Wansheng Liu

Wansheng Liu

  • Professor of Animal Genomics
  • Undergraduate Adviser
  • Graduate Faculty in Animal Science, Bioinformatics and Genomics (BG), and Molecular, Cellular and Integrative Biosciences (MCIBS) programs.
311 Animal, Veterinary and Biomedical Sciences Building
University Park, PA 16802

Areas of Expertise

  • Structural, functional and comparative genomics
  • Animal breeding and genetics
  • Epigenetics/epigenomics
  • Mammalian sex (X and Y) chromosome and evolution
  • Ruminant Y-linked genes and male fertility
  • Testis development and spermatogenesis


  • B.S., 1982, Animal Science, Northwest Agricultural University, China.
  • M.S., 1987, Animal Breeding and Genetics, Northwest Agricultural University, China.
  • Ph.D., 1993, Animal Breeding and Genetics, Northwest Agricultural University, China.
  • Post-Doctorate, 1996-1999, Department of Genetics, Uppsala University, Sweden.
  • Post-Doctorate, 1999-2003, Department of Animal Science, University of Minnesota.

Professional Experience

  • Professor (2018‐present) and Associate Professor (2007-2017): Department of Animal Science, Pennsylvania State University.
  • Associate Research Professor (2003‐2007): Department of Animal Biotechnology, University of Nevada, Reno.
  • Research Associate (1999‐2003): Department of Animal Science,University of Minnesota.
  • Researcher & Research Associate (1997‐1999), Department of Genetics, Uppsala University, Sweden. Department of Cell and Molecular Biology (CMB), Medical Nobel Institute, Karolinska Institute, Stockholm, Sweden. Department of Medicine, University of California, San Francisco (UCSF).
  • Visiting Scholar (1995‐1996): Department of Animal Breeding and Genetics, Swedish University of Agricultural Sciences, Uppsala, Sweden.
  • Associate Professor (1994‐1996), Institute of Developmental Biology, Department of Biology, Shandong University, Jinan, Shandong, China.
  • Associate Professor (1992‐1993): Department of Animal Science, Northwest Agricultural University, Shaanxi, China.
  • Lecturer (1987‐1991): Department of Animal Science, Northwest Agricultural University, Shaanxi, China.
  • Assistant Professor (1982‐1986): Department of Animal Science, Northwest Agricultural University, Shaanxi, China.


My research interests are in the areas of animal genetics, structural, functional and comparative genomics, mammalian Y‐chromosome, and male fertility. My current research projects are:

  • Functional and structural characterization of the ruminant Y‐chromosome, with a focus on Y gene content, gene expression and function
  • Characterization of Y-chromosome variations and their application in male health and reproduction in cattle and other livestock species
  • Functional characterization of the PRAME/PRAMEY gene family during spermatogenesis in cattle and mice.


  • ANSC 543 Animal Genomics 
  • ANSC 213 Introduction to Animal Biotechnology

Selected Publications

Since 2010 (for publications before 2010, please visit https://www.researchgate.net/profile/Wansheng_Liu)

