy of Sciences USA Published ahead of print January 18, 2019 https://doi.org/10.1073/pnas.1817018116.



Wang Z, Lee S, Oliver D, Yuan S, Tang C, Wang Y, Zheng H, Yan W (2018) Prps1l1, a testis specific gene, is dispensable for mouse spermatogenesis.Molecular Reproduction and Development 85(10): 802-804. DOI: 10.1002/mrd.23053

Nilsson E, Klukovich R, Sadler-Riggleman I, Beck D, Xie Y, Yan W*, Skinner MK* (2018) Environmental toxicant induced epigenetic transgenerational inheritance of ovarian pathology and granulosa cell epigenome and transcriptome alterations: ancestral origins of polycystic ovarian syndrome and primary ovarian insufficiency.Epigenetics 13 (8): 875-895 DOI:10.1080/15592294.2018.1521223. PMC6224216

Tang C, Klukovich R, Peng H, Wang Z, Yu T, Zhang Y, Zheng H, Klungland A, Yan W (2018) ALKBH5-dependent m6A demethylation controls splicing and stability of long 3’UTR mRNAs in male germ cells.Proceedings of the National Academy of Sciences USA 115, E325-E333 doi: 10.1073/pnas.1717794115

Barouki R, Melen E, Herceg Z, Beckers J, Chen J, Karagas M, Puga A, Xia Y, Chadwick L, Yan W, Audouze K, Slama R, Heindel JJ, Grandjean P, Kawamoto T, Nohara K (2018) Epigenetics as a mechanism linking developmental exposures to long-term toxicity.Environmental International 144: 77-86. doi.org/10.1016/j.envint.2018.02.014

Zhang Y, Zhang X, Shi J, Tuorto F, Li X, Liu Y, Liebers R, Zhang L, Qu Y, Qian J, Liu Y, Yan M, Cao Z, Lei X, Cao Y, Peng H, Zhai Q, Zhou T, Li L, Yan W, Lyko F, Zhang Y, Zhou Q, Duan E, Chen Q (2018) Dnmt2-dependent intergenerational transmission of paternally acquired metabolic disorders via sperm RNAs. Nature Cell Biology 20, 535-540.

Skinner M, Maamar MB, Sadler-Riggleman I, Klukovich R, Beck D, Xie Y, Nillson E, McBirney, Tang C, Yan W (2018) Alteration in sperm DNA methylation, noncoding RNA and histone retention associate with DD induced epigenetic transgenerational inheritance of diseaseEpigenetics and Chromatin 11:8. https://doi.org/10.1186/s13072-018-0178-0.

Grozdanov PN, Yu P, Yan W, MacDonald CC (2018) Cstf2t regulates expression of histone and histone-like proteins in male germ cells.  Andrology  doi: 10.1111/andr.12488

Tang C, Yu T, Xie Y, Wang Z, McSwiggen H, Zhang Y, Zheng H, Yan W (2018) Template switching causes artifactual junction formation and false identification of circular RNAs. Pre-print BioRxiv  doi: https://doi.org/10.1101/259556

Chuang T, Xie Y, Yan W, Khorram O (2018) Next-generation Sequencing Reveals Differentially Expressed Small Non-coding RNAs in Uterine Leiomyoma. Fertility and Sterility 109(5):919-929 doi: 10.1016/j.fertnstert.2018.01.034

Ben Maamar M, Sadler-Riggleman I, Beck D, McBirney M, Nilsson E, Klukovich R, Xie Y, Tang C, Yan W*, Skinner M* (2018) Alterations in sperm DNA Methylation, Non-Coding RNA expression, and histone retention mediate Vinclozolin induced epigenetic transgenerational inheritance of disease.  Environmental Epigenetics 4, dvy010. 


​Yu T, Tang C, Zhang Y, Zhang R, Yan W (2017) Microfluidics-based digital quantitative PCR for single cell small RNA quantification. Biology of Reproduction doi:10.1093/biolre/iox102

Yan W (2017) piRNAs-independent PIWI function in spermatogenesis and male fertility.Biology of Reproduction. doi:10.1093/biolre/iox055.

