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Curriculum Vitae
Dennis D.M. O'Leary

Current Position and Address

Vincent J. Coates Chair in Molecular Neurobiology
Molecular Neurobiology Laboratory
The Salk Institute
10010 N. Torrey Pines Rd
La Jolla, California 92037
Tel: (858) 453-4100 ext. 1415 or 1664
FAX: 858-558-6207


For peer-reviewed published papers and citation data click following URL: and enter Dennis O'Leary for User Profile.



1972 - 1976      B.S., Biology (Biochemistry), University of Illinois, Urbana
1979 - 1983     Ph.D., Neural Sciences, Washington Univ Sch Med - St. Louis (W. Maxwell Cowan - advisor)
1983 - 1985   Postdoctoral Fellow, The Salk Institute, La Jolla, CA (W. Maxwell Cowan - advisor)


Academic Appointments

1990 - present Professor (1993-present), Associate Professor (1990-1993), Molecular
Neurobiology Laboratory, The Salk Institute, La Jolla, CA
1991 - present  Adjunct Professor (1993-present), Associate Adjunct Professor (1990-1993), Univ California Sch Med, San Diego, Dept of Neuroscience, Dept of Biology, Graduate Programs in Neuroscience and in Biology, and Program in Cognitive Science
1986-1990 Associate Professor of Neurobiology with tenure (1990), Assistant Prof (1986-1990), Dept of Neurological Surgery, joint appointments in Dept of Anatomy and Neurobiol, Dept of Neurology, Washington Univ School of Medicine, St. Louis
1985-1986 Research Assistant Professor, Developmental Neurobiology Laboratory, The Salk Institute, La Jolla, California

Honors and Awards

2010-2014 President ('12-'14), President-elect ('10-'12), Cajal Club Association
(SFN affiliated)
2007 Krieg Cortical Discoverer Prize, Cajal Club Association
2004-2011 Javits Investigator Award in Neuroscience, NINDS, NIH
2003 Fellow, American Association for the Advancement of Science
2003 Ariens-Kappers Award Laureate, Royal Netherlands Academy of Arts and Sciences
2003 Highly Cited Researcher Award, Neuroscience, Original Member, Institute for Scientific information
1999-2001 McKnight Investigator Award
1999 Capputto Memorial Award, Argentina Society for Neurochemistry
1996 Decade of the Brain Medalist, American Association of Neurological Surgeons
1996 Presidential Special Lecture, Society for Neuroscience Annual Meeting
1993-2002 Editor, Journal of Neuroscience
1991 Young Investigator Award, Society for Neuroscience
1988 Krieg Cortical Explorer Award, Cajal Club Association
1987-1989  McKnight Scholars Award
1987-1989  Alfred P. Sloan Research Fellow


Editorial Boards

2010 - present Frontiers Reviews Editor
2008 – present Associate Editor, Frontiers in Neuroscience / Frontiers in Neural Circuits
2008 – present Editorial Board, Journal of Experimental Neuroscience
2007 – present Editorial Board, Neural Development
1993 - 2002   Senior Editor, Journal of Neuroscience, Development / Plasticity / Repair Section (handled entire editorial process for approximately 800 new submissions per year, and managed editorial office)
1990 - 1993   Associate Editor, Journal of Neuroscience
1993 - 2002 Receiving Editor, Neuroscience Research
1993 - present Associate Editor, Learning and Memory
1995 - present Editorial Board, Current Opinion in Neurobiology
2000 – present  Highlights Advisory Board, Nature Reviews Neuroscience
2001 – present Editorial Board, Biological Signals in Neuroscience
2003 – present  Editorial Board, Neurosignals
2001 Editorial Committee,  Annual Review of Neuroscience
1995 - 1997  Associate Editor, Neuron
1994 - 1995  Editorial Board, Trends in Neuroscience
1994 Volume Editor, Development, Current Opinion in Neurobiology, 1994, V 4, #1
1993 Guest Editor, Perspectives on Developmental Neurobiology, 1993, Vol 1, #2.

National Institutes of Health Committee Service (partial list)

2007 Development and Embryology of the Nervous System, Consensus Terminology Workshop, NIH Neuroscience Blueprint   
2003 - present  GENSAT Advisory Committee, NIH
2003 MDCN7 (NDPR) Scientific Review Group, NIH / CSR
2002 National Eye Institute Special Emphasis Review panel
2002 NIH Postgenomics evaluation workshop, Laguna Beach
2000 Chair, ZNS1 SRB-H SS Technical Evaluation Group-RFP 00-05, NINDS
2000 Special Emphasis Panel, ZRG1 SSS-R 03 S, NIH / CSR
1998 NICHD Board of Scientific Directors, Ad hoc reviewer of intramural program
1998 NINDS Board of Scientific Directors, Ad hoc reviewer of intramural program
1998 Neuro C Scientific Review Group, NIH / CSR
1997 NIH Working Group on Neuroscience Peer Review Reorganization
1992 - 1996 Visual Sciences B Scientific Review Group, NIH / CSR, Regular member
1993 National Institute of Mental Health Workshop (Redbook)
1991 National Eye Institute Workshop, Vice-Chair
1990 - 1991 NSP-B Scientific Review Group, Ad Hoc, NIH
1990 Visual Sciences A1 AHR-M1/M2 Scientific Review Group, NIH / CSR
1990 Visual Sciences B AHR-M3 Scientific Review Group, NIH / CSR

Other Professional Service (partial list)

2005 - present Howard Hughes Medical Institute, International Program Committee, Canada and Latin America
2004 - present Kirsch Foundation and Glaucoma Research Foundation Advisory Board
2004 External Review Committee, CNLM, Univ California-Irvine
2001 – present  External Advisory Comm., West Virginia Univ Med Cntr
2000 Organizer and Chair, Sam Hersch Symposium on Cerebral Palsy
1994 - present  Scientific Advisory Committee, Australasian Spinal Research Trust
1993 - present Organizer and Instructor (1993-1996), Instructor (current) Cold Spring Harbor Laboratory Summer Course - Developmental Neurobiology
1993 Co-organizer of NATO/NSF Conference "Neurospecificity: Maps and Molecules", Cargese, Corsica (May 23-29, 1993)

