The primary somatosensory cortex of the platypus (Ornithorhynchus anatinus) is characterized by a distinct array of functionally specific cytochrome oxidase (CO) modules, forming alternate CO-rich and CO-poor bands. In the current study, we undertook to establish whether the cellular morphology of layer V pyramidal neurones reflects this modular organization. To this end, we injected neurones with Lucifer Yellow in 250 μm thick, flat-mounted cortical slices and processed the tissue to reveal a light-stable reaction product. By aligning blood vessels in serial sections in which we injected individual neurones with sections processed for CO, we were able to establish the exact location of injected cells with respect to the pattern of CO bands. Pyramidal neurones in the CO-poor bands (which are responsive to both mechano- and electroreceptive stimuli) had basal dendritic fields that were larger than those in the CO-rich bands. The large basal dendritic fields of layer V pyramidal neurones in the CO-poor bands may allow for integration of a greater number of more diverse inputs, thus allowing for averaging of stimuli to improve the signal-to-noise ratio or enhance spatial discrimination of peripheral stimuli. In some instances, neurones located within approximately 100 μm of the boundaries of the CO bands had dendritic fields that appeared to conform to the CO bands, the dendrites preferentially arborizing within a single band and avoiding the neighbouring band. However, the bias was not absolute, as we observed many examples of cells with dendrites that crossed the boundary between bands. Furthermore, many cells had dendrites that showed distinct dendritic bias that bore no obvious relationship to the CO boundaries.

1.
Abbie, A.A. (1940) Cortical lamination in the monotremata. J. Comp. Neurol., 72: 429–467.
2.
Agmon, A., L.T. Tang, E.G. Jones, and D.K. O’Dowd (1995) Topological precision in the thalamic projection to neonatal mouse barrel cortex. J. Neurosci., 15: 549–561.
3.
Archer, M., F.A. Jenkins, S.J. Hand, P. Murray, and H. Godthelp (1992) Description of the skull and non-vestigial dentition of a Miocene platypus (Obdurodon dicksoni n. sp.) from Riversleigh, Australia, and the problem of monotreme origins. In Platypus and Echidnas, Royal Zoological Society of New South Wales, Australia (ed. by M.L. Augee), Beaver Press, Australia, pp. 15–27.
4.
Belford, G.R., and H.P. Killackey (1979) Vibrissae representation in subcortical trigeminal centers of the neonatal rat. J. Comp. Neurol., 183: 305–322.
5.
Buhl, E.H., and W. Schlote (1987) Intracellular Lucifer yellow staining and electronmicroscopy of neurones in slices of fixed epitumourous human cortical tissue. Acta Neuropathol, 75: 140–146.
6.
Casagrande, V.A., and J.H. Kaas (1994) The afferent, intrinsic and efferent connections of primary visual cortex in primates. In Cerebral Cortex, Vol. 10: Primary Visual Cortex in Primates (ed. by A. Peters and K.S. Rockland), Plenum, New York, pp. 201–259.
7.
Catania, K.C., R.G. Northcutt, J.H. Kaas, and P.D. Beck (1993) Nose stars and brain stripes. Nature, 364: 493.
8.
Colonnier, M. (1968) Synaptic patterns on different cell types in the different laminae of the cat visual cortex. Brain Res., 9: 268–287.
9.
DeFelipe, J., and I. Fariñas (1992) The pyramidal neuron of the cerebral cortex: morphological and chemical characteristics of the synaptic inputs. Prog. Neurobiol., 39: 563–607.
10.
DeFelipe, J., and E.G. Jones (1988) Cajal on the Cerebral Cortex. Oxford University Press, Oxford.
11.
Einstein, G. (1988) Intracellular injection of Lucifer Yellow into cortical neurons in lightly fixed sections and its application to human autopsy material. J. Neurosci. Methods, 26: 95–103.
12.
Elston, G.N., and M.G.P. Rosa (1997) The occipitoparietal pathway of the macaque monkey: comparison of pyramidal cell morphology in layer III of functionally related cortical visual areas. Cereb. Cortex, 7: 432–452.
13.
Elston, G.N., and M.G.P. Rosa (1998a) Morphological variation of layer III pyramidal neurones in the occipitotemporal pathway of the macaque monkey visual cortex. Cereb. Cortex, 8: 278– 294.
14.
Elston, G.N., and M.G.P. Rosa (1998b) Complex dendritic fields of pyramidal cells in the frontal eye field of the macaque monkey: comparison with parietal areas 7a and LIP. NeuroReport, 9: 127–131.
15.
