Researchers at UCD


Kay Nolan

Senior Lecturer

School Of Biology & Environment Science
Science Centre - West
Dublin 4

Tel: +353 1 7162112


1977 B.Sc (Hons) First class University College Dublin (Major subject: Zoology; Minor subject: Biochemistry)

1983 Ph.D.                             University College Dublin Ph.D. (Advisor: Dr Paddy Joyce)

Post-doctoral Training
1983-1984                             Washington University School of Medicine, St Louis, USA (Mentor: Dr William S. Sly)

1984-1989                             St Louis University School of Medicine, St Louis, USA (Mentor: Dr William S. Sly)

1990-1991                             Imperial College of Science, Technology and Medicine (Mentor: Prof. Colin R. Hopkins)

Academic Career
1991-2004                             University College Dublin (College Lecturer)
2005-Present                        University College Dublin (Senior Lecturer)




Employer: University College Dublin
Position: Graduate Student
Employer: Washington University School
Position: Research Associate
Employer: St. Louis University School of Medicine
Position: Research Associate
Employer: Imperial College, London.
Position: Research Associate
Employer: University College Dublin
Position: College Lecturer
Employer: Duke University Medical Center
Position: Visiting Scholar


Year 1977 Institution: University College Dublin
Qualification: BSc Subject:
Year 1983 Institution: University College Dublin
Qualification: PhD Subject:



Book Chapters

Nolan, CM (2013) 'Igf2r Locus' In: Maloy, S and Hughes, K (eds). Brenner's Online Encyclopedia of Genetics, 2E. Oxford: Elsevier. [Details]
Killian J.K., Nolan C.M. and Jirtle R.L.; (2000) 'The evolution of genomic imprinting in Jurassic mammals' In: Ryder O. San Diego CA (eds). Genetic resources for the next century. [Details]
Nolan, C.M.; (1992) 'Receptor-mediated endocytosis and lysosomal transport' In: Editors: A.I. Magee and T. Wileman. Publishers: Oxford University Press, Oxford (eds). Protein targeting: A practical approach. [Details]
Nolan, C.M.; (1992) 'Receptor-mediated endocytosis and lysosomal transport' In: Editors: A.I. Magee and T. Wileman. Publishers: Oxford University Press, Oxford (eds). Protein targeting: A practical approach. [Details]
Nolan, C.M. and W.S. Sly; (1989) 'I-cell disease and pseudo-Hurler polydystrophy' In: Editors: Shriver, C.R., Beaudet, A.L., Sly, W.S. and Vallee, D. Publisher: McGraw Hill Inc., New York (eds). Metabolic Basis of Inherited Disease, Sixth Edition. , pp.1589-1601 [Details]