  • Beiki, H., Murdoch, B.M., Park, C.A., Kern, C.H., Kontechy, D., Becker, G., Rincon, G., Jiang, H., Zhou, H., Thorne, J., Koltes, J.E., Michal, J.J., Davenport, K., Rijnkels, M., Ross, P.J., Hu, R.,  Corum, S., McKay, S., Smith, T.P.L., Liu, W.-S., Ma, W., Zhang, X., Xu, X., Han, X., Jiang, Z., Hu, Z.-L., Reecy, J.M. (2024)  Enhanced Bovine Genome Annotation Through Integration of Transcriptomics and Epi-Genetics Datasets Facilitates Genomic Biology. GigaScience 13, 1–21. DOI: https://orcid.org/0000-0003-4602-0990
  • Yang, M., Diaz, F., Krause, A.R.T., Lei, Y., Liu, W.-S. (2024) Synergistic enhancement of the mouse Pramex1 and Pramel1 in repressing retinoic acid (RA) signaling during gametogenesis. Cell & Bioscience 14(1):28. DOI: https://doi.org/10.1186/s13578-024-01212-w
  • Yang, M.Y., Ma, W.Z., Oatley, J., and Liu, W.-S. (2023) The mouse Pramel1 regulates spermatogonial development by inhibiting the retinoic acid (RA) signaling during spermatogenesis. Development  150, 1-15. https://doi.org/10.1242/dev.201907
  • Wang, J., Hua, G., Chen, J., Cui, K., Yang, Z., Han, D., Yang, X., Dong, X., Ma, Y, Cai, G., Zhang, Y., Li, J., Tai, Y., Da, L., Li, X., Ma, L., Ma, Q., Li, R., Liu, J., Darwish, H.Y.A., Wu, K., Rong, W., Liu, W.-S., Zhao, Y., Deng, X. (2023) Epigenetic mechanism of Gtl2-miRNAs causes the primitive sheep characteristics found in purebred Merino sheep. Cell & Bioscience 13, 190. DOI: https://doi.org/10.1186/s13578-023-01142-z
  • Kern, C, Lu, C., Wu, W.W., Zhang, J.B., Zhao, Y.Q., Ocon-Grove, O.M., Sutovsky, P., Diaz, F., Liu, W.-S. (2023) Role of the bovine PRAMEY protein in sperm function during in vitro fertilization (IVF). Cell and Tissue Research 391, 577–594. DOI: https://doi.org/10.1007/s00441-022-03717-7  
  • Kern, C.H., Feitosa, W.B., and Liu, W.-S. (2022) The dynamic of PRAMEY isoforms in testis and epididymis suggests their involvement in spermatozoa maturation. Frontiers in Genetics 13, 846345. DOI: https://doi.org/10.3389/fgene.2022.846345
  • Liu, W.-S. Lu, C., Mistry, B. V. (2021) Subcellular localization of the mouse PRAMEL1 and PRAMEX1 reveals multifaceted roles in the nucleus and cytoplasm of germ cells during spermatogenesis. Cell & Bioscience 11, 102. DOI: 10.1186/s13578-021-00612-6
  • Kern, C., Yang, M.Y., and Liu, W.-S. (2021) The PRAME family of cancer testis antigens is essential for germline development and gametogenesis. Biology of Reproduction, 105, 290-304. DOI: 10.1093/biolre/ioab074
  • Lu, C., Yang, M., Rossi, R.M., Wang, A., Feitosa, W.B., Diaz, F. J., Liu, W.-S. (2020) Deletion of the mouse X-linked Prame gene causes germ cell reduction in spermatogenesis. Mol. Reprod. Dev. 87(8), 666-679. DOI: 10.1002/mrd.23324
  • Yao, Y., Zhang, Y., Liu, W.-S., Deng, X. (2020) Highly efficient synchronization of sheep skin fibroblasts at G2/M phase and isolation of sheep Y chromosomes by flow cytometric sorting. Scientific Reports 10:9933. DOI:10.1038/s41598-020-66905-x
  • Chen, X., Zheng, Y.,  Lei, A., Zhang, H., Niu, H., Li, X., Zhang, P., Liao, M., Lv, Y., Zhu, Z., Pan, C., Dong, W., Chen, H., Wu, D., Liu, W.-S., Hamer, G., Zeng, S., Zeng, W. (2020) Early cleavage of preimplantation embryos is regulated by tRNAGln-TTG-derived small RNAs present in mature spermatozoa. J. Biol. Chem. 295 (32), 10885-10900. jbc.RA120.013003.
  • Dechow, C.D., Liu, W.-S., Specht, L.W., Blackburn, H. (2020) Reconstitution and modernization of lost Holstein male lineages using samples from a gene bank. J. Dairy Sci. 103, 4510-4516. DOI: https://doi.org/10.3168/jds.2019-17753. This article was selected by the Editor-in-Chief as the “Editor's Choice". 
