University of Kentucky College of Agriculture, Food & Environment


Gluck Center > Directory > Gluck Faculty >MacLeod, JN > Curriculum Vitae


James N. MacLeod, VMD, PhD
(859) 218-1099

Department of Veterinary Science
M.H. Gluck Equine Research Center
College of Agriculture, Food & Environment
University of Kentucky


1980; University of Delaware, College of Agricultural Sciences, B.S., magna cum laude, with distinction.

1984; University of Pennsylvania, School of Veterinary Medicine, V.M.D.

1990; University of Pennsylvania, Faculty of Arts and Sciences, Ph.D.

1992; University of Pennsylvania, School of Medicine, Endocrinology & Genetics Fellowship.

Current Academic Appointments:

2003-present - Adjunct Associate Professor of Molecular Genetics, Department of Biomedical Sciences, College of Veterinary Medicine, Cornell University, Ithaca, New York.

2003-present - John S. and Elizabeth A. Knight Chair / Professor of Veterinary Science, Maxwell H. Gluck Equine Research Center, University of Kentucky, Lexington, Kentucky.

2007-present - Professor of Orthopaedic Surgery, College of Medicine, University of Kentucky, Lexington, Kentucky.

Previous Academic Positions:

1984-1988 - Research Assistant (doctoral thesis research), University of Pennsylvania, College of Arts and Sciences - Biochemistry,  Philadelphia, Pennsylvania.

1988-1992 - Research Fellow, University of Pennsylvania, School of Medicine and the Howard Hughes Medical Institute, Philadelphia, Pennsylvania.

1992-1999 - Assistant Professor of Molecular Genetics, James A. Baker Institute for Animal Health, College of Veterinary Medicine, Cornell University, Ithaca, New York.

1999-2003 - Associate Professor of Molecular Genetics, James A. Baker Institute for Animal Health, College of Veterinary Medicine, Cornell University, Ithaca, New York.

Selected Academic Awards:

1984 - Student Chapter of the American Veterinary Medical Association Prize

1986 - Phi Zeta Research Award

1992 - Dorothy Donnelley Faculty Development Award

1993 - Marilyn M. Simpson Career Development Award

1998 - Pfizer Award for Research Excellence

2008 - Dickson Professor of Equine Science and Management

2009 - Prestigious Research Paper Award (co-recipient), College of Agriculture, University of Kentucky


Mobile Swivel and Tether Device for Attachment to Laboratory Animals, Patent No. 4,756,770, issued July 12, 1988, assigned to the University of Pennsylvania.

Canine Erythropoietin Gene and Recombinant Protein, No. 6,696,411, issued February 24, 2004, assigned to Cornell University.

Selected Grant History:

1995-1998 - Morris Animal Foundation. Clinical Efficacy and Safety of Recombinant Canine Erythropoietin, Principal Investigator, Total direct support: $79,796.

1997-2000 - Arthritis Foundation. Cartilage-Specific Fibronectin Isoform and Osteoarthritis,  Co-Principal Investigator (Nancy Burton-Wurster, Co-PI), Total direct support: $200,149.

1997-2002 - Department of Health and Human Services - Public Health Service, National Institutes of Health  RO1 AR44340. Cartilage-Specific Fibronectin Isoform,  Principal Investigator, Total direct support: $871,506.

1999-2002 - Pfizer, Inc. Clinical Efficacy and Safety of Recombinant Canine Erythropoietin, Principal Investigator, Total direct support: $98,963.

1999-2000 - Alumni Grants Fund, Cornell University. Development of an Internet-Based Program for Data Collection in Multicenter Studies, Principal Investigator, Total direct support $3,516.

1999-2000 - Feline Health Center, Cornell University. Expression of Recombinant Feline Erythropoietin, Principal Investigator, Total direct support: $30,000.

2000-2002 - The Winn Feline Foundation. Bioactivity of Recombinant Feline Erythropoietin, Principal Investigator, Total direct support: $30,000.

2001-2004 - Arthritis Foundation. Cartilage-Specific Fibronectin Isoform and Osteoarthritis, Principal Investigator (Nancy Burton-Wurster, Co-PI), Total direct support: $250,000.

2001-2004 - Arthritis Foundation. Cartilage-Specific Fibronectin Isoform and Osteoarthritis,  Principal Investigator, Total direct support: $250,000.