Ying Zhang, Tang C, Yu T, RR Zhang, Zheng H, Yan W (2016) MiRNAs control mRNA fate by compartmentalization based on 3’UTR length. Genome Biology In Press.   

Halim D, Wilson MP, Oliver D, Brosens E, Verheij JBGM, Han Y, Nanda V, Lyu Q, Doukas M, Stoop HJ, Brouwer RWW, van ljcken VFJ, Slivano OJ, Burns AJ, Christie CK, de Mesy Bentley KL, Brooks AS, Tibboel D, Xu S, Jin Z, Djuwantono T, Yan W, Alves MA, Hofstra RMW, Miano JM (2017) Loss of LMOD1 Impairs Cyto-contractile Coupling and Causes Megacystis Microcolon Intestinal Hypoperistalsis Syndrome in Humans and Mice.  Proceedings of the National Academy of Sciences USA doi: 10.1073/pnas.1620507114. 

Yan W* (2017) An interview with Magdalena Zernicka-Goetz.Biology of Reproduction doi:https://doi.org/10.1093/biolre/iox003.

Yan W* (2017) Mark it for destruction: A novel role of mRNA methylation in maternal-to-zygotic transition.Biology of Reproduction 10.1093/biolre/iox016 

Yanagimachi R, Harumi T, Matsubara H, Yan W, Yuan S, Yamaha E, Arai K, Hirohashi K, Tomoriha I, Matsubara T, Andoh T, Vines C, Cherr G. (2017) Chemical and Physical Guidance of Fish Spermatozoa into the Egg through the Micropyle. Biology of Reproduction  DOI:10.1093/biolre/iox015

Tang C*, Xie Y, Yan W* (2017) AASRA: An Anchor Alignment-Based Small RNA Annotation Pipeline.BioRxiv doi: https://doi.org/10.1101/132928


Chen Q*, Yan W*, Duan E* (2016) Epigenetic inheritance of acquired traits through sperm RNAs and sperm RNA modifications.Nature Reviews Genetics  doi:10.1038/nrg.2016.106. (*co-corresponding authors)

Yuan S, Tang C, Schuster A, Zhang Y, Zheng H, Yan W (2016) Paternal pachytene piRNAs are not required for fertilization, embryonic development and sperm-mediated epigenetic inheritance in mice.  Environmental Epigenetics doi:10.1093/eep/dvw021.

Fededa J, Esk C, Mierzwa B, Stanyte R, Yuan S, Zheng H, Ebnet K, Yan W, Knoblich J & Gerlich D (2016) microRNA-34/449 controls mitotic spindle orientation during mammalian cortex development.EMBO Journal doi:10.15252/embj.201694056

Schuster A, Tang C, Xie Y, Ortogero N, Yuan, S, Yan W (2016) SpermBase: a database for sperm RNA contents.Biology of Reproduction doi: 10.1095/ biolreprod.116.142190.

Zhang Y, Regalado J, Yan W (2016) Fat dad, fat kids: sperm small RNAs in control!Science Bulletin 61(6): 428-429.  doi: 10.1007/s11434-016-1031-0.

Schuster A, Skinner MK, Yan W (2016) Ancestral vinclozolin exposure alters the epigenetic transgenerational inheritance of sperm small noncoding RNAs. Environmental Epigenetics 2(1): 1-10. doi: 10.1093/eep/dvw001 dvw001.

Yuan S, Schuster AS, Tang C, Ortogero N, Bao J. Zheng H, Yan W (2016) Sperm-borne miRNAs and endo-siRNAs are important for fertilization and preimplantation development.  Development 143(4): 635-647. doi: 10.1242/dev.131755.

Bao J, Vitting-Seerup K, Waage J, Tang C, Ge Y, Porse BT, Yan W (2016) UPF2-mediated nonsense mRNA decay participates in 3'UTR shortening during male germ cell development.PLoS Genetics 12(5): e1005863. doi: 10.1371/journal.pgen.1005863.


Sosa E, Flores L, Yan W, McCarrey JR (2015) Escape of X-linked miRNA Genes from Meiotic Sex Chromosome Inactivation.Development 142(21): 3791-3800.

