Carla Shatz

Administrative Appointments

• Director, Bio-X , (2007– present )

Honors and Awards

• Elected Fellow, American Academy of Arts and Sciences (1992)
• Elected Member, European Academy of Sciences and Arts (1992)
• President, Society for Neuroscience (1994)
• Elected Member, National Academy of Sciences (1995)
• Charles A. Dana Award for Pioneering Achievement in Health and Education, Charles A. Dana Foundation (1995)
• Alcon Award for Outstanding Contributions to Vision Research, Alcon Research Institutre (1997)
• Elected Member, American Philosophical Society (1997)
• Elected Member, Institute of Medicine, National Academy of Sciences (1999)
• 2000 Weizmann Women and Science Award, Weizmann Institute (2000)
• Honorary Degree, Federal Institute of Technology, Lausanne, Switzerland (2002)
• Gill Prize in Neuroscience, Indiana University (2006)

Professional Education

• Postdoctoral, Harvard Medical School Neurobiology (1978)
• Ph.D., Harvard University Neurobiology (1976)
• M.Phil, University College London Physiology (1971)
• B.A., Radcliffe College, Cambridge, MA Chemistry (1969)

Postdoctoral Advisees

• Maja Djurisic, Han Mi Lee, Claus Thorsten Naserke

Graduate & Fellowship Program Affiliations

• Neurobiology
• Neurosciences

Web Site Links

• Shatz Lab Site

Research Interests

By studying the visual system of mammals, the Shatz Lab discovered that adult wiring emerges from dynamic interactions between neurons involving neural function and synaptic plasticity. Even before birth and long before vision, the eye spontaneously generates and sends coordinated patterns of neural activity to the brain. Blocking this activity in utero, or preventing vision after birth, disrupts normal tuning up of circuits and brain wiring. In turn, neural activity regulates the expression of genes involved in the process of circuit tuning. To discover cell and molecular underpinnings of circuit tuning, her lab has conducted functional screens for genes regulated by neural activity. Among these genes is the MHC (major histocompatibility) Class I family. This finding was very surprising because these genes- HLA genes in humans- are involved in cellular immunity and were previously not thought to be expressed by neurons at all! The Shatz Lab showed that other components of a signaling system for Class I MHC are also present in neurons, including a novel receptor, PirB. By studying and/or generating knockout mice, the lab is exploring a role for these molecules in synaptic plasticity, learning, memory and neurological disorders. The lab employs a variety of approaches in these studies, ranging from molecular biology to slice electrophysiology to in vivo imaging to behavior. Research has relevance not only for understanding brain wiring and developmental disorders such as Autism and Schizphrenia, but also for understanding how the nervous and immune systems interact.