  • Hughes, C. K., Maalouf, S.W., Liu, W.-S., Pate, J.L. (2019) Molecular profiling demonstrates modulation of immune cell function and matrix remodeling during luteal rescue. Biol. of Reprod 100(6), 1581–1596. DOI:10.1093/biolre/ioz03
  • Zhang, Y.Y., Han D.P., Dong, X.G., Wang, J.K., Chen, J.F., Yao, Y.Z., Darwish, H.Y.A., Liu, W.-S., Deng, X.M. (2019) Genome-wide profiling of RNA editing sites in sheep. Journal of Animal Science and Biotechnology 10, 31. DOI: 10.1186/s40104-019-0331-z.
  • Liu, W.-S. (2019) Mammalian sex chromosome structure, gene content and function in male fertility. Annu Rev Anim Biosci. 7, 103-124. DOI: 10.1146/annurev-animal-020518-115332
  • Dechow, C., Liu, W.-S. (2018) DNA methylation patterns in peripheral blood mononuclear cells from Holstein cattle with variable milk yield. BMC Genomics 19, 744. DOI: 10.1186/s12864-018-5124-9
  • Dechow, C., Liu, W.-S., Idun, J., Maness, W. (2018) Two dominant paternal lineages for North American Jersey artificial insemination sires. J. Dairy Sci. 101, 2281–84. DOI: 10.3168/jds.2017-13694. This article was selected by the Editor-in-Chief as the “Editor's Choice".
  • Liu, W.-S., Zhao, Y.Q., Lu, C., Ning, G., Ma, Y., Diaz, F., O'Connor, M. (2017) A novel testis-specific protein, PRAMEY, is involved in spermatogenesis in cattle. Reproduction 153, 847-863. DOI: 10.1530/REP-17-0013.
  • Maalouf, S.W., Liu, W.-S., Pate, J.L. (2016) MicroRNA in ovarian function. Cell Tissue Res 363, 7–18.
  • Zeng, W., Pan, C., Liu, W.-S. (2016). Regulatory role of noncoding RNAs during spermatogenesis. Spermatogenesis: Molecular Mechanisms, Regulation and Biological Perspectives (pp. 31-70). Nova Science Publishers, Inc.
  • Yue, X.-P. , Dechow, C., Liu, W.-S. (2015) A limited number of Y chromosome lineages present in North American Holsteins. J. Dairy Sci. 98, 2738–2745.  DOI: 10.3168/jds.2014-8601. This article was selected by the Editor-in-Chief as the “Featured Article" of the issue.
  • Maalouf, S.W., Liu, W.-S., Albert, I., Pate, J.L. (2014) Regulating life or death: Potential role of microRNA in rescue of the corpus luteum. Molecular and Cellular Endocrinology 398(1-2), 78-88. DOI: 10.1016/j.mce.2014.10.005 
  • Liu, W.-S., Chang, T.-C. (2014) Y chromosome-linked genes implicated in spermatogenesis in cattle.  Reproduction in Domestic Ruminants VIII. Ed: Jennifer Juengel, Akio Miyamoto, Christopher Price, Larry Raynolds, Mike Smith, and Robert Webb. Context Products Ltd., England. P239-255. DOI: 10.13140/2.1.1392.9922
  • Decker, J. E., McKay, S. D., Rolf,  M. M., Kim,  J. W., Alcalá, A. M., Sonstegard, T. S., Hanotte, O., Götherström,  A., Bradley, D. G., Seabury, C. M., Praharani, L., Babar, M. E., Yildiz, M. A., Michael P. Heaton, M. P., Liu, W.-S., Reecy, J.M., Saif-Ur-Rehman, M., Schnabel, R.D., Taylor, J.F. (2014) Worldwide patterns of divergence, migration and admixture in domesticated cattle. PLoS Genet. 10(3): e1004254. DOI: 10.1371/journal.pgen.1004254.
  • Yue, X.-P. , Dechow, C. D., Chang, T.-C., DeJarnette, J. M., Marshall, C. E., Lei, C. Z., Liu, W.-S. (2014) Copy number variations of the extensively amplified Y-linked genes, HSFY and ZNF280BY, in cattle and their association with male reproductive traits in Holstein bulls. BMC Genomics 15(1), 113.
  • Yue, X.-P. , Chang, T.-C., DeJarnette, J. M., Marshall, C. E., Lei, C. Z., Liu, W.-S. (2013) Copy number variation of PRAMEY across breeds and its association with male fertility in Holstein sires. J. Dairy Sci. 96,  8024-34. DOI: 10.3168/jds.2013-7037.