Yuan S, McSinggin H, Zheng H, Yan W (2015) A Testis-Specific Gene, Ubqlnl, Is Dispensable for Mouse Embryonic Development and Spermatogenesis. Molecular Reproduction and Development 82: 408-409. doi: 10.1002/mrd.22504.

Song HW, Bettegowda A, Oliver D, Yan W, Phan MH, de Rooij DG, Corbett M, Wilkinson MF. (2015) shRNA Off-Target Effects In Vivo: Impaired Endogenous siRNA Expression and Spermatogenic Defects.PLoS ONE 10(3): e0118549. doi:10.1371/journal.pone.0118549.

Oliver DK, Yuan S, Yan W (2015) Cas9/CRISPR-mediated genome editing results in pervasive genotypic mosaicism in founder mice derived from pronuclear injections. PLoS ONE 0(6): e0129457. doi:10.1371/journal.pone.0129457.

Yuan S, Qin W, Riordan CR, McSwiggin, H, Zheng H, Yan W (2015) Ubqln3, a testis-specific gene, is dispensable for embryonic development and spermatogenesis in mice. Molecular Reproduction and Development  82:266-267.

Yuan S, Oliver DK, Schuster A, Zheng H, Yan W (2015) Breeding scheme and maternal small RNAs affect the efficiency of transgenerational inheritance of a paramutation in mice.Scientific Reports 5, Article number: 9266, doi:10.1038/srep09266

Schuster A, Hennig GW, Ortogero N, Yan W (2015) In silico identification of novel endogenous small interference RNAs.Methods in Molecular Biology 1218:341-51. doi:10.1007/978-1-4939-1538-5_21. PubMed PMID: 25319662.

Ortogero N, Hennig GW, Yan W (2015) Computer-assisted annotation of small RNA transcriptomes. Methods in Molecular Biology 1218:353-64.doi:10.1007/978-1-4939-1538-5_22. PubMed PMID: 25319663.

Yuan S, Stratton CJ, Bao J, Zheng H, Bhetwal BP, Yanagimachi, R, Yan W (2015) Spata6 is required for normal assembly of the sperm connecting piece and tight head-tail conjunction.  Proceedings of the National Academy of Sciences USA 112(5): E430-E439.


Bao J, Tang C, Li J, Zhang Y, Bhetwal BP, Zheng H, Yan W (2014) RANBP9 Plays an essential role in spermatogenesis by regulating alternative splicing. PLoS Genetics 10 (12), e1004825.

Ortogero N, Hennig GW, Luong D, Bao J, Bhetwal BP, Ro S, McCarrey JR, Yan W (2014) A Novel Class of Somatic Small RNAs Similar to Germ Cell Pachytene PIWI-interacting Small RNAs. Journal of Biological Chemistry 289 (47): 32824-32834.

Yuan S, Tang C, Wu J, Zheng H, Xu C, Yan W (2014) Sperm-borne mir-34b/C and mir-449 Are Dispensable for The First Cleavage Division in Mice.  Biology Open doi: 10.1242/bio.201410959.

Bao J, Tang C, Porse BT, Yan W (2014) UPF2, a nonsense-mediated mRNA decay factor, is required for prepubertal Sertoli cell development and male fertility by ensuring transcriptomic fidelity. Development doi:10.1242/dev.115642.

Yan W (2014) Potential roles of noncoding RNAs in environmental epigenetic transgenerational inheritance.Molecular and Cellular Endocrinology 10.1016/j.mce.2014.09.008

Wu J, Bao J, Kim M, Yuan S, Tang C, Zheng H, Mastick GS, Xu C, Yan W (2014) Two miRNA clusters, miR-34b/c and miR-449, are essential for normal brain development, motile ciliogenesis and spermatogenesis. Proceedings of the National Academy of Sciences USA doi: 10.1073/pnas.1407777111; 111(28): E2851-E2857.

Yuan S, Ortogero N, Wu Q, Zheng H, Yan W (2014) Murine Follicular Development Requires Oocyte DICER, but Not DROSHA.Biology of Reproduction 114.119370.