Elston, G.N., and M.G.P. Rosa (1998c) Morphological variation of layer V pyramidal neurones in visual areas of the temporal lobe of the macaque monkey. Proc. Aust. Neurosci. Abstr., 9: 25.
16.
Elston, G.N., D.V. Pow, and M.B. Calford (1997) Neuronal composition and morphology in layer IV of two vibrissal barrel subfields of rat cortex. Cereb. Cortex, 7: 422–431.
17.
Elston, G.N., M.G.P. Rosa, and M.B. Calford (1996) Comparison of dendritic fields of layer III pyramidal neurones in striate and extrastriate visual areas of the marmoset: a lucifer yellow intracellular injection study. Cereb. Cortex, 6: 807– 813.
18.
Elston, G.N., M.G.P. Rosa, and M.B. Calford (1998) Reorganization of neuronal dendritic fields in adult monkey V1 following retinal lesions. Eur. Neurosci. Assn. Abstr., 10(suppl. 10): 227.
19.
Fitzpatrick, D., J.S. Lund, and G.G Blasdel (1985) Intrinsic connections of macaque striate cortex. Afferent and efferent connections of lamina 4C. J. Neurosci., 5: 3329–3349.
20.
Florence, S.L., and J.H. Kaas (1992) Ocular dominance columns in area 17 of Old World macaque and talapoin monkeys: complete reconstructions and quantitative analyses. Vis. Neurosci., 8: 449–462.
21.
Gilbert, C.D., and T.N. Wiesel (1979) Morphology and intracortical projections of functionally characterised neurones in the cat visual cortex. Nature, 280: 120–125.
22.
Gray, E.G. (1959) Axo-somatic and axo-dendritic synapses of the cerebral cortex; an electron microscope study. J. Anat., 93: 420–433.
23.
Greenough, W., and F. Chang (1988) Dendritic pattern formation involves both oriented regression and oriented growth in the barrels of mouse somatosensory cortex. Dev Brain Res., 43: 148–152.
24.
Harris, R.M., and T.A. Woolsey (1981) Dendritic plasticity in mouse barrel cortex following postnatal vibrissa follicle damage. J. Comp. Neurol., 196: 357–376.
25.
Hevner, R.F., and M.T.T. Wong-Riley (1992) Entorhinal cortex of the human, monkey and rat: metabolic map as revealed by cytochrome oxidase. J. Comp. Neurol., 326: 451–469.
26.
Horton, J.C., and D.R. Hocking (1996) Anatomical demonstration of ocular dominance columns in striate cortex of the squirrel monkey. J. Neurosci., 16: 5510–5522.
27.
Hubel, D.H., and T.N. Wiesel (1968) Receptive fields and functional architecture of monkey striate cortex. J. Physiol., 195: 215–243.
28.
Hubel, D.H., and T.N. Wiesel (1972) Laminar and columnar distribution of geniculocortical fibres in the macaque monkey. J. Comp. Neurol., 146: 421–450.
29.
Hübener, M., and J. Bolz (1988) Morphology of identified projection neurons in layer 5 of rat visual cortex. Neurosci. Lett., 94: 76–81.
30.
Hübener, M., and J. Bolz (1992) Relationship between dendritic morphology and cytochrome oxidase compartments in monkey striate cortex. J. Comp. Neurol., 324: 67–80.
31.
Ito, M. (1992) Simultaneous visualization of cortical barrels and horseradish peroxidase-injected layer 5b vibrissa neurons in the rat. J. Physiol., 454: 247–265.
32.
Jones, E.G. (1968) An electron microscopic study of the terminations of afferent fiber systems onto the somatic sensory cortex of the cat. J. Anat., 103: 595–597.
33.
Jones, E.G. (1975) Varieties and distribution of non-pyramidal cells in the somatic sensory cortex of the squirrel monkey. J. Comp. Neurol., 160: 205–268.
34.
Jones, E.G., and T.P.S. Powell (1969) Morphological variations in the dendritic spines of the neocortex. J. Cell Sci., 5: 509–529.
35.
Katz, L.C., and M. Constantine-Paton (1988) Relationship between segregated afferents and postsynaptic neurons in the optic tectum of three-eyed frogs. J. Neurosci., 8: 3160–3180.
36.
Katz, L.C., C.D. Gilbert, and T.N. Wiesel (1989) Local circuits and ocular dominance columns in monkey striate cortex. J. Neurosci., 9: 1389– 1399.
37.
Killackey, H. (1973) Anatomical evidence for cortical subdivisions based on vertically discrete thalamic projections from the ventral posterior nucleus to cortical barrels in the rat. Brain Res., 51: 326–331.
