Publications (selected):

For the cover article written for the lay public about genetics, genomics and the work of the Cheng lab, see The article is adapted from Keith’s concluding talk for the 2001 Penn State Frontiers of Science Lecture Series “Decoding Life’s Instruction Book: Genetics and Genomics.” The title of his talk, “A Model for Humans,” is also an article in Research Penn State, by Teresa Rafacz, September 2001, Volume 22, Issue 3.


All publications are PDF format unless otherwise noted:

  1. Cheng, KC, Katz SR, Lin AY, Xin X, Ding Y. Whole-Organism Cellular Pathology: A Systems Approach to Phenomics. In N. S. Foulkes (Ed.), Genetics, Genomics and Fish Phenomics. (2016) (pp. 89–115). ISBN: 9780128048009
  2. Canfield VA, Berg A, Peckins S, Wentzel SM, Ang KC, Oppenheimer S, Cheng KC. Molecular Phylogeography of a Human Autosomal Skin Color Locus Under Natural Selection. G3 3(11): 2059-67, 2013.
  3. Tsetskhladze ZR, Canfield VA, Ang KC, Wentzel SM, Reid KP, Berg AS, Johnson SL, Kawakami K, Cheng KC. Functional Assessment of Human Coding Mutations Affecting Skin Pigmentation Using Zebrafish. PloS ONE 7(10): e47398, October 2012 doi:10.1371/journal.pone.0047398
  4. Ang KC, Ngu MS, Reid KP, Teh MS, Aida ZS, Koh DXR, Berg A, Oppenheimer S, Salleh H, Clyde MM, Md-Zain BM, Canfield VA, Cheng K. Skin Color Variation in Orang Asli Tribes of Peninsular Malaysia. PLoS ONE 7(8): e42752, August 2012 doi:10.1371/journal.pone.0042752
  5. Valenzuela RK, Henderson MS, Walsh MH, Garrison NA, Kelch JT, Cohen-Barak O, Erickson DT, Meaney FJ, Walsh JB, Cheng KC, Ito S, Wakamatsu K, Frudakis F, Thomas M, Brilliant MH. Predicting Phenotype from Genotype: Normal Pigmentation. J Forensic Sci 55:315-322, 2010
  6. Cheng KC, Waldstein N. Unraveling genetic code: Social, moral questions arise from DNA (Text submitted as, “What does the human genome project mean to me”) Review & Opinion section, Harrisburg Patriot-News, Feb 15, 2009
  7. Zinnanti WJ, Lazovic J, Griffin K, Skvorak KJ, Paul HS, Homanics GE, Bewley MC, Cheng KC, LaNoue KF, Flanagan JM. Dual mechanism of brain injury and novel treatment strategy in maple syrup urine disease.Brain 2009; doi: 10.1093/brain/awp024. [For a copy of the paper, please send a request]
  8. Spitsbergen JM, Blazer VS, Bowser PR, Cheng KC, Cooper KR, Cooper TK, Frasca S Jr., Groman DB, Harper CM, Law JM, Marty GD, Smolowitz RM, St. Leger J, Wolf DC, Wolf JC. Finfish and aquatic invertebrate pathology resources for now and the future. Comparative Biochemistry and Physiology, Part C 149:249–257, 2009. [For a copy of the paper, please send a request to]
  9. Cheng KC. Zebrafish experts speak (editorial). Zebrafish 5:9-24, 2008
  10. Ekker SC, Parichy DM, Cheng KC. Research implications of pigment biology in zebrafish. Zebrafish 5:233-235, 2008. PMID: 19133820
  11. Cheng KC. Skin color in fish and humans: Impacts on science and society. Zebrafish 5:237-242, 2008. PMID: 19133821
  12. Canada BA, Thomas GK, Cheng KC, Wang JZ, Liu Y. Automatic Lattice Detection in Near-Regular Histology Array Images. Proceedings of the IEEE International Conference on Image Processing (ICIP), San Diego, California, IEEE, October 2008