Dong WW, Huang HL, Yang W, Liu J, Yu Y, Zhou SL, Wang W, Lv XC, Li ZY, Zhang MY, Zheng ZH, Yan W (2014) Testis-specific Fank1 knockdown in mice produces oligozoospermia. Asian Journal of Andrology 16(1): 124-30.  doi: 10.4103/1008-682X.122592.

Bao J, Zhang Y, Schuster A, Ortogero N, Nilsson EE, Skinner MK, Yan W (2014) Conditional inactivation of Miwi2 reveals that MIWI2 is only essential for prospermatogonial development in mice. Cell Death and Differentiation 21: 783-796 (May 2014) | doi:10.1038/cdd.2014.5.

Yan W (2014) Gene Knockouts That Affect Sertoli Cell Function. Chapter 15 in Sertoli Cell Biology, the 3rd Edition, edited by Michael Griswold. Academic Press, page 437.


Bao J, Wu J, Schuster AS, Hennig GW, Yan W (2013) Expression Profiling Reveals Developmentally Regulated lncRNA Repertoire in the Mouse Male Germline. Biology of Reproduction 89(5): 107, 1–12. doi: 10.1095/ biolreprod.113.113308.

Yuan S, Zheng H, Zheng Z, Yan W (2013) Proteomic Analyses Reveal a Role of Cytoplasmic Droplets as an Energy Source during Epididymal Sperm Maturation.PLoS ONE 8(10): e77466. doi:10.1371/journal.pone.0077466.

Bao J, Yuan S, Maestas A, Bhetwal B, Schuster A, Yan W (2013) Stk31 is dispensable for embryonic development and spermatogenesis in mice.  Molecular Reproduction and Development 80(10): 786.

Xu H, Yuan S, Zheng Z, Yan W (2013) The cytoplasmic droplet may be indicative of sperm motility and normal spermiogenesis.Asian Journal of Andrology 15, 799-805. doi:10.1038/aja.2013.69.

Bao J, Ma HY, Schuster A, Lin YM, Yan W (2013) Incomplete penetrance of Cre activity leads to phenotypic differences between Stra8-iCre; Mov10l1lox/lox and Stra8-iCre; Mov10l1lox/Δ mice. Genesis 51(7): 481-90. doi: 10.1002/dvg.22389.

Ro S, Ma HY, Park C, Ortogero N, Song R, Hennig GW, Zheng H, Lin YM, Moro L, Hsieh JT, Yan W (2013) The Mitochondrial Genome Encodes Abundant Small Noncoding RNAs.Cell Research 23,759-774. doi:10.1038/cr.2013.37.



Bao J, Yan W (2012) Male germline control of transposable elements.Biology of Reproduction 86 (5) 162, 1-14.

Bao J, Li D, Wang L, Wu J, Hu Y, Wang Z, Chen Y, Cao X, Jiang C, Yan W*, Xu C* (2012) microRNA-449 and microRNA-34b/c function redundantly in murine testes by targeting the E2F transcription factor-Retinoblastoma protein (E2F-pRb) pathway. Journal of Biological Chemistry 287(26): 21686-21698. (*co-corresponding author).

Wu Q, Song R, Ortogero N, Zheng H, Evanoff R, Small CL, Griswold MD, Namekawa SH, Royo H, Turner JM, Yan W (2012) The RNase III Enzyme DROSHA Is Essential for microRNA Production and Spermatogenesis. Journal of Biological Chemistry 287(30): 25173-25190.

Ortogero N, Hennig GW, Langille C, Ro S, McCarrey JR, Yan W (2012) Computer-Assisted Annotation of the Sertoli Cell Small RNA Transcriptome.Biology of Reproduction 88 (1) 3:1-10. doi:10.1095/biolreprod.112.102269

Idler, KR, Hennig, GW, Yan W (2012) Bioinformatic Identification of Novel Elements Potentially Involved in mRNA Fate Control during Spermatogenesis. Biology of Reproduction 87 (6) 138:1-10. doi:10.1095/biolreprod.112.102434.


Bao J, Zhang J, Song R, Zheng H, Yan W (2011) Ran-binding protein 17 is preferentially expressed in the testis and is associated with the XY body and manchette during spermatogenesis.Molecular and Cellular Endocrinology 333:134-142.