2005-2008 - Morris Animal Foundation.  Horse Genomics: Linear Mapping and Microarray Development, Co-Investigator (Ernest Bailey, PI), Total direct support: $15,000 (MacLeod laboratory portion).

2006-2008 - Morris Animal Foundation.  Articular Cartilage Maturation and Repair, Fellowship sponsor for Michael Mienaltowski, Total direct support: $70,000.

2007-2011 - Morris Animal Foundation.  Program in Equine Medical Genetics, Co‑Principal Investigator (James Mickelson, University of Minnesota, Program Director), Total direct support: $30,000 (MacLeod laboratory portion).

2008-2009 - University of Kentucky Health Care – Department of Orthopaedic Surgery, Metamorphosis of the Salamander as a Model of Postnatal Maturation of Articular Cartilge in Mammals.  Co-Principal Investigator (Christian Lattermann, Co-PI), Total direct support: $19,500.

2009-2012 - National Science Foundation.  Exon Splice Pattern Characterization of the Whole mRNA Transcriptome, Co-Principal Investigator (Jinze Liu, PI), Total direct support: $1,027,384.

2009-2011 - Morris Animal Foundation.  Orthopaedic Pathology and Genetic Associations in Cervical Stenotic Myelopathy.  Sponsor for Jennifer Janes, Total direct support: $100,000.

2010-2012 - Kentucky Horse Racing Commission.  Safety and anti-inflammatory efficacy of glucocorticoids for intra-articular therapy in racehorses.  Principal Investigator, Total direct support: $99,969.

2010-2012 - Grayson-Jockey Club Foundation. Orthopaedic Pathology and Genetic Associations with Cervical Stenotic Myelopathy.  Principal Investigator, Total direct support: $102,193.

2010-2015 - Lourie Foundation. Cellular Mechanisms of Articular Cartilage, Tendon, and Ligament Repair.  Principal Investigator, Total direct support: $500,000.

2011-2013 - Kentucky Horse Racing Commission.  Articular Cartilage Stem Cells.  Principal Investigator, Total direct support: $100,000.

2012-2016 - Department of Health and Human Services - Public Health Service, National Institutes of Health  RO1 HG006272. Unlocking transcript diversity via differential analyses of splice graphs, Co-Investigator (Jan Prins and Jinze Lui, Co-PIs), Total direct support: $900,000.

Selected Publications:

MacLeod J.N., Sorensen, M.P., Shapiro, B.H. 1987. Strain-independent elevation of hepatic monooxygenase enzymes in female mice. Xenobiotica 17:1095-1102.

MacLeod, J.N., Shapiro, B.H. 1988. Repetitive blood sampling in unrestrained and unstressed mice using a chronic indwelling right atrial catheterization apparatus. Laboratory Animal Science 38:603-608.

MacLeod, J.N., Shapiro, B.H. 1989. Growth hormone regulation of hepatic drug metabolizing enzymes in the mouse. Biochemical Pharmacology 38:1673 1677.

Shapiro BH, MacLeod JN, Pampori NA, Morrissey JJ, Lapenson DP, Waxman DJ. 1989. Signalling elements in the ultradian rhythm of circulating growth hormone regulating expression of sex-dependent forms of hepatic cytochrome P450. Endocrinology 125:2935-2944.

Waxman DJ, Morrissey JJ, MacLeod JN, Shapiro BH. 1990. Depletion of serum growth hormone in adult female rats by neonatal monosodium glutamate treatment without loss of female-specific hepatic enzymes P-450 2d (IIC12) and steroid 5 α-reductase. Endocrinology 126:712-720.

MacLeod, J.N., Worsley, I., Ray, J., Friesen, H.G., Liebhaber, S.A., Cooke, N.E. 1991. Human growth hormone-variant is a biologically active somatogen and lactogen. Endocrinology 128:1298-1302.

MacLeod, J.N., Liebhaber, S.A., MacGillivray, M.H., Cooke, N.E. 1991. Identification of a splice site mutation in the human growth hormone variant gene. Am. J. Hum. Genet. 48:1168-1174.

MacLeod, J.N., Pampori, N.A., Shapiro, B.H. 1991. Sex differences in the ultradian pattern of plasma growth hormone concentrations in mice. J. Endocrinol. 131:395-399.