Peer Reviewed Journals

Atencia-Fernandez S, Shiel RE, Mooney CT, Nolan CM (2015) 'Muscular dystrophy in the Japanese Spitz: an inversion disrupts the DMD and RPGR genes'. Animal Genetics, 46 (2):175-184. [DOI] [Details]
Shiel RE, Kennedy LJ, Nolan CM, Mooney CT, Callanan JJ (2014) 'Major histocompatability complex class II alleles and haplotypes associated with non-suppurative meningoencephalitis in greyhounds'. Tissue antigens, 84 (3):271-276. [Details]
Terzo E, McConnell JF, Shiel RE, McAllister H, Behr S, Priestnall SL, Smith KC, Nolan CM, Callanan JJ (2012) 'Unique topographic distribution of greyhound nonsuppurative meningoencephalitis'. Veterinary radiology & ultrasound : the official journal of the American College of Veterinary Radiology and the International Veterinary Radiology Association, . [Details]
Williams, C,Hoppe, HJ,Rezgui, D,Strickland, M,Forbes, BE,Grutzner, F,Frago, S,Ellis, RZ,Wattana-Amorn, P,Prince, SN,Zaccheo, OJ,Nolan, CM,Mungall, AJ,Jones, EY,Crump, MP,Hassan, AB (2012) 'An Exon Splice Enhancer Primes IGF2:IGF2R Binding Site Structure and Function Evolution'. Science, 338 :1209-1213. [DOI] [Details]
Gibney, E.R. and Nolan, C.M.; (2010) 'Epigenetics and gene expression'. Heredity, 105 (1):4-13. [Details]
Shiel, RE,Mooney, CT,Brennan, SF,Nolan, CM,Callanan, JJ; (2010) 'Clinical and clinicopathological features of non-suppurative meningoencephalitis in young greyhounds in Ireland'. Veterinary Record, 167 :333-337. [DOI] [Details]
Tsalavouta, M., Astudillo, O., Byrnes, L. and Nolan, C.M.; (2009) 'Regulation of expression of zebrafish (Danio rerio) insulin-like growth factor 2 receptor: implications for evolution at the IGF2R locus'. Evolution and Development, 11 (5):546-558. Available Online [Details]
Nolan, C.M., O'Sullivan, F.M., Brabazon, D.C. and Callanan, J.J. ; (2009) 'Genomic imprinting in Canis familiaris'. Reproduction in Domestic Animals, 44 (Suppl.2):16-21. [DOI] [Details]
Hosback, S., Hardiman, O., Nolan, C. M., Maria, A. C. D. B. B. B. A., Gorman, G., Lynch, C., O'Toole, O., Jakeman, P., ; (2007) 'Circulating insulin-like growth factors and related binding proteins are selectively altered in amyotrophic lateral sclerosis and multiple sclerosis'. Growth Hormone & Igf Research, 17 (6):472-479. [Details]
O'Sullivan, FM,Murphy, SK,Simel, LR,McCann, A,Callanan, JJ,Nolan, CM; (2007) 'Imprinted expression of the canine IGF2R, in the absence of an anti-sense transcript or promoter methylation'. Evolution and Development, 9 :579-589. [Details]
Murphy, SK,Nolan, CM,Huang, ZQ,Kucera, KS,Freking, BA,Smith, TPL,Leymaster, KA,Weidman, JR,Jirtle, RL; (2006) 'Callipyge mutation affects gene expression in cis: A potential role for chromatin structure'. Genome Research, 16 :340-346. [DOI] [Details]
Nolan, C.M., McCarthy, K., Eivers, E., Jirtle, R.L., Byrnes, L.; (2006) 'Mannose 6-phosphate receptors in an ancient vertebrate, zebrafish'. Development, Genes and Evolution, 216 (3):144-151. [Details]
Murphy, SK,Freking, BA,Smith, TPL,Leymaster, K,Nolan, CM,Wylie, AA,Evans, HK,Jirtle, RL; (2005) 'Abnormal postnatal maintenance of elevated DLK1 transcript levels in callipyge sheep'. Mammalian genome : official journal of the International Mammalian Genome Society, 16 :171-183. [DOI] [Details]
Weidman JR, Murphy SK, Nolan CM, Dietrich FS and Jirtle RL; (2004) 'Phylogenetic footprint analysis of Igf2 in extant mammals'. Genome Research, 14 (9):1726-1732. [Details]
Eivers E, McCarthy K, Glynn C, Nolan CM, Byrnes L.; (2004) 'Insulin-like growth factor (IGF) signalling is required for early dorso-anterior development of the zebrafish embryo. '. International Journal of Developmental Biology, 48 (10):1131-1140. [Details]