Professional Societies

American Association for the Advancement of Science
Society for Neuroscience
FASEB/American Association of Anatomists
Cajal Club
International Brain Research Organization

Service in Professional Societies

2010 – present  President-elect, Cajal Club
1996 - 1997  Cajal Club, Program Committee (chair, ‘97), Awards Committee, Officer Nomination Committee (chair, ‘96)
1992 - 1996 Executive Committee of the American Association of Anatomists
1986 - 1989  Advisory Committee of Young Anatomists to Executive Committee of American Association of Anatomists

Major Salk Institute Committees (current service on standing committees)

Appointments and Promotions Committee (APCOM)

Academic Planning and Facilities (APF)

Innovative Grants Committee (Chairman)

Summary of Research Contributions

Dennis O'Leary studies the development and plasticity of the vertebrate nervous system. His work focuses on two broad issues: development of neural connectivity and development of the cerebral cortex. To address these issues, O'Leary and his research group use genetic manipulations in mice and chick embryos, in vivo analyses and in vitro assays, and a range of techniques including molecular and cell biology, biochemistry, imaging, and behavioral testing.

O'Leary's goal is to understand fundamental developmental events in the nervous system, to establish a framework to design effective strategies to overcome birth defects, neurological diseases and disorders, and injury.

Development of Neural Connectivity

O'Leary discovered fundamental mechanisms involved in the development of neural connectivity that challenged and supplanted dogma at the time. These discoveries include (i) exuberant axonal projections, (ii) selective axon elimination and the underlying mechanisms, and (iii) interstitial axonal branching as the predominant mechanism of target recognition.

O'Leary's work demonstrated that specific axonal connections arise from initially exuberant projections followed by pruning through a mechanism of selective large-scale axon degeneration. He has established that this naturally-occurring axon pruning is a fundamental characteristic of vertebrate neural development and occurs through a caspase-dependent mechanism that is morphologically similar but molecularly distinct from “Wallerian” degeneration. O'Leary also has shown that the de novo interstitial branching of primary axons is the predominant mechanism of target recognition and innervation in vertebrates, and has determined cellular and molecular mechanisms of its control.

O'Leary's studies on the development of topographic maps in the visual system have defined the predominant mechanisms of topographic axon mapping and novel functions of ephrin-Eph signaling in axon guidance and mapping. This work has established novel cellular and molecular mechanisms for map development, including topographic axon branching as a crucial event in map development that requires unique functions for axon guidance molecules that are distinct from their control of growth cone targeting, and the relative roles of patterned activity and molecular guidance molecules. O'Leary has made prominent contributions to defining functions for ephrins and Ephs in axon guidance and mapping, has shown that distinct ephrins and Ephs regulate interstitial axon branching and direct branch extension, act bifunctionally as repellents and attractants and bidirectionally through forward- and reverse-signaling. His work is the first to determine receptor mechanisms of ephrin-A reverse signaling and the modulation of EphA forward signaling by complexing with neurotrophin receptors.

Development of the Cerebral Cortex

O'Leary has made pioneering contributions to cortical development, especially patterning of the neocortex into functionally-specialized areas. He has pioneered the role of intrinsic genetic mechanisms that specify area identities of cortical progenitors and their neuronal progeny, including the first identification of transcription factors that specify the area identity of progenitors and unique functions for morphogens in progenitor differentiation and regulation of cortical size. Based on this work, O'Leary has proposed a model of neuronal specification for higher brain centers distinct from the model developed from studies of the spinal cord. His group has also established genetic mechanisms for the determination of regional-fate in the cerebral cortex, for example the genetic regulation of the fate decision in telencephalic progenitors to generate olfactory cortex versus neocortex.

Complementing the genetic regulation of forebrain patterning, O'Leary has demonstrated mechanisms of plasticity in area patterning of the neocortex, including novel roles for afferent based mechanisms. Recently, he and his group have found a form of “top-down” plasticity not previously recognized in which the intrinsic genetic specification of cortical sensory areas determines the representation of the sensory periphery within them and drives the subsequent re-patterning of thalamic sensory nuclei to match their cortical counterparts.


For peer-reviewed published papers and citation data click following URL: and enter Dennis O'Leary for User Profile.

1. O'Leary, DDM, Fricke, RA, Stanfield, BB, Cowan, WM 1979 Changes in the associational afferents to the dentate gyrus in absence of its commissural input. Anatomy and Embryology 156: 283-299.

2. O'Leary, DDM, Stanfield, BB, Cowan, WM 1980 Evidence for the sprouting of the associational fibers to the dentate gyrus following removal of the commissural afferents in adult rats. Anatomy and Embryology 156: 151-161.

3. O'Leary, DDM, Stanfield, BB, Cowan, WM 1981 Evidence that early postnatal restriction of the cells of origin of the callosal projection is due to the elimination of axonal collaterals rather than to the death of neurons. Developmental Brain Research 1:607-617.

4. Stanfield, BB, O'Leary, DDM, Fricks, C 1982 Selective collateral elimination in early postnatal development restricts cortical distribution of rat pyramidal tract neurones. Nature 298: 371-373.

5. Gerfen, CR, O'Leary, DDM, Cowan, WM 1982 A note on the transneuronal transport of wheat germ agglutinin conjugated horseradish peroxidase in the avian and rodent visual systems. Experimental Brain Research 48: 443-448.

6. O'Leary, DDM, Cowan, WM 1982 Further studies on the development of the isthmo-optic nucleus with special reference to the occurrence and fate of ectopic and ipsilaterally projecting neurons. Journal of Comparative Neurology 212: 399-416.

7. Swanson, LW, Lindstrom, J, Tzartos, S, Schmued, LC, O'Leary, DDM, Cowan, WM 1983 Immunohistochemical localization of monoclonal antibodies to the nicotinic acetylcholine receptor in the midbrain of the chick. Proc. Natl. Acad. Sci. USA 80: 4532-4536.

8. O'Leary, DDM, Gerfen, CR, Cowan, WM 1983 The development and restriction of the ipsilateral retinofugal projection in the chick. Developmental Brain Research 10: 93-109.