Urbanek, M., MacLeod, J.N., Cooke, N.E., Liebhaber, S.A. 1992. Expression of a human growth hormone receptor isoform is predicted by tissue specific alternative splicing of exon 3 of the human growth hormone receptor gene transcript. Mol. Endocrinol. 6:279-287.

MacLeod, J.N., Lee, A.K., Liebhaber, S.A., Cooke, N.E. 1992. Developmental control and alternative splicing of the placentally expressed transcripts from the human growth hormone gene cluster. J. Biol. Chem. 267:14219-14226.

MacLeod, J.N., Burton-Wurster, N., Gu, D.N., Lust, G. 1996. Fibronectin mRNA splice variant in articular cartilage lacks basesencoding the V, III-15, and I-10 proteins segments. J. Biol. Chem. 271:18954-18960.

Albrecht BA, MacLeod JN, Daels PF. 1997. Differential transcription of steroidogenic enzymes in the equine primary corpus luteum during diestrus and early pregnancy. Biology of Reproduction, 56:821-829.

Burton-Wurster, N., Lust, G., MacLeod, J.N. 1997. Cartilage fibronectin isoforms: in search of functions for a special population of matrix glycoproteins. Matrix Biol. 15:441-454.

Stewart, M.C., Farnum, C.E., MacLeod, J.N. 1997. Expression of p21CIP1/WAF1 in chondrocytes. Calcif. Tissue Int. 61:199-204.

Todhunter RJ, Fubini SL, Vernier-Singer M, Wootton JAM, Lust G, Freeman KP, MacLeod JN. 1998. Experimental acute synovitis and intra-articular methylprednisolone acetate in ponies. Osteoarthritis and Cartilage, 6:94-105.

MacLeod, J.N., Fubini, S.L., Gu, D.N., Tetreault, J.W., Todhunter, R.J. 1998. Effect of synovitis and corticosteroids on transcription of cartilage matrix proteins. Am. J. Vet. Res. 59:1021-1026.

Burton-Wurster, N., Borden, C., Lust, G., MacLeod, J.N. 1998. Expression of the (V+C)–fibronectin isoform is tightly linked to the presence of a cartilaginous matrix. Matrix Biol. 17:193-203.

MacLeod, J.N., Tetreault, J.W., Lorschy, K.S., Gu, D.N. 1998. Expression and bioactivity of canine erythropoietin. Am. J. Vet. Res. 59:1144-1148.

Campbell BG, Wootton JAM, MacLeod JN, Minor RR. 1998. Sequence of canine COL1A2: Nucleotide substitutions affecting the cyanogen bromide peptide map of the a2(I) chain. Archives of Biochemistry and Biophysics, 357:67-75.

Randolph, J.F., Stokol, T., Scarlett, J.M., MacLeod, J.N. 1999. Comparison of the biologic activity and safety of recombinant canine erythropoietin to recombinant human erythropoietin in normal Beagle dogs. Am. J. Vet. Res. 60:636-642.

Burton-Wurster, N., Gendelman, R., Chen, H., Gu, D.N., Tetreault, J.W., Lust, G., Schwarzbauer, J.E., MacLeod, J.N. 1999. The cartilage-specific (V+C)– fibronectin isoform exist primarily in homodimeric and monomeric configurations. Biochemical J. 341:555-561.

Stewart, M.C., Saunders, K.M., Burton-Wurster, N., MacLeod, J.N. 2000. Phenotypic stability of articular chondrocytes in vitro: the effects of culture models, BMP-2 and serum supplementation. J. Bone Min. Res. 15:166-174.

Albrecht BA, MacLeod JN, Daels PF. 2000. Differential expression of steroidogenic enzymes by the primary corpora lutea of pregnant mares during equine chorionic gonadotropin secretion. Journal of Reproduction and Fertility, Suppl. 56:627-634.

Campbell BG, Wootton JAM, MacLeod JN, Minor RR.  2000. Sequence of normal canine COL1A1 cDNA and identification of a heterozygous a1(1) collagen Gly208Ala mutation in a severe case of canine osteogenesis imperfecta. Archives of Biochemistry and Biophysics, 384:37-46.