Publications

• Synaptogenesis in purified cortical subplate neurons. McKellar CE,  Shatz CJ.  Cereb Cortex.  2009; 19 (8): 1723-37
• MHC class I: an unexpected role in neuronal plasticity. Shatz CJ,  Neuron.  2009; 64 (1): 40-5
• Co-regulation of ocular dominance plasticity and NMDA receptor subunit expression in glutamic acid decarboxylase-65 knock-out mice. Kanold PO,  Kim YA, GrandPre T, Shatz CJ.  J Physiol.  2009; 587 (Pt 12): 2857-67
• H2-K(b) and H2-D(b) regulate cerebellar long-term depression and limit motor learning. McConnell MJ,  Huang YH, Datwani A, Shatz CJ.  Proc Natl Acad Sci U S A.  2009; 106 (16): 6784-9
• Regulation of CNS synapses by neuronal MHC class I. Goddard CA,  Butts DA, Shatz CJ.  Proc Natl Acad Sci U S A.  2007; 104 (16): 6828-33
• A burst-based "Hebbian" learning rule at retinogeniculate synapses links retinal waves to activity-dependent refinement. Butts DA,  Kanold PO, Shatz CJ.  PLoS Biol.  2007; 5 (3): e61
• Subplate neurons regulate maturation of cortical inhibition and outcome of ocular dominance plasticity. Kanold PO,  Shatz CJ.  Neuron.  2006; 51 (5): 627-38
• PirB restricts ocular-dominance plasticity in visual cortex. Syken J,  Grandpre T, Kanold PO, Shatz CJ.  Science.  2006; 313 (5794): 1795-800
• Effects of visual experience on activity-dependent gene regulation in cortex. Majdan M,  Shatz CJ.  Nat Neurosci.  2006; 9 (5): 650-9
• Lawrence C. Katz (1956-2005). Shatz CJ,  Nature.  2006; 439 (7073): 152
• Multiple periods of functional ocular dominance plasticity in mouse visual cortex. Tagawa Y,  Kanold PO, Majdan M, Shatz CJ.  Nat Neurosci.  2005; 8 (3): 380-8
• Changing scientific publishing. Raff MC,  Stevens CF, Roberts K, Shatz CJ, Roberts K, Shatz CJ, Newsome WT.  Science.  2004; 305 (5686): 945-6
• Immune signalling in neural development, synaptic plasticity and disease. Boulanger LM,  Shatz CJ.  Nat Rev Neurosci.  2004; 5 (7): 521-31
• Expression of T cell receptor beta locus in central nervous system neurons. Syken J,  Shatz CJ.  Proc Natl Acad Sci U S A.  2003; 100 (22): 13048-53
• Role of subplate neurons in functional maturation of visual cortical columns. Kanold PO,  Kara P, Reid RC, Shatz CJ.  Science.  2003; 301 (5632): 521-5
• Selective vulnerability of subplate neurons after early neonatal hypoxia-ischemia. McQuillen PS,  Sheldon RA, Shatz CJ, Ferriero DM.  J Neurosci.  2003; 23 (8): 3308-15
• A novel role for p75NTR in subplate growth cone complexity and visual thalamocortical innervation. McQuillen PS,  DeFreitas MF, Zada G, Shatz CJ.  J Neurosci.  2002; 22 (9): 3580-93
• An instructive role for retinal waves in the development of retinogeniculate connectivity. Stellwagen D,  Shatz CJ.  Neuron.  2002; 33 (3): 357-67
• Neuronal plasticity and cellular immunity: shared molecular mechanisms. Boulanger LM,  Huh GS, Shatz CJ.  Curr Opin Neurobiol.  2001; 11 (5): 568-78
• A novel p75NTR signaling pathway promotes survival, not death, of immunopurified neocortical subplate neurons. DeFreitas MF,  McQuillen PS, Shatz CJ.  J Neurosci.  2001; 21 (14): 5121-9
• Functional requirement for class I MHC in CNS development and plasticity. Huh GS,  Boulanger LM, Du H, Riquelme PA, Brotz TM, Shatz CJ.  Science.  2000; 290 (5499): 2155-9
• Netrin-1 promotes thalamic axon growth and is required for proper development of the thalamocortical projection. Braisted JE,  Catalano SM, Stimac R, Kennedy TE, Tessier-Lavigne M, Shatz CJ, O'Leary DD.  J Neurosci.  2000; 20 (15): 5792-801
• Dynamic regulation of BDNF and NT-3 expression during visual system development. Lein ES,  Hohn A, Shatz CJ.  J Comp Neurol.  2000; 420 (1): 1-18
• Rapid regulation of brain-derived neurotrophic factor mRNA within eye-specific circuits during ocular dominance column formation. Lein ES,  Shatz CJ.  J Neurosci.  2000; 20 (4): 1470-83