9. O'Leary, DDM, Cowan, WM 1983 Topographic organization of certain tectal afferent and efferent connections can develop normally in the absence of retinal input. Proc. Natl. Acad. Sci. USA 80: 6131-6135.

10. O'Leary, DDM, Cowan, WM 1984 Survival of isthmo-optic neurons after early removal of one eye. Developmental Brain Research 12: 293-310.

11. Fawcett, JW, O'Leary, DDM, Cowan, WM 1984 Activity and the control of ganglion cell death in the rat retina. Proc. Natl. Acad. Sci. USA 81: 5589-5593.

12. Cowan, WM, Fawcett, JW, O'Leary, DDM, Stanfield, BB 1984 Regressive events in neurogenesis. Science 225: 1258-1265.

13. Cowan, WM, O'Leary, DDM 1984 Cell death and process elimination: the role of regressive phenomena in the development of the vertebrate nervous system. In: Medicine, Science and Society: Symposia Celebrating the Harvard Medical School Bicentennial, K.J. Isselbacher (ed.), Wiley Press: New York, pp. 643-668.

14. Stanfield, BB, O'Leary, DDM 1985 Fetal occipital cortical neurons transplanted to rostral cortex develop and maintain a pyramidal tract axon. Nature 313: 135-137.

15. Crespo, D, O'Leary, DDM, Cowan, WM 1985 Changes in the number of optic nerve axons during late prenatal and postnatal development of the albino rat. Developmental Brain Research 19: 129-134.

16. O'Leary, DDM, Stanfield, BB 1985 Occipital cortical neurons with transient pyramidal tract axons extend and maintain collaterals to subcortical but not intracortical targets. Brain Research 336: 326-333.

17. Stanfield, BB, O'Leary, DDM 1985 The transient corticospinal projection from the occipital cortex during the postnatal development of the rat. Journal of Comparative Neurology 238: 236-248.

18. Fawcett, JW, O'Leary, DDM 1985 The role of electrical activity in the formation of topographic maps. Trends in Neuroscience 8: 201-206.

19. Cowan, WM, Fawcett, JW, O'Leary, DDM, Stanfield, BB 1985 Regressive events in neurogenesis. In: Neuroscience, P.H. Abelson, E. Butz, and S.H. Snyder (eds), Amer Assoc Adv Science, Washington D.C., pp.13-29.

20. O'Leary, DDM, Stanfield, BB 1986 A transient pyramidal tract projection from the visual cortex in the hamster and its removal by selective collateral elimination. Developmental Brain Research27: 87-99.

21. O'Leary, DDM, Crespo, D, Fawcett, JW, Cowan, WM 1986 The effect of intraocular tetrodotoxin on the postnatal reduction in numbers of optic nerve axons in the rat. Developmental Brain Research, 30: 96-103.

22. O'Leary, DDM, Fawcett, JW, Cowan, WM 1986 Topographic targeting errors in the retinocollicular projection and their elimination by selective ganglion cell death. Journal of Neuroscience 6: 3692-3705.

23. Stanfield, BB, Nahin, B, O'Leary, DDM 1987 A transient post-mammillary component of the rat fornix during development: implications for interspecific differences in mature axonal projections. Journal of Neuroscience7: 3350-3361.

24. Porter, LL, Cedarbaum, JM, O'Leary, DDM, Stanfield, BB, Asanuma, H 1987 The physiological identification of pyramidal tract neurons within transplants in the rostral cortex taken from the occipital cortex during development. Brain Research 436: 136-142.

25. O'Leary, DDM 1987 The remodeling of early axonal projections through the selective loss of neurons and axon collaterals. In: Selective Neuronal Death, M. O'Connor (ed.), (Ciba Foundation Symposium No. 126), Wiley: Chichester, pp. 113-142.

26. Stanfield, BB, O'Leary, DDM 1988 Neurons in the subiculum with transient postmammillary collaterals during development maintain projections to the mammillary complex. Experimental Brain Research 72: 185-190.

27. O'Leary, DDM, Terashima, T 1988 Cortical axons branch to multiple subcortical targets by interstitial axon budding: implications for target recognition and "waiting periods." Neuron1: 901-910.

28. O'Leary, DDM, Stanfield, BB 1989 Selective elimination of axons extended by developing cortical neurons is dependent on regional locale. Experiments utilizing fetal cortical transplants. Journal of Neuroscience 9:2230-2246.

29. Nakamura, H, O'Leary, DDM 1989 Inaccuracies in initial growth and arborization of chick retinotectal axons followed by course corrections and axon remodeling to develop topographic order. Journal of Neuroscience 9:3776-3795.

30. O'Leary, DDM 1989 Do cortical areas emerge from a proto-cortex? Trends in Neuroscience 12:400-406.

31. Simon, DK, O'Leary, DDM 1990 Limited topographic specificity in the targeting and branching of mammalian retinal axons. Developmental Biology 137:125-134.

32. Heffner, CA, Lumsden, AGS, O'Leary, DDM 1990 Target control of collateral extension and directional axon growth in the mammalian brain. Science 247:217-220.

33. O'Leary, DDM, Bicknese, AR, De Carlos, JA, Heffner, CD, Koester, SE, Kutka, LJ, Terashima, T 1990 Target selection by cortical axons: Alternative mechanisms to establish axonal connections in the developing brain. Cold Spring Hbr Symp Quant Biol. 55:453-468.

34. O'Leary, DDM 1990 Growth, branching, and target selection by developing cortical axons. In: FESN Symp #8: Neural Development, T Wiesel, L Katz and D Anderson (Eds.), Elsevier: Amsterdam.

35. Simon, DK, O'Leary, DDM 1991 Relationship of retinotopic ordering of axons in the optic pathway to the formation of visual maps in central targets. Journal of Comparative Neurology307:393-404.

36. Schlaggar, BL, O'Leary, DDM 1991 Potential of visual cortex to develop arrays of functional units unique to somatosensory cortex. Science252:1556-1560.