Albrecht BA, MacLeod JN, Daels PF. 2001. Expression of 3β-hydroxysteroid dehydrogenase, cytochrome P450 17a-hydroxylase/17,20-lyase and cytochrome P450 aromatase enzymes in corpora lutea of diestrous and early pregnant mares. Theriogenology, 55:551-561.

Fubini, S.L., Todhunter, R.J., Burton-Wurster, N., Vernier-Singer, M., MacLeod, J.N. 2001. Corticosteroids alter the differentiaed phenotype of articular chondrocytes. J. Orthop. Res. 19:688-695.

Campbell BG, Wootton JAM, MacLeod JN, Minor RR.  2001. Canine COL1A2 mutation resulting in C-terminal truncation of pro-α2(1) and severe osteogenesis imperfecta.  Journal of Bone and Mineral Research, 16:1147-1153.

Chen, H., Gu, D.-N., Burton-Wurster, N., MacLeod, J.N. 2002. Absence of the I-10 protein segment mediates restricted dimerization of the cartilage-specific fibronectin isoform. J. Biol. Chem. 277:20095-20103.

Burton-Wurster, N., Chen, H., Gendelman, R., Jackson, M.L., Gagliardo, L.F., Gu, D.N., Zelko, R.R., Lust, G, MacLeod, J.N. 2002. Specific immunological detection of the (V+C)– fibronectin isoform. Matrix Biology, 21:393-398.

Gendelman, R., Burton-Wurster, N., MacLeod, J.N., Lust, G. 2003. The cartilage-specific fibronectin isoform has a high affinity binding site for the small proteoglycan decorin. Journal of Biological Chemistry, 278:11175-11181.

Mertens WD, MacLeod JN, Fubini SL, Vernier-Singer M, Nixon AJ, Todhunter RJ.  2003. Polysulfated glycosaminoglycans modulate transcription of interleukin-1 treated chondrocytes in monolayer culture.  Veterinary Comparative Orthopaedics and Traumatology, 2:93-98.

Randolph, J.F., Scarlett, J.M., Stokol, T., MacLeod, J.N. 2004. Clinical efficacy and safety of recombinant canine erythropoietin in dogs with anemia of chronic renal failure and dogs with recombinant human erythropoietin-induced red cell aplasia. Journal of Veterinary Internal Medicine, 18:81-91.

Stewart, M.C., Kadlcek, R.M., Robbins, P., MacLeod, J.N., Ballock, T.R. 2004. Expression and activity of the CDK inhibitor p57Kip2in chondrocytes undergoing hypertrophic differentiation. Journal of Bone and Mineral Research, 19:123-132.

Randolph, J.F., Scarlett, J.M., Stokol, T., Saunders, K.M., MacLeod, J.N. 2004. Expression, bioactivity, and clinical assessment of recombinant feline erythropoietin. American Journal of Veterinary Research, 65:1355-1366.

Steffey, M.A., Miura, N., Todhunter, R.J., Nykamp, S.G., Freeman, K.P., Scarpino, V., Vernier-Singer, M.A., Erb, H.N., MacLeod, J.N., Lust, G., Burton-Wurster, N. 2004. The potential and limitations of cartilage-specific (V+C)– fibronectin and cartilage oligomeric matrix protein as osteoarthritis biomarkers in canine synovial fluid, Osteoarthritis and Cartilage, 12:818-825.

Walker, M. C., Mandell, T. C., Crawford,  P. C., Simon, G. G., Cahill, K. S., Fernandes, P. J., MacLeod, J. N., Byrne, B. J., Levy, J. K. 2005. Expression of erythropoietin in cats treated with a recombinant adeno-associated viral vector.  American Journal of Veterinary Research, 66:450-456.

Coleman, S. J., Gong, G., Gaile, D. P., Chowdhary, B. P., Bailey, E., Liu, L., MacLeod, J. N. 2007. Evaluation of COMPASS as a comparative mapping tool for ESTs using horse radiation hybrid maps.  Animal Genetics, 38:294-302.

Huang L, Zhu W, Saunders CP, MacLeod JN, Zhou M, Stromberg AJ, Bathke A.  2008. A novel application of quantile regression for identification of cartilage biomarkers in equine microarray data. BMC Bioinformatics, 9:300.