Galvin, C.D., O. Hardiman, and C.M. Nolan,; (2003) 'IGF-1 receptor mediates differentiation of primary cultures of mouse skeletal myoblasts'. Molecular and Cellular Endocrinology, 200 :19-29. [Details]
Wylie, A.A., Pulford, D.J., McVie-Wylie, A.J., Waterland, R.A., Evans, H.K., Chen, Y.T., Nolan, C.M., Orton, T.C. and Jirtle, R.L.; (2003) 'Tissue-specific inactivation of murine M6P/IGF2R'. American Journal of Pathology, 162 :321-328. [Details]
Ayaso, E., C.M. Nolan, and L Byrnes,; (2002) 'Zebrafish insulin-like growth factor-1 receptor: molecular cloning and developmental expression'. Molecular and Cellular Endocrinology, 191 :137-148. [Details]
Oka, Y,Waterland, RA,Killian, JK,Nolan, CM,Jang, HS,Tohara, K,Sakaguchi, S,Yao, T,Iwashita, A,Yata, Y,Takahara, T,Sato, SI,Suzuki, K,Masuda, T,Jirtle, RL; (2002) 'M6P/IGF2R tumor suppressor gene mutated in hepatocellular carcinomas in Japan'. Hepatology, 35 :1153-1163. [DOI] [Details]
Nolan, C.M., Killian, J.K. Petitte, J.N. and Jirtle, R.L.; (2001) 'Imprint status of M6P/IGF2R and IGF2 in chickens'. Development Genes and Evolution, 211 :179-183. [Details]
Killian, J.K., Nolan, C.M., Stewart, N., Munday, B.L., Anderson, N.A., Nicol, S. and Jirtle, R.L.; (2001) 'Monotreme IGF2 expression and ancestral origin of genomic imprinting'. Journal of Experimental Zoology, 291 :205-212. [Details]
Killian, J.K., Nolan, C.M., Wylie, A.A., Li, T/., Vu, T.H., Hoffman, A.R. and Jirtle, R.L.; (2001) 'Divergent evolution in M6P/IGF2R imprinting from the Jurassic to the Quaternary'. Human Molecular Genetics, 10 :1721-1728. [Details]
Galvin C.D., Hardiman O. and Nolan C.M.; (2000) 'Binding of 125I-insulin-like growth factor-II to cells cultured in fetal bovine serum: a complication'. International Journal of Biochemistry & Cell Biology, (39):189-200. [Details]
Nolan, C.M. and M.A. Lawlor,; (1999) 'Variable accumulation of insulin-like growth factor II in mouse tissues deficient in insulin-like growth factor II receptor'. International Journal of Biochemistry & Cell Biology, 31 :1419-1431. [Details]
Nolan, C.M., Kyle, J.W., Watanabe, H. and Sly, W.S.; (1990) 'Binding of insulin-like growth factor II (IGF-II) by human cation-independent mannose 6-phosphate receptor/IGF-II receptor expressed in receptor-deficient mouse L cells'. Cell Regulation, 1 :197-213. [Details]
Kyle, J.W., Nolan, C.M., Oshima, A. and Sly, W.S.; (1988) 'Expression of human cation-independent mannose 6-phosphate receptor cDNA in receptor-negative mouse P388D1 cells following gene transfer'. Journal of Biochemistry, 263 :16230-16235. [Details]
Oshima, A., Nolan, C.M., Kyle, J.W. and Sly, W.S.; (1988) 'The human cation-independent mannose 6-phosphate receptor: cloning and sequence of the full-length cDNA and expression of functional receptor in COS cells'. Journal of Biological Chemistry, 263 :2553-2562. [Details]
Nolan, C.M., Creek, K.E., Grubb, J.H. and Sly, W.S.; (1987) 'Antibody to the phosphomannosyl receptor inhibits recycling of the receptor in fibroblasts'. Journal of Cellular Biochemistry, 35 :137-151. [Details]
Williams C, Hoppe HJ, Rezgui D, Strickland M, Forbes BE, Grutzner F, Frago S, Ellis RZ, Wattana-Amorn P, Prince SN, Zaccheo OJ, Nolan CM, Mungall AJ, Jones EY, Crump MP, Hassan AB (2012) 'An exon splice enhancer primes IGF2:IGF2R binding site structure and function evolution'. Science (New York, N.Y.), 338 (6111):1209-1213. [DOI] [Details]
Murphy S.K., Nolan C.M., Huang Z., Kucera K.S., Freking B.A., Smith T.P., Leymaster K.A., Weidman J.R., Jirtle R.L.; (2006) 'Callipyge mutation affects gene expression in cis: a potential role for chromatin structure'. GENOME RESEARCH, 16 (3):340-346. [Details]
Hartnett, L,Glynn, C,Nolan, CM,Grealy, M,Byrnes, L; (2010) 'Insulin-like growth factor-2 regulates early neural and cardiovascular system development in zebrafish embryos'. Development, 54 :573-583. [DOI] [Details]