Park C, Yan W, Ward SM, Hwang SJ, Wu Q, Hatton W, Park J, Sanders KM, Ro S (2011) MicroRNAs dynamically remodel gastrointestinal smooth muscle cells.PLoS ONE 6(4): e18628.

Park C, Hennig GW, Sanders KM, Cho JH, Hatton WJ, Redelman D, Park JK, Ward SM, Miano JM, Yan W, Ro S (2011) SRF-dependent microRNAs regulate gastrointestinal smooth muscle cell phenotypes.  Gastroenterology 141(1): 164-175.

Song R, Hennig G, Wu Q, Jose C, Zheng H, Yan W (2011) Male germ cells express abundant endogenous siRNAs.Proceedings of the National Academy of Sciences USA 108:13159-64.

Idler RK, Yan W (2011) Control of messenger RNA fate by RNA binding proteins: an emphasis on mammalian spermatogenesis.  Journal of Andrology 33: 309-337

Bao J, Wang L, Lei J, Hu Y, Liu Y, Shen H, Yan W, Xu C (2011) STK31 (TDRD8) is dynamically regulated throughout mouse spermatogenesis and interacts with MIWI protein.  Histochemistry and Cell Biology 137(3): 377-89.



Ro S, Park C, Jin J, Zheng H, Blair P, Redelman D, Ward SM, Yan W, Sanders KM (2010) A model to study the phenotypic changes of interstitial cells of Cajal (ICC) in gastrointestinal diseases. Gastroenterology 138(3): 1068-78.e1-2.  

Ro S, Yan W (2010) Detection and quantitative analysis of small RNAs by PCR.Methods in Molecular Biology 629:295-305.

Ro S, Yan W (2010) Small RNA cloning. Methods in Molecular Biology 629:273-85.

Song R, Ro S, Yan W (2010) In situ hybridization detection of microRNAs.Methods in Molecular Biology 629:287-94.

Yan W, Slaymaker S, Si Y, Jin J, Zheng H, Aslanian A, Saunders L, Verdin E, Charo IF (2010) Zmynd15 encodes a transcription repressor essential for late spermiogenesis and male fertility.  Journal of Biological Chemistry 285:31418-31426.

Bao J, Zhang J, Zheng H, Yan W (2010) UBQLN1 interacts with SPEM1 and participates in spermiogenesis. Molecular and Cellular Endocrinology 327:89-97.


Papaioannou MD, Pitetti JL, Ro S, Park C, Aubry F, Schaad O, Vejnar CE, Descombes P, Zdobnov EM, McManus MT, Guillou F, Harfe BD, Yan W, Jégou B, Nef S (2009)  Sertoli cell Dicer is essential for spermatogenesis in mice. Developmental Biology 326:250-259

Song R, Michaels JD, Ro S, McCarrey JR, Yan W (2009)  X-linked microRNAs escape meiotic sex chromosome inactivation. Nature Genetics 41:488-493

Yan W (2009) Male Infertility caused by spermiogenic defects: Lessons from gene knockouts.Molecular and Cellular Endocrinology 306:24-32. doi:10.1016/j.mce.2009.03.003.

Ma L, Buchold GM, Greenbaum MP, Roy A, Burns KH, Zhu H, Han DY, Harris RA, Coarfa C, GunaratnePH, Yan W, Matzuk MM (2009) GASZ is Essential for Male Meiosis and Suppression of Retrotransposon Synthesis in the Male Germline. PLoS Genetics 5(9): e1000635. doi: 10.1371/journal.pgen.1000635.

Yan W, McCarrey JR (2009) Noise amidst the silence: sex chromosome inactivation in the male.Epigenetics 4(7):452-456

Zhu MH, Kim TW, Ro S, Yan W, Ward SM, Koh SD, Sanders KM (2009) A Ca(2+)-activated Cl(-) conductance in interstitial cells of Cajal linked to slow wave currents and pacemaker activity. Journal of Physiology 587:4905-4918


Yan W*, Morozumi K, Zhang J, Ro S, Park C, Yanagimachi R* (2008) Birth of mice after intracytoplasmic injection of single purified sperm nuclei and detection of messenger RNAs and microRNAs in the sperm nuclei.Biology of Reproduction 78:896-902 (*corresponding author)


Zheng H, Stratton CJ, Morozumi K, Jin J, Yanagimachi R, Yan W (2007) Lack of Spem1 causes aberrant cytoplasm removal, sperm deformation and male infertility. Proceedings of the National Academy of Sciences USA 104:6852-6857.