Raudsepp T, Gustafson-Seabury A, Durkin K, Wagner ML, Goh G, Seabury CM, Brinkmeyer-Langford C, Lee E-J, Agarwala R, Stallknecht-Rice E, Schäffer AA, Tozaki T, Yasue H, Penedo MC, Lyons LA, Khazanehdari KA, Binns MM, MacLeod JN, Distl O, Guérin G, Leeb T, Mickelson JR, Chowdhary BP. 2008. A 4103 marker integrated map of the horse genome. Cytogenetic and Genome Research, 122:28-36.

Mienaltowski MJ, Huang L, Stromberg AJ, MacLeod JN.  2008. Differential gene expression associated with postnatal articular cartilage maturation.  BMC Musculoskeletal Disorders, 9:149.

Haneda S, Nagaoka K, Nambo Y, Kikuchi M, Nakano Y, Matsui M, Miyake Y, MacLeod JN, Imakawa K.  2009. Expression of interleukin-1 receptor antagonist (IL-1RN) in the equine endometrium is regulated by a steroid hormone derived from the conceptus.  Domestic Animal Endocrinology, 36:209-218.

Astakhova NM, Perelygin AA, Zharkikh AA, Lear TL, Coleman SJ, MacLeod JN, Brinton MA. 2009.   Characterization of equine and other vertebrate TLR3, TLR7, and TLR8 genes.  Immunogenetics. 61:529-39.

Mienaltowski MJ, Huang L, Frisbie DD, McIlwraith CW, Stromberg AJ, Bathke A, MacLeod JN.  2009. Transcriptional profiling differences for articular cartilage and repair tissue in equine joint surface lesions.  BMC Medical Genomics, 14;2:60.

Wade CM, Giulotto E, Sigurdsson S, Zoli M, Gnerre S, Imsland F, Lear TL, Adelson DL, Bailey E, Bellone RR, Blöcker H, Distl O, Edgar RC, Garber M, Leeb T, Mauceli E, MacLeod JN, Penedo MC, Raison JM, Sharpe T, Vogel J, Andersson L, Antczak DF, Biagi T, Binns MM, Chowdhary BP, Coleman SJ, Della Valle G, Fryc S, Guérin G, Hasegawa T, Hill EW, Jurka J, Kiialainen A, Lindgren G, Liu J, Magnani E, Mickelson JR, Murray J, Nergadze SG, Onofrio R, Pedroni S, Piras MF, Raudsepp T, Rocchi M, Røed KH, Ryder OA, Searle S, Skow L, Swinburne JE, Syvänen AC, Tozaki T, Valberg SJ, Vaudin M, White JR, Zody MC; Broad Institute Genome Sequencing Platform; Broad Institute Whole Genome Assembly Team, Lander ES, Lindblad-Toh K. 2009.  Genome sequence, comparative analysis, and population genetics of the domestic horse.  Science, 326:865-7.

McGivney BA, Eivers SS, MacHugh DE, MacLeod JN, O’Gorman GM, Park SDE, Katz LM,  and Hill EW.  Transcriptional modifications following exercise in Thoroughbred horse skeletal muscle highlight molecular mechanisms that lead to muscle hypertrophy. BMC Genomics, 10:638, 2009.

Mienaltowski MJ, Huang L, Bathke A, Stromberg AJ, MacLeod JN.  Transcriptional comparisons between articular repair tissue, neonatal cartilage, cultured chondrocytes, and mesenchymal stromal cells. Briefings in Functional Genomics & Proteomics, doi:10.1093/bfgp/elq007, 2010.

Wang K, Singh D, Zeng Z, Coleman SJ, Huang Y, Savich GL, Xiaping H, Mieczkowski P, Grimm SA, Perou CM, MacLeod JN, Chiang DY, Prins JF, Liu J.  Accurate mapping of RNA-seq reads for splice junction discovery. Nucleic Acids Research, doi:10.1093/nar/gkq622, 2010.

Coleman SJ, Zeng Z, Wang K, Luo S, Khrebtukova I, Mienaltowski MJ, Schroth GP, Liu, J., MacLeod JN.  Structural annotation of equine protein-coding genes determined by mRNA sequencing. Animal Genetics, 41 (Suppl. 2):121–130, 2010.