37. O'Leary, DDM, Heffner, CD, Kutka, L, Lopez-Mascaraque, L, Missias, A, Reinoso, BS 1991 A target-derived chemoattractant controls the development of the corticopontine projection by a novel mechanism of axon targeting. Development2:123-130.

38. O'Leary, DDM, Schlaggar, BL, Stanfield, BB 1992 The specification of sensory cortex: Lessons from cortical transplantation. Experimental Neurology 115:121-126.

39. Simon, DK, O'Leary, DDM 1992 Development of topographic order in the mammalian retinocollicular projection. Journal of Neuroscience 12:1212-1232.

40. De Carlos, JA, O'Leary, DDM 1992 Growth and targeting of subplate axons and establishment of major cortical pathways. Journal of Neuroscience 12:1194-1211.

41. Koester, SE, O'Leary, DDM 1992 Functional classes of cortical projection neurons develop dendritic distinctions by class-specific sculpting of an early common pattern. Journal of Neuroscience 12:1382-1394.

42. Simon, DK, O'Leary, DDM 1992 Influence of position along the medial-lateral axis of the superior colliculus on the topographic targeting and survival of retinal axons. Developmental Brain Research 69:167-172.

43. Simon, DK, Prusky, GT, O'Leary, DDM, Constantine-Paton, M 1992 NMDA receptor antagonists disrupt the development of a mammalian neural map. Proc. Natl. Acad. Sci. USA, 89:10593-10597.

44. Simon, DK, O'Leary, DDM 1992 Responses of retinal axons in vivo and in vitro to molecules encoding position in the embryonic superior colliculus. Neuron 9:977-989.

45. O'Leary, DDM 1992 Development of connectional diversity and specificity in the mammalian brain by the pruning of collateral projections. Current Opinion in Neurobiology 2:70-77.

46. O'Leary, DDM, Simon, DK 1992 Position-encoding molecules and the development of retinal maps. Seminars in Neuroscience 4:365-371.

47. Schlaggar, BL, De Carlos, JA, O'Leary, DDM 1993 Acetylcholinesterase as an early marker of the differentiation of dorsal thalamus in embryonic rats. Developmental Brain Research 75:19-30.

48. Schlaggar, BL, Fox, K, O'Leary, DDM 1993 Postsynaptic control of plasticity in developing somatosensory cortex. Nature364:623-626.

49. Koester, SE, O'Leary, DDM 1993 Connectional distinction between callosal and subcortically projecting cortical neurons is determined before axon extension. Developmental Biology 160:1-14.

50. Tuttle, R, O'Leary, DDM 1993 Cortical connections form in culture. Current Biology 3:70-72.

51. O'Leary, DDM 1993 Adding neurons to the adult mammalian brain. Proc Natl Acad Sci USA 90:2101-2102.

52. O'Leary, DDM, Koester, SE 1993 Development of projection neuron types, axonal pathways and patterned connections of the mammalian cortex. Neuron 10:991-1006.

53. Schlaggar, BL, O'Leary, DDM 1993 Patterning of the barrelfield in somatosensory cortex with implications for the specification of neocortical areas. Perspectives on Developmental Neurobiology 1:81-92.

54. Shatz, CJ, O'Leary, DDM for the panel. 1993 Repair and replacement to restore sight report from panel on ganglion cell connectivity. Archs. Opthalmol., 111:472-477.

55. Simon, DK, Roskies, AL, O'Leary, DDM 1994 Plasticity in the development of topographic order in the mammalian retinocollicular projection. Developmental Biology 162:384-393.

56. Bicknese, AR, Sheppard, AM, O'Leary, DDM, Pearlman, AL 1994 Thalamocortical axons extend along a chondroitin sulfate proteoglycan-enriched pathway coincident with the neocortical subplate and distinct from the efferent path. Journal of Neuroscience 14:3500-3510.

57. Schlaggar, BL, O'Leary, DDM 1994 Early development of the somatotopic map and barrel patterning in rat somatosensory cortex. Journal of Comparative Neurology 346:80-96.

58. Roskies, A, O'Leary, DDM 1994 Control of topographic retinal axon branching by inhibitory membrane-bound molecules. Science 265:799-802.

59. Sato, M, Lopez-Mascaraque, L, Heffner, CD, O'Leary, DDM 1994 Action of a diffusible target-derived chemoattractant on cortical axon branch induction and directed growth. Neuron 13:791-803.

60. Koester, SE, O'Leary, DDM 1994 The corpus callosum is pioneered by early generated neurons of the cingulate cortex. Journal of Neuroscience 14:6608-6621.

61. O'Leary, DDM, Schlaggar, BL, Tuttle, R 1994 Specification of neocortical areas and thalamocortical connections. Annual Review of Neuroscience 17: 419-439.

62. Lumsden, AGS, O'Leary, DDM 1994 Development: editorial overview. Current Opinion in Neurobiology4: 1-7.

63. O'Leary, DDM, Ruff, NL, Dyck, RL 1994 Development, critical period plasticity, and adult reorganizations of mammalian somatosensory systems. Current Opinion in Neurobiology 4: 535-544.

64. O'Leary, DDM 1994 Attractive guides for axons. Nature 371:15-16.

65. Koester, SE, O'Leary, DDM 1994 Development of projection neurons in mammalian cerebral cortex. Progress in Brain Research 102:207-215.

66. Tuttle, R, Braisted, J, Schlaggar, BL, O'Leary, DDM 1995 Maturation-dependent upregulation of growth promoting molecules in developing cortical plate controls thalamic and cortical neurite growth. Journal of Neuroscience 15:3039-3052.

67. De Carlos, JA, Schlaggar, BL, O'Leary, DDM 1995 Development of acetylcholinesterase-positive thalamocortical and basal forebrain afferents to embryonic rat cortex. Experimental Brain Research104:385-401.

68. Roskies, A, Friedman, G, O'Leary, DDM 1995 Molecules and mechanisms in the development of retinotopic maps. Perspectives on Developmental Neurobiology 3:63-75.

69. O'Leary, DDM, Borngasser, DJ, Fox, K, Schlaggar, BL 1995 Plasticity in the development of neocortical areas. In: Development of the Cerebral Cortex. Ciba Fnd Symp No. 193. G. Bock and C. Blakemore, Eds. pp.214-230.