Jin J, Jin N, Zheng H, Ro S, Tafolla D, Sanders KM, Yan W (2007) Catsper3 and Catsper4 are essential for sperm hyperactivated motility and male fertility.Biology of Reproduction 77:37-44.

Ro S, Park C, Song R, Nguyen D, Jin J, Sanders KM, McCarrey JR, Yan W (2007) Cloning and expression profiling of testis-expressed piRNA-like RNAs.RNA 13:1693-1702.

Zheng Z, Zheng H, Yan W (2007) Fank1 is a testis-specific gene encoding a nuclear protein exclusively expressed during the transition from the meiotic to the haploid phase of spermatogenesis. Gene Expression Patterns 7:777-783.

Ro S, Park C, Young D, Sanders KM, Yan W (2007) Tissue-dependent paired expression of miRNAs.Nucleic Acids Research 35:5944-5953.

Lin YN, Roy A, Yan W, Burns KH, Matzuk MM (2007) Loss of zona pellucida binding proteins in the acrosomal matrix disrupts acrosome biogenesis and sperm morphogenesis.Molecular and Cellular Biology 27:6794-6805.

Ro S, Park C, Young D, Sanders KM, McCarrey JR, Yan W (2007)  Cloning and expression profiling of testis-expressed microRNAs.  Developmental Biology 311: 592-602.

Ro S, Song R, Park C, Zheng H, Sanders KM, Yan W (2007) Cloning and expression profiling of small RNAs expressed in the mouse ovary.  RNA  13:2366-2380.


Wang SH, Zheng HL, Esaki Y, Kelly F, Yan W (2006) Cullin3 is a KLHL10-interacting protein preferentially expressed during late spermiogenesis.  Biology of Reproduction 74:102-108.

Greenbaum MP*, Yan W*, Wu MH, Lin YN, Agno JE, Sharma M, Braun RE, Rajkovic A, Matzuk MM. (2006) TEX14 is essential for intercellular bridges and fertility in male mice.  Proceedings of the National Academy of Sciences USA 103:4982-7. (*co-first authors)

Yatsenko AN, Roy A, Chen R, Ma L, Murthy LJ, Yan W, Lamb DJ, Matzuk MM (2006) Non-invasive genetic diagnosis of male infertility using spermatozoal RNA: KLHL10 mutations in oligozoospermic patients impair homodimerization.  Human Molecular Genetics 15:3411-3419.

Ro S, Park C, Jin J, Sanders KM, Yan W (2006) A PCR-based method for detection and quantification of small RNAs.  Biochemical and Biophysical Research Communications 351:756-763.



Huang JX, Yan W, Song ZX, Qian RY, Chen P, Salminen E, Toppari J (2005) Relationship between proliferative activity of cancer cells and clinic pathological factors in patients with esophageal squamous cell carcinoma.World Journal of Gastroenterology 11:2956-295.

Yan W, Ma L, Stein P, Pangas SA, Burns KH, Bai, YC, Schultz, RM, Matzuk, MM (2005) Mice deficient in oocyte-specific oligoadenylate synthetase-like protein OAS1D display reduced fertility.Molecular and Cellular Biology 25:4615-4624.

Jin JL, O’Doherty AM, Wang SH, Zheng HL, Sanders KM, Yan W (2005) Catsper3 and Catsper4 encode two cation channel-like proteins exclusively expressed in the testis. Biology of Reproduction 73:1235-1242.



Yan W, Ma L, Zilinski CA, Matzuk MM. (2004) Identification and characterization of evolutionarily conserved pufferfish, zebrafish, and frog orthologs of GASZ. Biology of Reproduction 70:1619-1625.

Roy A, Yan W, Burns KH, Matzuk MM (2004) Tektin3 encodes an evolutionarily conserved putative testicular microtubules-related protein expressed preferentially in male germ cells.Molecular Reproduction and Development 67:295-302.