  • Chang, T.-C., Yang, Y., Retzel, E., Liu, W.-S. (2013) Male-specific region of the bovine Y chromosome is gene rich with a high transcriptomic activity in testis development. Proc. Natl. Acad. Sci. U.S.A. 110(30), 12373-12378.
  • Mistry, B.V., Chang, T.-C., Yasue, H., Chiba, M., Zhao, Y.-Q., Oatley, J., Diaz, F., Liu, W.-S. (2013) Differential expression of PRAMEL1, a cancer/testis antigen, during spermatogenesis in the mouse. PLoS ONE 8, e60611.
  • Vallimont, J.E., Dechow, C.D., Daubert, J.M., Dekleva, M.W., Blüm, J., Barlieb, C.M., Liu, W.-S, Varga, G.A., Heinrichs, A.J., Baumrucker C.R. (2013) Feed utilization and its associations with fertility and productive life in 11 Commercial Pennsylvania Tie Stalls. J. Dairy Sci. 96, 1251-4. 
  • Groenen, M., Archibald, A., Uenishi, H., Tuggle, C., Takeuchi, Y., Rothschild, M., Rogel-Gaillard, C., Park, C., Milan, D., Megens, H-jan., et al. (2012) Analyses of pig genomes provide insight into porcine demography and evolution. Nature 491, 393-398.
  • Brown, D. E., Dechow, C. D., Liu, W.-S., Harvatine, K.J., Ott, T. (2012) Association of telomere length with age, herd and culling in lactating Holsteins. J. Dairy Sci. 95, 6384-6387.
  • Dekleva, M.W., Dechow, C.D., Daubert, J.M., Liu, W.-S., Varga, G.A., Bauck, S., Woodward B. (2012)  Interactions of milk, fat and protein yield genotypes with herd feeding characteristics. J Dairy Sci. 95(3),1559-1564.
  • Chang, T.-C., Klabnik, J., Liu, W.-S. (2011) Regional selection acting on OFD1 genes. PLoS ONE 6(10), e26195.
  • Chang, T.-C., Yang, Y., Yasue, H., Bharti, A., Retzel, E., Liu, W.-S. (2011) The expansion of the PRAME gene family in eutheria. PLoS ONE 6(2), e16867.
  • Yang, Y., Chang, T.-C., Yasue, H., Bharti, A, Retzel, E., Liu, W.-S. (2011) ZNF280BY and ZNF280AY: autosome derived Y chromosome gene families in Bovidae.  BMC Genomics 12, 13.
  • Ma, J.-G., Chang, T.-C., Yasue, H., Farmer, A.D., Crow, J.A., Eyer, K., Hiraiwa, H., Shimogiri, T., Meyers, S.N., Beever, J.E., Schook, L.B., Retzel, E., Beattie, C.W.,  Liu, W.-S. (2011) A high resolution comparative map of porcine chromosome 4 (SSC4). Animal Genetics 42(4), 440-444.
  • Vallimont, J. E., C.D. Dechow, J.M. Daubert, M.W. Dekleva, J.W.  Blum, C.M. Barlieb, W.-S. Liu, G.A. Varga, A.J. Heinrichs, and C.R. Baumrucker (2011) Heritability of Gross Feed Efficiency and Associations with Yield, Intake, Residual Intake, Body Weight and Body Condition Score in 11 Commercial Pennsylvania Tie Stalls. J. of Dairy Sci. 94(4), 2108-2113.
  • Chang, T.-C, Liu, W.-S. (2010) The molecular evolution of PL10 homologs. BMC Evolutionary Biology 10, 127.
  • Vallimont, J.E., Dechow, C.D., Daubert, J.M., Dekleva, M. W., Blüm, J.,Barlieb,C. M., Liu, W.-S, Varga, G.A., Heinrichs, A.J., Baumrucker C.R. (2010) Genetic parameters of feed intake, production, body weight, body condition score, and selected type traits of Holstein cows in commercial tie-stall barns. J Anim Sci  93(10), 4892-4901.
  • Porada, C.D., Sanada, C., Long, C.R., Wood, J.A., Desai, J.,Frederick, N., Millsap, L., Bormann, C., Menges, S.L., Hanna, C., Flores-Foxworth, G., Shin, T., Westhusin, M.E., Liu, W.-S., Glimp, H., Zanjani, E.D., Lozier, J.N., Pliska, V., Stranzinger, G., Joerg, H., Kraemer, D.C., Almeida-Porada, G. (2010) Clinical and molecular characterization of a re-established line of sheep exhibiting hemophilia A. Journal of Thrombosis and Haemostasis 8(2), 276-85. ePub: Nov 23, 2009.
  • For more publications, please visit https://www.researchgate.net/profile/Wansheng_Liu