• Subplate neuron ablation alters neurotrophin expression and ocular dominance column formation. Lein ES,  Finney EM, McQuillen PS, Shatz CJ.  Proc Natl Acad Sci U S A.  1999; 96 (23): 13491-5
• Dynamics of retinal waves are controlled by cyclic AMP. Stellwagen D,  Shatz CJ, Feller MB.  Neuron.  1999; 24 (3): 673-85
• Dynamic regulation of cpg15 during activity-dependent synaptic development in the mammalian visual system. Corriveau RA,  Shatz CJ, Nedivi E.  J Neurosci.  1999; 19 (18): 7999-8008
• Retinal waves are governed by collective network properties. Butts DA,  Feller MB, Shatz CJ, Rokhsar DS.  J Neurosci.  1999; 19 (9): 3580-93
• Brain waves and brain wiring: the role of endogenous and sensory-driven neural activity in development. Penn AA,  Shatz CJ.  Pediatr Res.  1999; 45 (4 Pt 1): 447-58
• Establishment of patterned thalamocortical connections does not require nitric oxide synthase. Finney EM,  Shatz CJ.  J Neurosci.  1998; 18 (21): 8826-38
• Regulation of class I MHC gene expression in the developing and mature CNS by neural activity. Corriveau RA,  Huh GS, Shatz CJ.  Neuron.  1998; 21 (3): 505-20
• Major glutamatergic projection from subplate into visual cortex during development. Finney EM,  Stone JR, Shatz CJ.  J Comp Neurol.  1998; 398 (1): 105-18
• Activity-dependent cortical target selection by thalamic axons. Catalano SM,  Shatz CJ.  Science.  1998; 281 (5376): 559-62
• Many major CNS axon projections develop normally in the absence of semaphorin III. Catalano SM,  Messersmith EK, Goodman CS, Shatz CJ, Chédotal A.  Mol Cell Neurosci.  1998; 11 (4): 173-82
• Competition in retinogeniculate patterning driven by spontaneous activity. Penn AA,  Riquelme PA, Feller MB, Shatz CJ.  Science.  1998; 279 (5359): 2108-12
• Form from function in visual system development. Shatz CJ,  Harvey Lect.  1997-1998; 93 17-34
• Dendritic development of retinal ganglion cells after prenatal intracranial infusion of tetrodotoxin. Campbell G,  Ramoa AS, Stryker MP, Shatz CJ.  Vis Neurosci.  1997 Jul-Aug; 14 (4): 779-88
• Dynamic processes shape spatiotemporal properties of retinal waves. Feller MB,  Butts DA, Aaron HL, Rokhsar DS, Shatz CJ.  Neuron.  1997; 19 (2): 293-306
• Blockade of endogenous ligands of trkB inhibits formation of ocular dominance columns. Cabelli RJ,  Shelton DL, Segal RA, Shatz CJ.  Neuron.  1997; 19 (1): 63-76
• Activity-dependent regulation of NMDAR1 immunoreactivity in the developing visual cortex. Catalano SM,  Chang CK, Shatz CJ.  J Neurosci.  1997; 17 (21): 8376-90
• Migration of neocortical neurons in the absence of functional NMDA receptors. Messersmith EK,  Feller MB, Zhang H, Shatz CJ.  Mol Cell Neurosci.  1997; 9 (5-6): 347-57
• Developmental changes revealed by immunohistochemical markers in human cerebral cortex. Honig LS,  Herrmann K, Shatz CJ.  Cereb Cortex.  1996 Nov-Dec; 6 (6): 794-806
• Synaptic activity and the construction of cortical circuits. Katz LC,  Shatz CJ.  Science.  1996; 274 (5290): 1133-8
• Thalamic relay of spontaneous retinal activity prior to vision. Mooney R,  Penn AA, Gallego R, Shatz CJ.  Neuron.  1996; 17 (5): 863-74
• Emergence of order in visual system development. Shatz CJ,  Proc Natl Acad Sci U S A.  1996; 93 (2): 602-8
• Emergence of order in visual system development. Shatz CJ,  J Physiol Paris.  1996; 90 (3-4): 141-50
• Changing patterns of expression and subcellular localization of TrkB in the developing visual system. Cabelli RJ,  Allendoerfer KL, Radeke MJ, Welcher AA, Feinstein SC, Shatz CJ.  J Neurosci.  1996; 16 (24): 7965-80
• Requirement for cholinergic synaptic transmission in the propagation of spontaneous retinal waves. Feller MB,  Wellis DP, Stellwagen D, Werblin FS, Shatz CJ.  Science.  1996; 272 (5265): 1182-7
• Early functional neural networks in the developing retina. Wong RO,  Chernjavsky A, Smith SJ, Shatz CJ.  Nature.  1995; 374 (6524): 716-8