70. Bastmeyer, M, O'Leary, DDM 1996 Dynamics of target recognition by interstitial axon branching along developing cortical axons. Journal of Neuroscience 16:1450-1459.

71. Moriyoshi, K, Richards, L, Akazawa, C, O'Leary, DDM, Nakanishi, S 1996 Labeling neural cells using adenoviral gene transfer of membrane-targeted GFP. Neuron16:255-260.

72. Fox, K, Schlaggar, BL, O'Leary, DDM 1996 Glutamate receptor blockade at cortical synapses disrupts development of thalamocortical and columnar organization of somatosensory cortex. Proc. Natl. Acad. Sci. USA 93:5584-5589.

73. Friedman, GC, O'Leary, DDM 1996 Retroviral misexpression of engrailed genes in the chick optic tectum perturbs the topographic targeting of retinal axons. Journal of Neuroscience 16:5498-5509.

74. Daston, MM, Bastmeyer, M, Rutishauser, U, O'Leary, DDM 1996 Spatially restricted increase in polysialic acid influences target recognition by corticospinal axons. Journal of Neuroscience 16:5488-5497.

75. Nakamoto, M, Cheng, H-J, Friedmann, GC, McLaughlin, T, Hansen, MJ, Yoon, CH, O'Leary, DDM, Flanagan, JG 1996 Topographically specific effects of ELF-1 on retinal axon guidance in vitro and retinal axon mapping in vivo. Cell86:755-766.

76. O'Leary, DDM 1996 Areal specialization of the developing neocortex: differentiation, developmental plasticity and genetic specification. In: Nobel Symposium: The Lifespan Development of Individuals -Behavioral, Neurobiological and Psychosocial Perspectives. D. Magnusson, Ed., Elsevier Press. pp.23-37.

77. Friedman, GC, O'Leary, DDM 1996 Eph receptor tyrosine kinases and their ligands in neural development. Current Opinion in Neurobiology 6:127-133.

78. O'Leary, DDM, Bastmeyer, M, Daston, ME, Koester, SE, Richards, L, Yee, K 1996 Development of Cortical Output Projections: Axon Guidance, Target Recognition, and Plasticity. In: Integrative and Molecular Approach to Brain Function: Uehara Memorial Foundation Symposium-1996. M. Ito and Y. Miyashita, Eds. Elseveir:Amsterdam. pp.165-178.

79. Richards, LR, Koester, SE, Tuttle, R, O'Leary, DDM 1997 Directed growth of early cortical axons is influenced by a chemoattractant released from an intermediate target. Journal of Neuroscience 17:2445-2458.

80. Braisted, JE, McLaughlin, T, Wang, HU, Friedman, GC, Anderson, DJ, O'Leary, DDM 1997 Graded and lamina-specific distributions of ligands of EphB receptor tyrosine kinases in the developing retinotectal system. Developmental Biology 191:14-28.

81. Chenn, A, Braisted, JE, McConnell, SK, O'Leary, DDM 1997 Development of the Cerebral Cortex: Mechanisms Controlling Cell Fate, Laminar and Areal Patterning, and Axonal Connectivity. In, Molecular and Cellular Approaches to Neural Development, pp. 440-473. Oxford Univ. Press. W.M. Cowan, L. Zipursky, and T. Jessell, Eds.

82. Bastmeyer, B, Daston, MM, Possel, H, O'Leary, DDM 1998 Collateral branch formation related to cellular structures in the axon tract during corticopontine target recognition. Journal of Comparative Neurology 392:1-18.

83. Tuttle, R, Braisted, JE, Richards, LJ, O'Leary, DDM 1998 Retinal axon guidance by region specific cues in the diencephalon. Development 125:791-801.

84. Frisén J1, Yates PA1, McLaughlin T, Friedman GC, O'Leary DDM2, Barbacid M2 1998 Ephrin-A5 (AL-1/RAGS) is essential for proper retinal axon guidance and topographic mapping in the mammalian visual system. Neuron 20:235-243. [1, co-first authors; 2, co-communicating authors]

85. Tuttle, R, O'Leary, DDM 1998 Neurotrophins rapidly modulate growth cone response to the axon guidance molecule, collapsin-1. Mol Cell Neurobio11:1-8.

86. Braisted, JE, Tuttle, R, O'Leary, DDM 1999 Thalamocortical axons are influenced by chemorepellent and chemoattractant activities localized to decision points along their path. Developmental Biology 208:430-440.

87. Tuttle, R1, Nakagawa, Y1, Johnson, JE, O'Leary, DDM 1999 Defects in thalamocortical axon pathfinding correlate with altered cell domains in embryonic Mash-1 deficient mice. Development126:1903-1916. [1, co-first authors]

88. O'Leary, DDM, Wilkinson, DG 1999 Eph receptors and ephrins in neural development. Current Opinion in Neurobiology 9:55-73.

89. O'Leary, DDM, Yates, P, McLaughlin, T, 1999 Mapping sights and smells in the brain: Distinct mechanisms to achieve a common goal. Cell96:255-269.

90. McLaughlin, T, O'Leary, DDM 1999 Functional consequences of coincident expression of EphA receptors and ephrin-A ligands. Neuron22:636-639.

91. Bertuzzi, S, Hindges, R, Mui, S, O'Leary, DDM, Lemke, G 1999 The homeodomain protein Vax1 is required for axon guidance and development of major axon tracts in the brain. Genes and Development13:3092-3105.

92. Yee, KT, Simon, HH, Tessier-Lavigne, M, O'Leary, DDM 1999 Extension of long leading processes and neuronal migration in the mammalian brain directed by the chemoattractant netrin-1. Neuron 24:607-622.

93. Nakagawa, Y, Johnson, JE, O'Leary, DDM 1999 Graded and areal expression patterns of regulatory genes and cadherins in embryonic neocortex independent of thalamocortical input. Journal of Neuroscience 19:10877-10885.

94. Bishop, KM, Goudreau, G, O'Leary, DDM 2000 Emx2 and Pax6 regulate area identity in the mammalian neocortex. Science288:344-349.