Yan W, Ma L, Matzuk MM (2004) Haploinsufficiency of KLHL10 causes male infertility.Proceedings of the National Academy of Sciences USA 101:7793-7798.

Pangas SA, Yan W, Matzuk MM, Rajkovic A (2004) Restricted germ cell expression of a gene encoding a novel mammalian HORMA domain containing protein.Gene Expression Patterns 5: 257-263.


Yan W, Burns KH, Matzuk MM (2003) Genetic engineering to study testicular tumorigenesis. APMIS 111: 174-183.

Toppari J, Suominen JS, Yan W (2003) The role of retinoblastoma protein family in the control of germ cell proliferation, differentiation, and survival.APMIS 111:245-251. 

Suzumori N, Burns KH, Yan W, Matzuk MM (2003) RFLP-4 interacts with proteins of the ubiquitin-proteosome degradation pathway in oocytes. Proceedings of the National Academy of Sciences USA 100:550-555.

Yan W, Assadi AH, Wynshaw-Boris A, Eichele G, Matzuk MM, Clark GD (2003) Previously uncharacterized role for platelet-activating factor acetylhydrolase 1b in mouse spermatogenesis.Proceedings of the National Academy of Sciences USA 100:7189-7194.

Wu MH, Rajkovic A, Burns KH, Yan W, Lin YN, Matzuk MM (2003) Sequence and expression of testis-expressed gene 14 (Tex 14): a gene encoding a protein kinase expressed exclusively during spermatogenesis.Gene Expression Patterns 3:231-236.

Yan W, Huang JX, Lax AS, Pelliniemi L, Salminen E, Poutanen M, Toppari J (2003) Overexpression of Bcl-w in the testis disrupts spermatogenesis: revelation of a role of BCL-W in male germ cell cycle control. Molecular Endocrinology 17:1868-1879.

Yan W, Santti H, Jänne OA, Palvimo JJ, Toppari J (2003) Expression of the E3 SUMO-1 ligases PIASx and PIAS1 during spermatogenesis in the rat. Gene Expression Patterns 3:301-308.

Yan W, Ma L, Burns KH, Matzuk MM (2003) HILS1 is a spermatid-specific linker histone H1-like protein implicated in chromatin remodeling during mammalian spermiogenesis.   Proceedings of the National Academy of Sciences USA 100:10546-10551.



Yan W, Rajkovic A, Viveiros MM, Burns KH, Eppig JJ, Matzuk MM (2002) Identification of Gasz, an evolutionarily conserved gene expressed exclusively in germ cells and encoding a protein with four ankyrin repeats, a sterile-alpha motif, and a basic leucine zipper. Molecular Endocrinology 16: 1168-1184.

Rajkovic A, Yan C, Yan W, Klysik M, Matzuk M (2002) Obox, a family of homeobox genes preferentially expressed in germ cells.Genomics 79: 711-717.

Yan W, Hirvonen SJ, Palvimo J, Toppari J, Janne OA (2002) Expression of the Nuclear RING Finger Protein SNURF/RNF4 during Rat Testis Development Suggests a Role in Spermatid Maturation. Mechanisms of Development 118: 247-253.

Yan W, Burns KH, Ma L, Matzuk MM (2002) Identification of Zfp393, a germ cell-specific gene encoding a novel zinc finger protein. Mechanisms of Development 118: 233-239.



Li X, Nokkala E, Yan W, Streng T, Saarinen N, Wärri A, Huhtaniemi I, Santti R, Mäkelä S, Poutanen M (2001) Altered structure and function of reproductive organs in transgenic male mice overexpressing human aromatase.Endocrinology 142:2435–2442.

Yan W, Kero J, Suominen J, Toppari J (2001) Differential expression and regulation of retinoblastoma protein (pRb) family members during testicular development and spermatogenesis: a model for the control of germ cell proliferation, differentiation and apoptosis in vivo. Oncogene 20: 1343–1356.