• Semaphorin III can function as a selective chemorepellent to pattern sensory projections in the spinal cord. Messersmith EK,  Leonardo ED, Shatz CJ, Tessier-Lavigne M, Goodman CS, Kolodkin AL.  Neuron.  1995; 14 (5): 949-59
• Blockade of action potential activity alters initial arborization of thalamic axons within cortical layer 4. Herrmann K,  Shatz CJ.  Proc Natl Acad Sci U S A.  1995; 92 (24): 11244-8
• Inhibition of ocular dominance column formation by infusion of NT-4/5 or BDNF. Cabelli RJ,  Hohn A, Shatz CJ.  Science.  1995; 267 (5204): 1662-6
• Viktor Hamburger Award review. Role for spontaneous neural activity in the patterning of connections between retina and LGN during visual system development. Shatz CJ,  Int J Dev Neurosci.  1994; 12 (6): 531-46
• Ultrastructural evidence for synaptic interactions between thalamocortical axons and subplate neurons. Herrmann K,  Antonini A, Shatz CJ.  Eur J Neurosci.  1994; 6 (11): 1729-42
• Segregation of geniculocortical afferents during the critical period: a role for subplate neurons. Ghosh A,  Shatz CJ.  J Neurosci.  1994; 14 (6): 3862-80
• The subplate, a transient neocortical structure: its role in the development of connections between thalamus and cortex. Allendoerfer KL,  Shatz CJ.  Annu Rev Neurosci.  1994; 17 185-218
• Regulation of neurotrophin receptors during the maturation of the mammalian visual system. Allendoerfer KL,  Cabelli RJ, Escandón E, Kaplan DR, Nikolics K, Shatz CJ.  J Neurosci.  1994; 14 (3 Pt 2): 1795-811
• Subplate pioneers and the formation of descending connections from cerebral cortex. McConnell SK,  Ghosh A, Shatz CJ.  J Neurosci.  1994; 14 (4): 1892-907
• Independent control of dendritic and axonal form in the developing lateral geniculate nucleus. Dalva MB,  Ghosh A, Shatz CJ.  J Neurosci.  1994; 14 (6): 3588-602
• Neuronal coupling in the developing mammalian retina. Penn AA,  Wong RO, Shatz CJ.  J Neurosci.  1994; 14 (6): 3805-15
• Enhancement of transmission at the developing retinogeniculate synapse. Mooney R,  Madison DV, Shatz CJ.  Neuron.  1993; 10 (5): 815-25
• Transient period of correlated bursting activity during development of the mammalian retina. Wong RO,  Meister M, Shatz CJ.  Neuron.  1993; 11 (5): 923-38
• Developmental mechanisms that generate precise patterns of neuronal connectivity. Goodman CS,  Shatz CJ.  Cell.  1993; 72 Suppl 77-98
• A role for subplate neurons in the patterning of connections from thalamus to neocortex. Ghosh A,  Shatz CJ.  Development.  1993; 117 (3): 1031-47
• Repair and replacement to restore sight. Report from the Panel on Ganglion Cell/Connectivity. Shatz CJ,  O'Leary DD.  Arch Ophthalmol.  1993; 111 (4): 472-7
• How are specific connections formed between thalamus and cortex? Shatz CJ,  Curr Opin Neurobiol.  1992; 2 (1): 78-82
• Synaptic Contacts and the Transient Dendritic Spines of Developing Retinal Ganglion Cells. Wong RO,  Yamawaki RM, Shatz CJ.  Eur J Neurosci.  1992; 4 (12): 1387-1397
• Involvement of subplate neurons in the formation of ocular dominance columns. Ghosh A,  Shatz CJ.  Science.  1992; 255 (5050): 1441-3
• Synapses formed by identified retinogeniculate axons during the segregation of eye input. Campbell G,  Shatz CJ.  J Neurosci.  1992; 12 (5): 1847-58
• The developing brain. Shatz CJ,  Sci Am.  1992; 267 (3): 60-7
• Dividing up the neocortex. Shatz CJ,  Science.  1992; 258 (5080): 237-8
• Neuronal death, a tradition of dying. Oppenheim RW,  Schwartz LM, Shatz CJ.  J Neurobiol.  1992; 23 (9): 1111-5
• Pathfinding and target selection by developing geniculocortical axons. Ghosh A,  Shatz CJ.  J Neurosci.  1992; 12 (1): 39-55
• Morphology of pioneer and follower growth cones in the developing cerebral cortex. Kim GJ,  Shatz CJ, McConnell SK.  J Neurobiol.  1991; 22 (6): 629-42