95. Ringstedt, T1, Braisted, JE1, Brose, K, Goodman, C, Kidd, T, Tessier-Lavigne, M, O'Leary, DDM 2000 Slit inhibition of retinal axon growth and its role in retinal axon pathfinding and innervation patterns within the diencephalon Journal of Neuroscience 20:4983-4991. [1, co-first authors]

96. Brown, A1, Yates, PA1, Burrola, P, Ortuño, D, Vaidya, A, Jessell, TM, Pfaff, SL, O'Leary, DDM2, Lemke, G2 2000 Topographic mapping from the retina to the midbrain is controlled by relative but not absolute levels of EphA receptor signaling. Cell102:77-88. [1, co-first authors; 2, co-senior authors].

97. Braisted, JE, Catalano, SM, Stimac, R, Kennedy, T, Tessier-Lavigne, M, Shatz, CJ, O'Leary, DDM 2000 Netrin-1 promotes thalamic axon growth and is required for proper development of the thalamocortical projection. Journal of Neuroscience 20:5792-5801.

98. Liu, Q1, Dwyer, ND1, O'Leary, DDM 2000 Differential expression of COUP-TF1, CHL-1 and two novel genes in developing neocortex identified by differential display PCR. Journal of Neuroscience20:7682-7690. [1, co-first authors]

99. Erkman, L, Yates, PA, McLaughlin, T, McEvilly, RJ, Whisenhunt, T, O'Connell, SM, Krones, AL, Kirby, MA, Rapaport, DH, Bermingham, JR, O'Leary, DDM1, Rosenfeld, MG1 2000 A POU domain transcription factor dependent program regulates axon pathfinding in the vertebrate visual system. Neuron28:779-792. [1, co-communicating authors]

100. Dwyer, ND, O'Leary, DDM 2001 Tbr1 conducts the orchestration of early cortical development. Neuron29:309-311.

101. Nakagawa, Y, O'Leary, DDM 2001 Combinatorial patterns of expression of LIM-homeodomain and other regulatory genes parcellate developing thalamus. Journal of Neuroscience, 21:2711-2725.

102. Simon, HH, Saueressig, H, Wurst, W, Goulding, MD, O'Leary, DDM 2001 Fate of midbrain dopaminergic neurons controlled by the engrailed genes. Journal of Neuroscience, 21:3126-3134.

103. Gesemann, M1, Litwack, ED1, Yee, KT, O'Leary, DDM 2001 Identification of candidate genes for controlling development of the basilar pons by differential display PCR. Mol Cell Neurobio18:1-12. [1, co-first authors]

104. Yates, PA1, Roskies, AR1, McLaughlin, T, O'Leary, DDM 2001 Topographic specific axon branching controlled by ephrin-As is the critical event in retinotectal map development. Journal of Neuroscience, 21:8548-8563. [1, co-first authors]

105. Mui, SH, Hindges, R, O'Leary, DDM, Lemke, G, Bertuzzi, S 2002 The homeodomain protein Vax2 patterns the dorsoventral and nasotemporal axes of the eye. Development 129:797-804.

106. O'Leary, DDM, Nakagawa, Y 2002 Patterning centers, regulatory genes and extrinsic mechanisms controlling arealization of the neocortex. Curr. Op. Neurobio.,12:14-25.

107. Woodring, PJ, Litwack, ED, O'Leary, DDM, Lucero, GR, Wang, JYJ, Hunter, T 2002 Modulation of the F-actin cytoskeleton by c-Abl tyrosine kinase in cell spreading and neurite extension. Journal of Cell Biology, 156:879-892.

108. Hindges, R1, McLaughlin, T1, Genoud, N, Henkemeyer, M, O'Leary, DDM 2002 EphB forward signaling controls directional branch extension and arborization required for dorsal-ventral retinotopic mapping. Neuron, 35: 475-487. [1, co-first authors]

109. Bishop, KM, Rubenstein JRL, O'Leary, DDM 2002 Distinct actions of Emx1, Emx2 and Pax6 in regulating the specification of areas in the developing neocortex. Journal of Neuroscience, 22:7627-7638.

110. Burden, S, Berg, D, O'Leary, DDM 2003 Chapter 18: Target Selection, Topographic Maps, and Synapse Formation. pp. 469-498. In: Fundamentals of Neuroscience, 2nd Edition. (Squire, Bloom, McConnell, Roberts, Spitzer, Zigmond, eds.). Academic Press.

111. McLaughlin, T, Hindges, R, O'Leary, DDM 2003 Regulation of axial patterning of the retina and its topographic mapping in the brain. Curr Op Neurobio, 13:57-69.

112. Bishop, KM, Garel S, Nakagawa, Y, Rubenstein JRL, O'Leary, DDM 2003 Emx1 and Emx2 cooperate to regulate cortical size, lamination, neuronal differentiation, development of cortical efferents, and thalamocortical pathfinding. Journal of Comparative Neurology 457:345-360.

113. McLaughlin, T, Hindges, R, Yates, PA, O'Leary, DDM 2003 Bifunctional action of ephrin-B1 as a repellent and attractant to control bidirectional branch extension in dorsal-ventral retinotopic mapping. Development 130:2407-2418.

114. Leingartner, A, Richards, LJ, Dyck, RH, Akazawa, C, O'Leary, DDM 2003 Cloning, patterned cortical expression, and adenovirus-mediated ectopic expression of Emx2 to assess its regulation of area-specific thalamocortical projections. Cerebral Cortex 13:648-660.

115. Nakagawa Y, O'Leary DDM 2003 Dynamic patterned expression of orphan nuclear receptor genes RORa and RORb in developing mouse forebrain. Developmental Neuroscience, 25:234-244.

116. McLaughlin, T1, Torborg, CL1, Feller, MB2, O'Leary, DDM2 2003 Retinotopic map refinement requires spontaneous retinal waves during a brief critical period of development. Neuron, 40:1147-1160. [1, co-first authors; 2, co-communicating authors]

117. Yates, PA, Holub, AD, McLaughlin, T, Sejnowski, TJ, O'Leary, DDM 2004 Computational modeling of retinotopic map development to define contributions of EphA-ephrin-A gradients, axon-axon interactions and patterned activity. Journal of Neurobiology 59:95-113.