Cosden RS, Lattermann C, Romine S, Gao J, Voss SR, MacLeod JN.  Intrinsic repair of full-thickness articular cartilage defects in the axolotl salamander. Osteoarthritis and Cartilage, 19:200-205, 2011.

Kikuchi M, Nakano Y, Nambo Y, Haneda S, Matsui M, Miyake Y, MacLeod JN, Nagaoka K, Imakawa K.  Production of Calcium Maintenance Factor Stanniocalcin-1 (STC1) by the Equine Endometrium During the Early Pregnant Period.  Journal of Reproduction and Development, 57:203-211, 2011.

Vanderman KS, Tremblay M, Zhu W, Shimojo M, Mienaltowski MJ, Coleman SJ, MacLeod JN.  Brother of CDO (BOC) Expression in Articular Cartilage.  Osteoarthritis and Cartilage, 19:435-438, 2011.

Go YY, Bailey E, Cook DG, Coleman SJ, MacLeod JN, Chen KC, Timoney PJ, Balasuriya UB.  Genome-Wide Association Study Among Four Horse Breeds Identifies a Common Haplotype Associated with the in vitro CD3+ T Cell Susceptibility/Resistance to Equine Arteritis Virus Infection.  Journal of Virology, 85:13174-13184. 2011.

Cosden RS, Bickett MM, Lattermann C, MacLeod JN.  Structural and functional analysis of intra-articular interzone tissue in axolotl salamanders. Osteoarthritis and Cartilage, 20:1347-1356, 2012.

Huang Y, Hu Y, Jones CD, MacLeod JN, Chiang DY Liu Y, Prins JF, Liu J. A Robust Method for Transcript Quantification with RNA-seq Data. Journal of Computational Biology, 20:167-187, 2013.

Coleman SJ, Zeng Z, Hestand MS, Liu J, MacLeod JN. Analysis of unannotated equine transcripts identified by mRNA sequencing. PLoS One, 8: e70125. doi: 10.1371/journal.pone.0070125. 2013.

Janes J, Garrett KS, McQuerry KJ, Pease AP, Williams NM, Reed SM, MacLeod JN.  Comparison of cervical radiographs and MRI to assess vertebral canal stenosis in Wobbler Syndrome.  Equine Veterinary Journal, 46:681-686, doi: 10.1111/evj.12221, 2014.

Mikkel S, Jónsson H, Chang D, Sarkissian C, Ermini L, Ginolhac A, Albrechtsen A, Dupanloup I, Foucal A, Petersen B, Raghavan M, Seguin-Orlando, A, Korneliussen T, Velazquez AMV, Stenderup J, Hoover CA, Rubin CJ, Alfarhan AH, Alquraishi SA, Al-Rasheid KAS, MacHugh DE, Kalbfleish T, MacLeod JN, Rubin EM, Sicheritz-Ponten T, Andersson L, Hofreiter M, Marques-Bonet T, Gilbert MTP, Nielsen R, Excoffier L, Willerslev E, Shapiro B, Orlando L. Prehistoric horse genomes reveal the genetic foundation and cost of domestication.  Proceedings National Academy of Sciences USA, 111:E5661-5669, doi: 10.1073/pnas.1416991111, 2014.

Hestand MS, Kalbfleisch TS, Coleman SJ, Zeng Z, Orlando L, MacLeod JN.  Annotation of the protein-coding regions of the horse genome.  PLoS One, 10:e0124375. doi: 10.1371/journal.pone.0124375, 2015.

Rebolledo-Mendez J, Hestand MS, Coleman S, Zeng Z, Orlando L, MacLeod JN, Kalbfleisch T.  Comparison of the equine reference sequence with its Sanger source data and new Illumina reads, PLoS One, 10:e0126852. doi: 10.1371/journal.pone.0126852, 2015.

Janes J, Garrett KS, McQuerry KJ, Waddell S, Voor M, Reed SM, Williams NM, MacLeod JN.  Skeletal Pathology of the Cervical Articular Processes in Equine Cervical Stenotic Myelopathy. Veterinary Pathology, pii: 0300985815593127, 2015. [Epub ahead of print].


Maxwell H.Gluck Equine Research Center
Department of Veterinary Science, University of Kentucky
Lexington, Kentucky 40546-0099

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