Other Journals

Murphy, S.K., Freking, B.A., Smith, T.P.L., Leymaster, K., Nolan, C.M., Wylie, A.A., Evans, H.K. and Jirtle, R,L. ; (2005) 'Abnormal postnatal maintenance of elevated DLK1 transcript levels in callipyge sheep' Mammalian Genome 16 :171-183. [Details]

Conference Publications

Atencia, S; Mooney, CT; Shiel RE; Jones, BR; Nolan CM (2013) Muscular dystrophy in japanese spitz: description of the mutation and carrier detection using polymerase chain reaction Forum of the American College of Veterinary Internal Medicine (2013) [Details]
Nolan, CM,O'Sullivan, FM,Brabazon, DC,Callanan, JJ (2009) REPRODUCTION IN DOMESTIC ANIMALS Genomic Imprinting in Canis familiaris , pp.16-21 [DOI] [Details]
Walpole, C., O'Coigligh, O., Nolan, C. and Stewart, G. (2011) Investigation of UT-B urea transporter isoforms in the human gastrointestinal tract Epithelial and Membrane Transport Group meeting of The Physiological Society London, [Details]
Nolan, C.M. and W.S. Sly,; (1987) Intracellular traffic of the mannose 6-phosphate receptor and its ligands Proceedings of the Fourth International Symposium on the Immunobiology of Proteins and Peptides; 1987 [Details]

Published abstracts

Watanabe, H., Kyle, J.W., Nolan, C.M. and Sly, W.S.; (1988) The human man 6-P/IGF-II receptor: Structure/function analysis of the mutationally altered cytoplasmic domain following gene transfer and gene amplification in receptor-negative mouse L-cells. Published abstracts [Details]
Nolan, C.M., Kyle, J.W. and Sly, W.S.; (1988) Co-expression of IGF-II receptor activity and man 6-P receptor activity in receptor-negative mouse L-cells stably transformed with the human cation-independent man 6-P receptor cDNA. Published abstracts [Details]
Galvin, C.D., Nolan, C.M. and Hardiman, O.; (1998) The insulin-like growth factor family in skeletal muscle differentiation. Published abstracts [Details]
Galvin, C.D., Hardiman, O., and Nolan, C.M.; (1999) IGF-II binding to cultured cells: A complication. Published abstracts [Details]
Galvin, C.D., Hardiman, O., and Nolan, C.M.; (1999) Suramin inhibits differentiation of primary mouse myoblasts. Published abstracts [Details]
Ayaso, E., Nolan, C.M. and Byrnes, L.; (1999) Zebrafish IGF-I receptor expression during development. Published abstracts [Details]
Murphy, S.K., Wylie, A.A., McVie-Wylie, A.J., Pulford, D., Nolan, C.M., Orton, T.C. and Jirtle, R.L.; (2001) Development and characterization of a conditional M6p/Igf2r knock-out mouse. Published abstracts [Details]
Hosback, S., Doyle, M., Gorman, G., Hardiman, O., Nolan, C.M. and Jakeman P.; (2002) Circulating insulin-like growth factor system in chronic neurological disease states. Published abstracts [Details]
Nolan CM; (2004) Genomic imprinting and cancer. Published abstracts [Details]


Research Interests

The overall goal of our research is to understand the molecular basis of the related topics of animal development and evolution. While this is basic research, the results aim to increase our understanding of human cell and molecular physiology, so that this knowledge can be applied to the physiology of disease, and ultimately to the development of novel therapeutic strategies. There are currently several inter-related strands to our research:

IGF-signaling in development and disease
The IGFs are thought to control cell proliferation, differentiation, survival and apoptosis. We recently used primary mouse myoblasts to show that this system regulates differentiation of skeletal muscle, one of the most abundant tissues in vertebrates. We are now examining the involvement of IGF signaling in the muscle degeneration that is a feature of neurological diseases such as motor neuron disease. (Collaborators: Dr Orla Hardiman; Prof. Phil Jakeman).