  13. Canada BA, Thomas GK, Cheng KC, Wang JZ, Liu Y. Towards Efficient Automated Characterization of Irregular Histology Images via Transformation to Frieze-Like Patterns. Proceedings of the ACM International Conference on Image and Video Retrieval (CIVR 2008), pp. 581-590, Niagara Falls, Canada, July 2008
  14. Cheng KC. Demystifying Skin Color and “Race”. In R.E. Hall (ed.): Racism in the 21st Century, Chapter 1, pp. 3-23, Springer, 2008
  15. Canada BA, Thomas GK, Cheng KC*, Wang JZ.* Automated segmentation and classification of zebrafish histology images for high-throughput phenotyping. Proceedings of the Third IEEE-NIH Life Science Systems and Applications Workshop, pp. 245-248, Washington, D.C., November 2007. (*equal contributors) ISBN number: 978-1-4244-1813-8
  16. Zinnanti WJ, Lazovic J, Housman C, LaNoue K, O’Callaghan JP, Simpson I, Woontner M, Goodman SI, Connor JR, Jacobs RE, Cheng KC. Mechanism of age-dependent susceptibility and novel treatment strategy in glutaric acidemia type I. J Clin Invest 117:3258-3270, 2007. PMID: 17932566
  17. Norton HL, Kittles RA, Parra E, McKeigue P, Mao X, Cheng KC, Canfield VA, Bradley DG, McEvoy B, Shriver MD. Genetic evidence for the convergent evolution of light skin in Europeans and East Asians. Molecular Biol Evol 24:710-722, 2007. PMID: 17182896
  18. Chi A, Valencia JC, Hu Z-Z, Watabe H, Yamaguchi H, Mangini NM, Huang H, Canfield VA, Cheng KC, Yang OF, Abe R, Yamagishi S, Shabanowitz J, Hearing VJ, Wu C, Appella E, Hunt DF. Proteomic and bioinformatic characterization of the biogenesis and function of melanosomes. J Proteome Res 5:3135-3144, 2006. PMID: 17081065
  19. Canada BA, Cheng KC, Wang JZ. QCHARM: A Novel Computational and Scientific Visualization Framework for Facilitating Discovery and Improving Diagnostic Reliability in Medicine. Proceedings of the American Medical Informatics Association Annual Symposium, pp. 870, Washington, D.C., November 2006
  20. Cheng KC, Canfield VA. The role of SLC24A5 in skin color. Invest Dermatol 16:836-838, 2006
  21. Moore JL, Rush LM, Breneman C, Mohideen M-AP, Cheng, KC. Zebrafish genomic instability mutants and cancer susceptibility. Genetics 174:585-600, 2006 (October cover article), 2006. PMID: 16888336
  22. Sabaliauskas NA, Foutz CA, Mest JR, Budgeon LR, Sidor A, Gershenson J, Joshi S, Cheng KC. High-throughput zebrafish histology. Methods 39:246-254, 2006. PMID: 16870470
  23. Zinnanti WJ, Lazovic J, Wolpert EB, Antonetti DA, Smith MB, Connor JR, Woontner M, Goodman SI, Cheng KC. New insights for glutaric aciduria type I. Brain 129:E55, 2006. PMID: 16870880
  24. Blasiole B, Canfield VA, Mohideen M-APK, Vollrath MA, Huss D, Dickman JD, Cheng KC, Fekete DM, Levenson, RL. Separate Na,K-Atpase genes are required for otolith formation and semicircular canal development in zebrafish. Dev Biol 294:148-160, 2006. PMID: 16566913
  25. Cheng KC. Bach’s Goldberg Variations: An artistic inspiration for art, science, medicine, and life. Wild Onions20:14-15, 2006