38.
Killackey, H.P., and S. Leshin (1975) The organization of specific thalamocortical projections to the posteromedial barrel subfield of the rat somatic sensory cortex. Brain Res., 86: 469– 472.
39.
Kim, C.B.Y., L.P. Pier, and P.D. Spear (1997) Effects of ageing on numbers and sizes of neurons in histochemically defined subregions of monkey striate cortex. Anat. Rec., 247: 119– 128.
40.
Kirsch, J.A., and G.C. Mayer (1998) The platypus is not a rodent: DNA hybridisation, amniote physiology, and the palimpsest theory. Phil. Trans. R. Soc. Lond. B., 353: 1221–1237.
41.
Kossel, A., S. Lowel, and J. Bolz (1995) Relationships between dendritic fields and functional architecture in striate cortex of normal and visually deprived cats. J. Neurosci., 15: 3913– 3926.
42.
Krubitzer, L.A., P. Manger, J.D. Pettigrew, and M. Calford (1995) Organization of somatosensory cortex in monotremes: in search of the prototypical plan. J. Comp. Neurol., 351: 261–306.
43.
Lachica, E.A., and V.A. Casagrande (1992) Direct W-like geniculate projections to the cytochrome oxidase (CO) blobs in primate visual cortex. J. Comp. Neurol., 319: 141–158.
44.
Lachica, E.A., P.D. Beck, and V.A. Casagrande (1993) Intrinsic connections of layer III of striate cortex in squirrel monkey and bush baby: correlation with patterns of cytochrome oxidase. J. Comp. Neurol., 329: 163–187.
45.
Larramendi, L.M.H., L. Fickenscher, and L. Lemkey-Johnston (1967) Synaptic vesicles of inhibitory and excitatory terminals in the cerebellum. Science, 156: 967–969.
46.
LeVay, S., M.P. Stryker, and C.J. Schatz (1978) Ocular dominance columns and their development in layer IV of the cat’s visual cortex: a quantitative study. J. Comp. Neurol., 179: 223–244.
47.
Livingstone, M.S. (1996) Ocular dominance columns in New World monkeys. J. Neurosci., 16: 2086–2096.
48.
Livingstone, M.S., and D.H. Hubel (1984) Anatomy and physiology of a color system in the primate visual cortex. J. Neurosci., 4: 309–356.
49.
Lorente De Nó, R. (1938) The cerebral cortex, architecture, intracortical connections and motor projections. In Physiology of the nervous system (ed. by J.F. Foulton), Oxford University Press, London, pp. 291–325.
50.
Lund, J.S. (1984) Spiny stellate neurons. In Cerebral Cortex, Vol. 1: Cellular components of the cerebral cortex (ed. by A. Peters and E.G. Jones), Plenum, New York, pp. 255–308.
51.
Lund, J.S., R.D. Lund, A.E. Hendrickson, A.H. Bunt, and A.F. Fuchs (1975) The origin of efferent pathways from the primary visual cortex, area 17, of the macaque monkey as shown by retrograde transport of horseradish peroxidase. J. Comp. Neurol., 164: 287–304.
52.
Lund, J., T. Yoshioka, and J.B. Levitt (1993) Comparison of intrinsic connectivity in different areas of macaque monkey cerebral cortex. Cerebral Cortex, 3: 148–162.
53.
Malach, R. (1992) Dendritic sampling across processing streams in monkey striate cortex. J. Comp. Neurol., 315: 303–312.
54.
Manger, P.R. (1994) Platypus electroreception: neuroethology of a novel mammalian sensory system. PhD thesis, The University of Queensland, Australia.
55.
Manger, P.R., and J.D. Pettigrew (1995) Electroreception and the feeding behaviour of the platypus. Phil. Trans. R. Soc. Lond. B., 347: 359–381.
56.
Manger, P.R., and J.D. Pettigrew (1996) Ultrastructure, number, distribution and innervation of electroreceptors and mechanoreceptors in the bill skin of the platypus. Brain Behav. Evol., 48: 27–54.
57.
Manger, P.R., M.B. Calford, and J.D. Pettigrew (1996) Properties of electrosensory neurons in the cortex of the platypus (Ornithorhynchus anatinus): implications for processing of electrosensory stimuli. Proc. R. Soc. Lond. B, 263: 611–617.
58.
Manger, P.R., J.D. Pettigrew, J. Keast, and A. Bauer (1995) Nerve terminals of mucous gland electroreceptors in the platypus. Proc. R. Soc. Lond. B, 260: 13–19.
59.