Because of the complexity of the IGF-family, conventional gene knock-outs in mice are of limited value in determining the molecular basis of IGF-1 signaling during development. Therefore, we are using an alternative model vertebrate, the zebrafish Danio rerio, for such studies. The beauty of the zebrafish is that it allows the adoption of experimental embryology combined with genetic approaches. By inhibiting the function of the IGF-1receptor, we showed its importance in brain-, eye- and heart formation. (Collaborators: Dr Lucy Byrnes; Dr Maura Grealy)

Evolution of genomic imprinting
One of our approaches to this is to track the evolution of the structure and function of one imprinted gene, the CI-MPR, also known as the IGF2R in mammals. We have established the zebrafish as a model organism for this by determining the cDNA sequence, gene organization and expression pattern of the zebrafish receptor, and have identified its potential role in neural development. We are currently testing the ability of the zebrafish receptor to interact with IGF2 and are examining how this gene is regulated in zebrafish (Collaborators: Dr Lucy Byrnes; Dr Randy Jirtle; Prof Bass Hassan; Dr Oliver Zaccheo).

Although the evolution of genomic imprinting is a subject of much debate, the phenomenon has only been well documented in a limited number of mammalian orders (chiefly rodent, primates and artiodactyls). We are currently examining the occurrence of genomic imprinting in a novel order, the Carnivora. Using the canine M6P/IGF2R, we established the existence of genomic imprinting in the dog. We found some interesting differences between the imprinted canine IGF2R and its ortholog in mouse. We are pursuing this study and expanding the study to other candidate imprinted genes. The dog may be a unique model organism for analysis of imprinting in behavior/cancer/evolution of morphological diversity. This study is extremely timely because of recent advances in the genetics and genomics of dogs (Collaborators: Prof Sean Callanan; Dr Amanda McCann; Dr Susan Murphy; Dr Randy Jirtle).

The LUPA project

The EU-funded LUPA project is using dog genetics to study predisposition to complex diseases (those that involve several genes and/or environmental factors). In collaboration with Prof Sean Callanan (UCD School of Agriculture, Food Science and Veterinary Medicine), we are attempting to determine the basis of a novel brain inflammatory disease that has recently been identified in Irish greyhounds. 

Environmental epigenomics

We have recently begun a study aimed at developing zebrafish (Danio rerio) as a model system for examining the response of the epigenome to environmental agents. Zebrafish offer many advantages in toxicological studies, but in spite of the extensive use of zebrafish as a model of vertebrate development, its epigenome has not yet been well characterized.



Internal Collaborators

Prof John J Callanan, UCD School of Agriculture, Food Science and Veterinary Medicine

Dr Amanda McCann, UCD School of Medicine and Medical Science


External Collaborators

Dr Orla Hardiman, Department of Neurology, Beaumont Hospital, Dublin, Ireland.

Prof Phil Jakeman, School of Exercise Physiology, University of Limerick, Ireland.

Prof Bass Hassan, Weatherall Institute for Molecular Medicine, University of Oxford, UK.

Dr Susan K Murphy, Department of Obstetrics and Gynecology, Duke University Medical Center, Durham, North Carolina, USA.

European Canine Consortium (LUPA project members). Scientific Co-ordinator: Anne-Sophie Lequarre, Faculty of Veterinary Medicine, University of Liege, Belgium

Dr Randy L Jirtle, Department of Radiation Oncology, Duke University Medical Center, Durham, North Carolina, USA.