118. Litwack, ED, Babey, R, Buser, R, Gesemann, M, O'Leary, DDM 2004 Identification and characterization of two novel brain-derived immunoglobulin superfamily members that display a unique structural organization. Mol Cell Neurobio, 25:263-274.

119. Thuret, S, Bhatt, L, O'Leary, DDM, Simon, HH 2004 Identification and developmental analysis of genes expressed by dopaminergic neurons of the substantia nigra pars compacta. Mol Cell Neurobio, 25:394-405.

120. Hamasaki, T1, Leingartner, A1, Ringstedt, T, O'Leary DDM 2004 EMX2 regulates sizes and positioning of the primary sensory and motor areas in neocortex by direct specification of cortical progenitors. Neuron 43:359-72. [1, co-first authors]

121. Pak, W, Hindges, R, Lim, Y-S, Pfaff, SL, O'Leary, DDM 2004 Magnitude of binocular vision controlled by Islet-2 repression of a genetic program that specifies laterality of retinal axon pathfinding. Cell 119:567-578.

122. O'Leary, DDM, McLaughlin, T 2005 Mechanisms of retinotopic map development: Ephs, ephrins, and spontaneous correlated retinal activity. In: Development, Dynamics and Pathology of Neuronal Networks: From Molecules to Functional Circuits, J. van Pelt, M. Kamermans, C.N. Levelt, A. van Ooyen, G.J.A. Ramakers, P.R. Roelfseman (Eds.), Progress in Brain Research Vol. 147. Elsevier, Amsterdam, pp. 43 - 65.

123. Simon, HH, Scholtz, C, O'Leary, DDM 2005 Engrailed genes control developmental fate of serotonergic and noradrenergic neurons in mid- and hindbrain in a gene dose-dependent manner. Mol Cell Neurobio 28:96-105.

124. McLaughlin, T, O'Leary, DDM 2005 Molecular gradients and development of retinotopic maps. Annual Review of Neuroscience28:327-356.

125. Luo, L., O'Leary, DDM 2005 Axon retraction and degeneration in development and disease. Annual Review of Neuroscience 28:127-156.

126. Kroll, TT, O'Leary, DDM 2005 Ventralized dorsal telencephalic progenitors in Pax6 mutant mice generate GABA interneurons of a lateral ganglionic eminence fate. Proc. Natl. Acad. Sci. USA 102:7374-7379.

127. Quina, L, Pak, W, Lanier J, Banwait P, Gratwick K, Liu L, Velasquez T, O'Leary DDM, Goulding M, Turner EE 2005 Brn3a-expressing retinal ganglion cells project specifically to thalamocortical and collicular visual pathways. Journal of Neuroscience 25:11595-11604.

128. Takeuchi, A, O'Leary, DDM 2006 Radial migration of superficial layer cortical neurons controlled by novel Ig cell adhesion molecule, MDGA1. Journal of Neuroscience 26:4460-4464.

129. O'Leary DDM, Borngasser D 2006 Cortical ventricular zone progenitors and their progeny maintain spatial relationships and radial patterning during preplate development indicating an early protomap. Cerebral Cortex 16:i46-i56.

130. Hoopfer, ED1, McLaughlin, T1, Watts, RJ, Schuldiner, O, O'Leary, DDM2, Luo, L22006 Wlds protection distinguishes axon degeneration following injury from naturally-occurring developmental pruning. Neuron 50:883-895. [1, co-first authors; 2, co-communicating authors]

131. Li, H1, Bishop, KM1, O'Leary, DDM 2006 Potential target genes of EMX2 include Odz/Ten-M and other gene families with implications for cortical patterning. Mol Cell Neurosci, 33:136-149. [1, co-first authors]

132. Leingartner, A1, Thuret, S1, Kroll, TT, Chou, S-J, Leasure, JL, Gage, FH, O'Leary, DDM 2007 Sizes of somatosensory and motor cortical areas determine proficiency at tactile and motor behaviors. Proc. Natl. Acad. Sci. USA, 104:4153-4158. [1, co-first authors]

133. Takeuchi, A1, Hamasaki, T1, Litwack, ED, O'Leary DDM 2007 Novel IgCAM, MDGA1, expressed in unique cortical area- and layer-specific patterns and transiently by distinct forebrain populations of Cajal-Retzius neurons. Cerebral Cortex, 17:1531-1541. [1, co-first authors]

134. Sahara, S, Kawakami, Y, Izpisua Belmonte, JC, O'Leary, DDM 2007 Sp8 exhibits reciprocal induction with Fgf8 but has an opposing effect on anterior-posterior cortical area patterning. Neural Development, 2:10 doi:10.1186/1749-8104-2-10. PMCID: PMC1890288.

135. Burden, S, O'Leary, DDM, Scheiffele, P. 2007 Chapter 18: Target Selection, Topographic Maps, and Synapse Formation. pp. 469-498. In: Fundamentals of Neuroscience, 3rd Edition. (L. Squire, ed.). Academic Press.

136. Armentano, M1, Chou, S-J1, Tomassy, GS, Leingärtner, A, O'Leary, DDM2, Studer, M22007 COUP-TFI regulates the balance of cortical patterning between frontal/motor and sensory areas. Nature Neuroscience 10:1277-1286. [1, co-first authors; 2, co-communicating authors]

137. O'Leary, DDM, Chou, S-J, Hamasaki, T, Sahara, S, Takeuchi, A, Thuret, S, Leingartner, A 2007 Regulation of laminar and area patterning of mammalian neocortex and behavioural Implications. In: Development of the Cerebral Cortex. Novartis Foundation Symposium. G. Bock, J. Goode, Eds. Oxford Press. pp.141-164, PMID: 18494257.

138. O'Leary, DDM, Chou, S-J, Sahara, S 2007 Area patterning of the mammalian cortex. Neuron, 56:252-269, PMID: 17964244.