Suominen J, Yan W, Toppari J, Kaipia A (2001) Expression and regulation of Bcl-2-related ovarian killer (Bok) mRNA in the developing and adult rat testis.European Journal of Endocrinology 145:771-778

Blaeser F, Toppari J, Heikinheimo M, Yan W, Wallace M, Ho N, Chatila T (2001) Normal Spermatogenesis in CaMKIV/Gr-Deficient Mice.American Journal of Physiology, Endocrinology, and Metabolism 281: E931-E937.



Yan W, Suominen J, Toppari J (2000) Stem cell factor protects germ cells from apoptosis in vitro.Journal of Cell Science 113:161-168.

Yan W, Samson M, Jégou B, Toppari J (2000) Bcl-w forms complexes with Bax and Bak and elevated ratios of Bax/Bcl-w and Bak/Bcl-w correspond to spermatogonial and spermatocyte apoptosis in the testis.Molecular Endocrinology 14:682-699.

Zheng H, Yan W, Toppari J, Härkönen P (2000) Improved non-radioactive RT-PCR method for relative quantification of mRNA. BioTechniques 28:832-834.

Yan W, Kero J, Huhtaniemi I, Toppari J (2000) Stem cell factor functions as a survival factor for mature Leydig cells and a growth factor for precursor Leydig cells after ethylene dimethane sulfonate treatment: implication of the role of stem cell factor/c-kit interaction in Leydig cell development. Developmental Biology 227:169-182.

Yan W, Suominen J, Samson M, Jégou B, Toppari J (2000) Involvement of Bcl-2 family proteins in apoptosis during testicular development and pro-survival effect of stem cell factor on germ cells in vitro. Molecular and Cellular Endocrinology 165:115 – 129.


1999 and prior

Tena-Sempere M, Kero J, Rannikko A, Yan W, Huhtaniemi I (1999) The pattern of inhibin/activin a and bB subunit mRNA expression in rat testis after selective Leydig cell destruction by ethylene dimethane sulfonate.Endocrinology 140: 5761-5770.

Schultz R, Yan W, Toppari J, Völkl A, Gustafsson J, Huikko MP (1999) Expression of peroxisome proliferator-activated receptor a mRNA and protein in human and rat testis. Endocrinology 140:2968-2975.

Moilanen AM, Karvonen U, Poukka H, Yan W, Toppari J, Jänne OA, Palvimo J (1999) A testis-specific androgen receptor coregulator that belongs to a novel family of nuclear proteins. Journal of Biological Chemistry 274:3700-3704.

Yan W, Linderborg J, Suominen J, Toppari J (1999) Stage-specific regulation of stem cell factor (SCF) gene expression in the rat seminiferous epithelium. Endocrinology 140:1499-1504.

Hakovirta H*, Yan W*, Kaleva M, Zhang F, Vänttinen K, Morris PL, Söder O, Parvinen M, Toppari J (1999) Function of stem cell factor as a survival factor of spermatogonia and localization of messenger ribonucleic acid in the rat seminiferous epithelium.Endocrinology 140:1492-1498 (*co-first authors).

Nata M, Kimura T, Hashiyada M, He P, Yan W, Li X, Funayama M, Sagisaka K (1999) Allele frequencies of eight STRs in Japanese and Chinese. International Journal of Legal Medicine 112:396-399.

Yan W, West A, Toppari J, Lähdetie J (1997) Stage-specific expression and phosphorylation of retinoblastoma protein (pRb) in the rat seminiferous epithelium.Molecular and Cellular Endocrinology 132:137-148.

He P, Nata M, Kanetake J, Ji G, Yan W, Li X, Sagisaka K (1997) Polymorphism of short tandem repeat (STR) loci PLA2, D3S2459, D8S315 (kw38), CYP19, D3S1359, FGA, and D8S1132 in the Japanese and Chinese populations. Tohoku Journal of Experimental Medicine 183:251-262

Sun YX, Zhao GC, Yan W (1995) Age estimation on the female sternum by quantification theory and stepwise regression analysis.Forensic Science International 74:57-62

Jiang XH, Yan W (1993) Parentage determination of placental tissues through DNA fingerprints.Annals of Academy of Medicine 22:8-10.

University of Nevada, Reno School of Medicine

University of Nevada, Reno School of Medicine