Manger, P.R., M. Sum, M. Szymanski, S. Ridgway, and L.A. Krubitzer (1998) Modular subdivisions of dolphin insular cortex: does evolutionary history repeat itself? J. Cog. Neurosci., 10: 153–166.
60.
Martinich, S., M.G.P. Rosa, and C.E. Rocha-Miranda (1990) Patterns of cytochrome oxidase activity in the visual cortex of a South American opossum (Didelphis marsupialis aurita). Braz. J. Med. Biol. Res., 23: 883–887.
61.
Matsubara, J.A., R. Chase, and M. Thejomayen (1996) Comparative morphology of three types of projection-identified pyramidal neurons in the superficial layers of cat visual cortex. J. Comp. Neurol., 366: 93–108.
62.
Murphy, K.M., D.G. Jones, and C. van Sluyters (1995) Cytochrome-oxidase blobs in cat primary visual cortex. J. Neurosci., 15: 4196– 4208.
63.
Musser, A.M., and M. Archer (1998) New information about the skull and dentary of the Miocene platypus Obdurodon dicksoni and a discussion of ornithorhynchid relationships. Phil. Trans. R. Soc. Lond. B., 353: 1059–1061.
64.
Nieuwenhuys, R. (1994) The Neocortex. An overview of its evolutionary development, structural organization and synaptology. Anat. Embryol., 190: 307–337.
65.
Pettigrew, J.D., P.R. Manger, and P.J. Fine (1998) The sensory world of the platypus. Phil. Trans. R. Soc. Lond. B, 353: 1199–1210.
66.
Rosa, M.G.P., R. Gattass, and M. Fiorani (1988) Complete pattern of ocular dominance stripes in V1 of a New World monkey, Cebus apella. Exp. Brain Res., 72: 645–648.
67.
Rosa, M.G.P., R.Gattass, and J.G.M. Soares (1991) A quantitative analysis of cytochrome oxidase-rich patches in the primary visual cortex of Cebus monkeys: topographic distribution and effects of late monocular enucleation. Exp. Brain Res., 84: 195–209.
68.
Rosa, M.G.P., R. Gattass, M. Fiorani, and J.G.M. Soares (1992) Laminar, columnar and topographic aspects of ocular dominance in the primary visual cortex of Cebus monkeys. Exp. Brain Res., 88: 249–264.
69.
Schmued, L.C. (1990) A rapid, sensitive histochemical stain for myelin in frozen brain sections. J. Histochem. Cytochem., 38: 717–720.
70.
Shatz, C.J., S. Lindström, and T.N. Wiesel (1997) The distribution of afferents representing the right and left eyes in the cat’s visual cortex. Brain Res., 131: 103–116.
71.
Sholl, D.A. (1953) Dendritic organization in the neurons of the visual and motor cortices of the cat. J. Anat., 87: 387–406.
72.
Simons, D.J., and T.A. Woolsey (1984) Morphology of Golgi-Cox-impregnated barrel neurons in rat SmI cortex. J. Comp. Neurol., 230: 119– 132.
73.
Tootell, R.B.H., M.S. Silverman, R.L. De Valois, and G.H. Jacobs (1983) Functional organization of the second cortical visual area of primates. Science, 220: 737–739.
74.
Uchizono, K. (1968) Inhibitory and excitatory synapses in vertibrate and invertibrate animals. In Structure and function of inhibitory neuronal mechanisms (ed. by V.C. Euler, S. Skoglund, and U. Soderburg), Pergamon, Oxford, pp. 33– 60.
75.
Welker, C. (1971) Microelectrode delineation of fine grain somatotopic organisation of SmI cerebral neocortex in albino rat. Brain Res., 26: 259–275.
76.
Weller, W.L. (1993) SmI cortical barrels in an Australian marsupial, Trichosurus vulpecula (brush-tailed possum): Structural organisation, patterned distribution, and somatotopic relationships. J. Comp. Neurol, 337: 471–492.
77.
Wong-Riley, M. (1979) Changes in the visual system of monocularly sutured or enucleated cats demonstrable with cytochrome oxidase histochemistry. Brain Res., 171: 11–28.
78.
Woolsey, T.A., and H. Van der Loos (1970) The structural organisation of layer IV in the somatosensory region (SI) of mouse cerebral cortex. Brain Res., 17: 205–242.
79.
Woolsey, T.A., N.L. Dierker, and D.F. Wann (1975) Mouse SmI cortex: qualitative and quantitative classification of Golgi-impregnated barrel neurons. Proc. Natl. Acad. Sci. USA, 72: 2165– 2169.
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