139. Zhou, L, Bar, I, Achouri, Y, Campbell, K, De Backer, O, Hebert, J, Jones, K, Kessaris, N, Lambert de Rouvroit, C, O'Leary, DDM, Richardson, W, Goffinet, A, Tissir, F 2008 Early Forebrain Wiring: Genetic Dissection Using Conditional Celsr3 Mutant Mice. Science 320: 946 – 949, PMCID: PMC2746700.

140. O'Leary, DDM, Sahara, S. 2008 Genetic regulation of arealization of the neocortex. Current Opinion in Neurobiology, 18:90-100, PMCID: PMC2677555.

141. Lim, Y-S1, McLaughlin, T1, Sung, T-C1, Santiago, A, Lee, K-F, O'Leary, DDM 2008 p75NTR mediates ephrin-A reverse signaling required for axon repulsion and mapping. Neuron, 59:746-758. [1, co-first authors], PMCID: PMC2677386.

142. McLaughlin, T, O'Leary, DDM 2009 Chemoaffinity Hypothesis: Development of Topographic Axonal Projections. In: Squire LR (ed.) Encyclopedia of Neuroscience, vol 2, pp. 819-825. Oxford: Academic Press.

143. O'Leary, DDM, Kroll, TK, 2009 Neural Patterning: Arealization of the Cortex. In: Squire LR (ed.) Encyclopedia of Neuroscience, vol 2, pp. 189-197. Oxford: Academic Press.

144. Nikolaev, A, McLaughlin, T, O'Leary, DDM, Tessier-Lavinge, M 2009 APP binds DR6 to trigger axon pruning and neuron death via distinct caspases. Nature 457:981-990. PMCID: PMC2677572.

145. Braisted, JE1, Ringstedt, T1, O'Leary, DDM 2009 Slits are chemorepellents endogenous to hypothalamus and steer thalamocortical axons into ventral telencephalon. Cerebral Cortex 19:i144-i155. [1, co-first authors]. PMCID: PMC2693534.

146. Sahara, S, O'Leary, DDM 2009 Fgf10 regulates transition period of cortical stem cell differentiation to radial glia controlling generation of neurons and basal progenitors. Neuron 63: 48-62. PMCID: PMC2746711.

147. Chou, S-J, Perez-Garcia, CG, Kroll, TT, O'Leary, DDM 2009 Lhx2 specifies regional fate in Emx1 lineage of telencephalic progenitors generating cerebral cortex. Nature Neurosci. 12:1381-1389. PMCID: PMC2897740.

148. Feldheim, DA, O'Leary, DDM 2010 Visual Map Development: Bidirectional Signaling, Bifunctional Guidance Molecules, and Competition. Cold Spring Harbor Perspectives in Biology. Web Pub Date 09-29-10. PMCID: PMC2964178.

149. Wu, CS, Zhu, J, Wager-Miller, J, Wang, S, O'Leary, DDM, Monory, K, Lutz, B, Mackie, K, Lu, HC 2010 Requirement of cannabinoid CB1 receptor in cortical pyramidal neurons for appropriate development of corticothalamic and thalamocortical projections. Eur. J. of Neuroscience 32:693-706. PMID: 21050275.

150. Dwyer, ND, Manning, DK, Moran, JL, Mudbhary, R, Fleming, MS, O'Leary, DDM, Walsh, CA, Beier, DR. 2011 A forward genetic screen with a thalamocortical axon reporter mouse yields novel neurodevelopment mutants and a distinct Emx2 mutant phenotype. Neural Development, 6:3. PMCID: PMC3024922.

151. Kawakami, Y, Marti, M, Kawakami, H, Itoh, J, Quach, T, Johnson, A, Sahara, S, O'Leary, DDM, Nakagawa, Y, Lewandoski, M, Pfaff, S, Evans, S, Izpisua Belmonte, JC 2011 Islet1-mediated activation of the ß-catenin pathway is necessary for hindlimb initiation in mice. Development 138:4465-4473. PMCID: PMC3177316.

152. Hurtado de Mendoza, T, Perez-Garcia, CG, Kroll T, Hoong, N, O'Leary, DDM, Verma, I. 2011 Antiapoptotic protein Lifegard is required for survival and maintenance of Purkinje and Granular Cells. PNAS 108:17189-17194. PMCID: PMC3193209.

153. Simon, DJ, Weimer, RM, McLaughlin, T, Kallop, D, Stanger, K, Yang, J, O'Leary, DDM, Hannoush, RN, Tessier-Lavigne, M. 2011 The caspase cascade of axon degeneration. Submitted.

154. Chou, S-J, O'Leary, DDM 2011 Role for Lhx2 in corticogenesis through regulation of progenitor differentiation. Submitted.

155. Pao, G, Zhu, Q, Perez-Garcia, C, Chou, S-J, Suh, H, Gage, FH, O'Leary, DDM, Verma, IM 2011 A role of tumor suppressor BRCA1 in embryonic brain development and neurogenesis Submitted.

156. Li H, Chou SJ, Hamasaki T, Perez-Garcia CG, O'Leary DD. Neuregulin repellent signaling via ErbB4 restricts GABAergic interneurons to migratory paths from ganglionic eminence to cortical destinations. Neural Dev 2012 Feb 29;7(1):10. PMCID: PMC3353847.

157. Sahara S, Yanagawa Y, O'Leary DD, Stevens CF. The fraction of cortical GABAergic neurons is constant from near the start of cortical neurogenesis to adulthood. J Neurosci 2012 Apr 4;32(14):4755-61. PMCID: PMC3325497.

158. Simon, DJ, Weimer, RM, McLaughlin, T, Kallop, D, Stanger, K, Yang, J, O'Leary, DDM, Hannoush, RN, Tessier-Lavigne, M. 2012 A Caspase Cascade Regulating Developmental Axon Degeneration. Journal of Neuroscience, in press.

159. O'Leary, DDM, Stocker, AM, Zembrzycki, A 2012 Area Patterning of the Mammalian Cortex. Patterning and Cell Type Specification in the Developing CNS and PNS: Comprehensive Developmental Neuroscience, Volume 1, Chapter 21, 25 pages. [].