• Remodeling of retinal ganglion cell dendrites in the absence of action potential activity. Wong RO,  Herrmann K, Shatz CJ.  J Neurobiol.  1991; 22 (7): 685-97
• Changing patterns of synaptic input to subplate and cortical plate during development of visual cortex. Friauf E,  Shatz CJ.  J Neurophysiol.  1991; 66 (6): 2059-71
• Synchronous bursts of action potentials in ganglion cells of the developing mammalian retina. Meister M,  Wong RO, Baylor DA, Shatz CJ.  Science.  1991; 252 (5008): 939-43
• The Effects of Prenatal Intracranial Infusion of Tetrodotoxin on Naturally Occurring Retinal Ganglion Cell Death and Optic Nerve Ultrastructure. Friedman S,  Shatz CJ.  Eur J Neurosci.  1990; 2 (3): 243-253
• Relation Between Putative Transmitter Phenotypes and Connectivity of Subplate Neurons During Cerebral Cortical Development. Antonini A,  Shatz CJ.  Eur J Neurosci.  1990; 2 (9): 744-761
• Functional synaptic circuits in the subplate during fetal and early postnatal development of cat visual cortex. Friauf E,  McConnell SK, Shatz CJ.  J Neurosci.  1990; 10 (8): 2601-13
• Competitive interactions between retinal ganglion cells during prenatal development. Shatz CJ,  J Neurobiol.  1990; 21 (1): 197-211
• Nerve growth factor receptor immunoreactivity is transiently associated with the subplate neurons of the mammalian cerebral cortex. Allendoerfer KL,  Shelton DL, Shooter EM, Shatz CJ.  Proc Natl Acad Sci U S A.  1990; 87 (1): 187-90
• Pioneer neurons and target selection in cerebral cortical development. Shatz CJ,  Ghosh A, McConnell SK, Allendoerfer KL, Friauf E, Antonini A.  Cold Spring Harb Symp Quant Biol.  1990; 55 469-80
• Impulse activity and the patterning of connections during CNS development. Shatz CJ,  Neuron.  1990; 5 (6): 745-56
• Requirement for subplate neurons in the formation of thalamocortical connections. Ghosh A,  Antonini A, McConnell SK, Shatz CJ.  Nature.  1990; 347 (6289): 179-81
• Retinal ganglion beta cells project transiently to the superior colliculus during development. Ramoa AS,  Campbell G, Shatz CJ.  Proc Natl Acad Sci U S A.  1989; 86 (6): 2061-5
• Subplate neurons pioneer the first axon pathway from the cerebral cortex. McConnell SK,  Ghosh A, Shatz CJ.  Science.  1989; 245 (4921): 978-82
• The earliest-generated neurons of the cat cerebral cortex: characterization by MAP2 and neurotransmitter immunohistochemistry during fetal life. Chun JJ,  Shatz CJ.  J Neurosci.  1989; 9 (5): 1648-67
• Interstitial cells of the adult neocortical white matter are the remnant of the early generated subplate neuron population. Chun JJ,  Shatz CJ.  J Comp Neurol.  1989; 282 (4): 555-69
• Prenatal tetrodotoxin infusion blocks segregation of retinogeniculate afferents. Shatz CJ,  Stryker MP.  Science.  1988; 242 (4875): 87-9
• Dendritic growth and remodeling of cat retinal ganglion cells during fetal and postnatal development. Ramoa AS,  Campbell G, Shatz CJ.  J Neurosci.  1988; 8 (11): 4239-61
• Modification of retinal ganglion cell axon morphology by prenatal infusion of tetrodotoxin. Sretavan DW,  Shatz CJ, Stryker MP.  Nature.  1988; 336 (6198): 468-71
• Axon arbors of X and Y retinal ganglion cells are differentially affected by prenatal disruption of binocular inputs. Garraghty PE,  Shatz CJ, Sretavan DW, Sur M.  Proc Natl Acad Sci U S A.  1988; 85 (19): 7361-5
• Redistribution of synaptic vesicle antigens is correlated with the disappearance of a transient synaptic zone in the developing cerebral cortex. Chun JJ,  Shatz CJ.  Neuron.  1988; 1 (4): 297-310
• Abnormal pigmentation and unusual morphogenesis of the optic stalk may be correlated with retinal axon misguidance in embryonic Siamese cats. Webster MJ,  Shatz CJ, Kliot M, Silver J.  J Comp Neurol.  1988; 269 (4): 592-611