  26. Cheng KC. The golden mutation in zebrafish. Commentary #1, February 2006 PASPCR web-site posting.
  27. Zinnanti WJ, Lazovic J, Wolpert EB, Antonetti DA, Smith MB, Connor JR, Woontner M, Goodman SI, Cheng KC. A diet-induced mouse model for glutaric aciduria type I. Brain 129:899-910, 2006. PMID: 16446282
  28. Lamason RL, Mohideen M-AP, Mest JR, Wong AC, Norton HL, Aros MC, Juurynec MJ, Mao X, Humphreville VR, Humbert JE, Sinha S, Moore JL, Jagadeeswaran P, Ning G, Makalowska I, Zhao W, McKeigue PM, O’Donnell D, Kittles R, Parra EJ, Mangini NJ, Grunwald DJ, Shriver MD, Canfield VA, Cheng KC. SLC24A5 affects pigmentation in zebrafish and man. Science 310:1782-1786, 2005. (cover article) PMID: 16357253 [due to the large file size of the supplement and cover, we offer three versions: SciencePaperNoSuppl.pdf (the paper as presented in the journal; 457kb), SciencePaper&Suppl.pdf (the paper in the journal, including the supplementary data; 2857kb), and Science16Dec2005cover-full.jpg (just the cover, in high resolution, originally submitted with the caption, “Skin color is only gene deep”; 5814kb)]. Science 16 December 2005 “News of the Week”.             – Learning Materials Using the Golden Story
  29. Croushore JA, Blasiole B, Riddle RC, Thisse C, Thisse B, Canfield VA, Robertson GP, Cheng KC, Levenson R. ptena and ptenb genes play distinct roles in zebrafish embryogenesis. Developmental Dynamics 234:911-921, 2005. PMID: 16193492
  30. Duffy KT, McAleer MF, Davidson WR, Kari L, Kari C, Liu C-G, Farber SA, Cheng KC, Mest JR, Wickstrom E, Dicker AP, Rodeck U. Coordinate control of cell cycle regulatory genes in zebrafish development tested by cyclin D1 knockdown with morpholino phosporodiamidates and hydroxyprolyl-phosophono peptide nucleic acids. Nucleic Acids Res 33:4914-4921, 2005. PMID: 16284195
  31. Cheng KC. As I See It: Use science to support global policy. Harrisburg Patriot-News, December 12, 2004, F1, 6.
  32. Ostrander GK, Cheng KC, Wolf JC, Wolfe MJ. Shark cartilage, cancer and the growing threat of pseudoscience. Cancer Res 64:8485-8491, 2004. PMID: 15574750
  33. Cheng KC. Zebrafish experts speak (editorial). Zebrafish 1:85-103, 2004
  34. Cheng KC. A life-span atlas for the zebrafish (announcement). Zebrafish 1:69, 2004
  35. Gerhard GS, Malek RL, Keller E, Murtha J, Cheng KC. Zebrafish, Killifish, neither fish, both fish? J Gerontol59:B873-B875, 2004. PMID: 15472148
  36. Moore JL, Gestl EE, Cheng KC. Mosaic eyes, genomic instability mutants, and cancer susceptibility. Meth Cell Biol 76:555-568, 2004. PMID: 15602892
  37. Cheng KC, Levenson RL, Robishaw JD. Functional genomic dissection of multimeric protein families in zebrafish. Developmental Dynamics 228:555-567, 2003. PMID: 14579392
  38. Mohideen M-APK, Beckwith LG, Tsao-Wu GS, Moore JL, Wong ACC, Chinoy MR, Cheng KC. Histology-based screen for zebrafish mutants with abnormal cell differentiation. Developmental Dynamics 228:414-423, 2003. PMID: 14579380
  39. Gerhard GS, Cheng KC. A call to fins! Zebrafish as a gerontological model. Aging Cell 1:104-111, 2002.
  40. Gerhard GS, Kaufmann EJ, Wang X, Stewart R, Moore JL, Kasales CJ, Cheng KC. Life spans and senescent phenotypes in an outbred and an inbred strain of zebrafish (Danio rerio). Exp. Gerontology 37:1055-1068, 2002. PMID: 12213556
  41. Moore JL, Aros M, Steudel KG, Cheng KC. Fixation and decalcification of adult zebrafish for histological, immunocytochemical, and genotypic analysis. Biotechniques 32:296-298, 2002. PMID: 11848405
  42. Rajarao SJR, Canfield VA, Mohideen M-APK, Postlethwait JH, Cheng KC, Levenson R. The repertoire of Na,K-ATPase a and b subunit genes expressed in the zebrafish, Danio rerio. Genome Research 11:1211-1220, 2001. PMID: 11435403
  43. Mohideen M-APK, Moore JL, Cheng KC. Centromere-linked microsatellite markers for linkage groups 3, 4, 6, 7, 13 and 20 of zebrafish (Danio rerio). Genomics 67:102-106, 2000. PMID: 10945477
  44. Beckwith LG, Moore JL, Tsao-Wu GS, Harshbarger JC, Cheng KC. Ethylnitrosourea induces neoplasms in zebrafish (Danio rerio). Lab Invest 80:379-385, 2000 (cover article). PMID: 11956074
  45. Cheng KC, Beckwith L, Wang X. Update to: Agarose embedded tissue arrays for histological and genetic analysis. In Expression Genetics: High-Throughput Methods, M. McClelland and A. Pardee (eds.), Eaton Publishing, p. 37, 1999.
  46. Tsao-Wu GS, Weber CH, Budgeon LR, Cheng KC. Agarose embedded tissue arrays for histologic and genetic analysis. In Expression Genetics: High-Throughput Methods, Chapter 4, M. McClelland and A. Pardee (eds.), Eaton Publishing, pp. 31-36, 1999.
  47. Tsao-Wu GS, Weber CH, Budgeon LR, Cheng KC. Agarose embedded tissue arrays for histologic and genetic analysis. Biotechniques 25:614-618, 1998. PMID: 9793642
  48. Cheng K. You can’t win the Indy 500 in a Yugo: Help NSF funding: An op-ed. In HMS Beagle: The BioMedNet Magazine, Issue 28, Apr. 17, 1998.
  49. Cheng KC. A day well-spent for science: a first experience with Scientific Congressional Advocacy. The ASCB Newsletter 21(3), April 1998.
  50. Cheng KC, Moore JL. Genetic dissection of vertebrate processes in zebrafish: A comparison of uniparental and two-generation screens. Biochemistry and Cell Biology 75:525-533, 1997. PMID: 9551177
  51. Gestl EE, Kauffman EJ, Moore JL, Cheng KC. New conditions for generation of gynogenetic half-tetrad embryos in the zebrafish, Danio rerio. J Heredity 88:76-79, 1997.
  52. Cheng KC, Loeb LA. Genomic stability and instability: A working paradigm. In Current Topics in Microbiology and Immunology, Chapter 2, M.B. Kastan (ed.), Springer-Verlag, Berlin, pp. 5-18, 1997. PMID: 8979437
  53. Hite JM, Eckert KE, Kauffman EJ, Cheng KC. Factors affecting fidelity of DNA synthesis during PCR amplification of d(CA)n d(G-T)n microsatellite repeats. Nucleic Acids Res 24:2429-2434, 1996. PMID: 8710517
  54. Ham P, Cheng K. Construction of zebrafish spawning cages. The Zebrafish Science Monitor, pp. 5-8, December 18, 1995.
  55. Kauffman EJ, Gestl EE, Kim DJ, Walker C, Hite JM, Yan G, Rogan PK, Johnson SL, Cheng KC. Microsatellite-centromere mapping in the zebrafish (Danio rerio). Genomics 30:337-341, 1995. PMID: 8586435
  56. Cheng KC, Loeb LA. Genomic Instability and tumor progression: mechanistic considerations. Advances in Cancer Research 60:121-156, 1993. PMID: 8417498
  57. Cheng KC, Cahill DS, Kasai, Nishimura S, Loeb LA. 8-Hydroxyguanine, an abundant form of oxidative DNA damage, causes G – T and A – C substitutions. J Biol Chem 267:166-172, 1992. PMID: 1730583 “1112 ISI citations on 4-19-2010”
  58. Cheng KC, Preston BD, Cahill DS, Dosanjh MK, Singer B, Loeb LA. The vinyl chloride DNA derivative, N2,3-ethenoguanine, causes G to A transitions in E. coli. Proc Natl Acad Sci, USA. 88:9974-9978, 1991. PMID: 1946466
  59. Cheng KC, Dias M. Genomic Instability and Cancer: Cause and Effect (invited Keystone Meeting review).Cancer Cells 3:188-192, 1991. PMID: 1679993
  60. Cheng KC, Smith GR. Distribution of Chi-stimulated recombinational exchanges and heteroduplex endpoints in phage lambda. Genetics 123:5-17, 1989. PMID: 2530132
  61. Cheng KC, Smith GR. Cutting of Chi-like sequences by RecBCD enzyme of Escherichia coli. J Mol Biol 194:747-750, 1987. PMID: 2958631
  62. Cheng KC, Smith GR. Recombinational hotspot activity of Chi-like sequences. J Mol Biol 180:371-377, 1984. PMID: 6239928