• Prenatal disruption of binocular interactions creates novel lamination in the cat's lateral geniculate nucleus. Garraghty PE,  Shatz CJ, Sur M.  Vis Neurosci.  1988; 1 (1): 93-102
• A fibronectin-like molecule is present in the developing cat cerebral cortex and is correlated with subplate neurons. Chun JJ,  Shatz CJ.  J Cell Biol.  1988; 106 (3): 857-72
• Transient cells of the developing mammalian telencephalon are peptide-immunoreactive neurons. Chun JJ,  Nakamura MJ, Shatz CJ.  Nature.  1987 Feb 12-18; 325 (6105): 617-20
• Transient morphological features of identified ganglion cells in living fetal and neonatal retina. Ramoa AS,  Campbell G, Shatz CJ.  Science.  1987; 237 (4814): 522-5
• Axon trajectories and pattern of terminal arborization during the prenatal development of the cat's retinogeniculate pathway. Sretavan DW,  Shatz CJ.  J Comp Neurol.  1987; 255 (3): 386-400
• Development of the mammalian visual system. Shatz CJ,  Mead Johnson Symp Perinat Dev Med.  1987; (29): 19-26
• Prenatal development of cat retinogeniculate axon arbors in the absence of binocular interactions. Sretavan DW,  Shatz CJ.  J Neurosci.  1986; 6 (4): 990-1003
• Development of glutamic acid decarboxylase immunoreactivity in the cat's lateral geniculate nucleus. Shotwell SL,  Shatz CJ, Luskin MB.  J Neurosci.  1986; 6 (5): 1410-23
• The relationship between the geniculocortical afferents and their cortical target cells during development of the cat's primary visual cortex. Shatz CJ,  Luskin MB.  J Neurosci.  1986; 6 (12): 3655-68
• Prenatal development of retinal ganglion cell axons: segregation into eye-specific layers within the cat's lateral geniculate nucleus. Sretavan DW,  Shatz CJ.  J Neurosci.  1986; 6 (1): 234-51
• Interactions between retinal ganglion cells during the development of the mammalian visual system. Shatz CJ,  Sretavan DW.  Annu Rev Neurosci.  1986; 9 171-207
• Neurogenesis of the cat's primary visual cortex. Luskin MB,  Shatz CJ.  J Comp Neurol.  1985; 242 (4): 611-31
• Abnormal development of the retinogeniculate projection in Siamese cats. Kliot M,  Shatz CJ.  J Neurosci.  1985; 5 (10): 2641-53
• Visual Neurobiology: Development of Visual Pathways in Mammals. Shatz CJ,  Science.  1985; 228 (4695): 67-68
• Studies of the earliest generated cells of the cat's visual cortex: cogeneration of subplate and marginal zones. Luskin MB,  Shatz CJ.  J Neurosci.  1985; 5 (4): 1062-75
• Prenatal development of individual retinogeniculate axons during the period of segregation. Sretavan D,  Shatz CJ.  Nature.  1984 Apr 26-May 2; 308 (5962): 845-8
• Prenatal development of functional connections in the cat's retinogeniculate pathway. Shatz CJ,  Kirkwood PA.  J Neurosci.  1984; 4 (5): 1378-97
• The prenatal development of the cat's retinogeniculate pathway. Shatz CJ,  J Neurosci.  1983; 3 (3): 482-99
• Prenatal misrouting of the retinogeniculate pathway in Siamese cats. Shatz CJ,  Kliot M.  Nature.  1982; 300 (5892): 525-9
• The genesis of efferent connections from the visual cortex of the fetal rhesus monkey. Shatz CJ,  Rakic P.  J Comp Neurol.  1981; 196 (2): 287-307
• Siamese cat: altered connections of visual cortex. Shatz CJ,  LeVay S.  Science.  1979; 204 (4390): 328-30
• Ocular dominance in layer IV of the cat's visual cortex and the effects of monocular deprivation. Shatz CJ,  Stryker MP.  J Physiol.  1978; 281 267-83
• Ocular dominance columns and their development in layer IV of the cat's visual cortex: a quantitative study. LeVay S,  Stryker MP, Shatz CJ.  J Comp Neurol.  1978; 179 (1): 223-44
• The distribution of afferents representing the right and left eyes in the cat's visual cortex. Shatz CJ,  Lindström S, Wiesel TN.  Brain Res.  1977; 131 (1): 103-16
• Anatomy of interhemispheric connections in the visual system of Boston Siamese and ordinary cats. Shatz CJ,  J Comp Neurol.  1977; 173 (3): 497-518