Olfactory Bulb: Synaptic Organization

Chemosensory information transduced by olfactory receptors in the periphery is mapped into a two dimensional array of glomeruli in the olfactory bulb, creating an orderly spatial code of odour identity. In the bulb, local interneurons implement multiple layers of recurrent and lateral inhibition both within and across glomeruli. These synaptic operations can rescale, filter and modulate neural activity patterns of principal neurons, dynamically modifying odour representations being relayed to higher cortical centres.

Keywords: olfactory bulb; glomeruli; dendrodendritic

Figure 1. Olfactory bulb circuits. Schematic showing some of the major synaptic connections in the mammalian olfactory bulb. Green arrows indicate glutamate release at excitatory synapses, red arrows indicate GABA release at inhibitory synapses (both activating ionotropic postsynaptic receptors). ON axons arriving from the periphery enter glomeruli (gray ovals) and release glutamate. This activates a cascade of local circuits: (1) mitral/tufted (M/T) to PG cell reciprocal feedback; (2) PG cell interglomerular lateral inhibition; (3) SA cell interglomerular lateral inhibition; (4) endothelin (ET) cell to PG cell reciprocal feedback; (5) mitral/tufted (M/T) cell intraglomerular synchronization by gap junctions and glutamate spillover; (6) M/T cell lateral inhibition through granule cells (GC); (7) M/T to granule reciprocal feedback; (8) Blanes cell (BC) inhibition of GC; (9) M/T axon collateral excitation of GC; (10) feedback excitation of granule cells from olfactory cortex; (11) neuromodulatory inputs to glomeruli and granule cells, including serotonergic (5-hydroxytryptamine (HT)), cholinergic (acetylcholine (ACh)) and noradrenergic (NA) fibres.
Figure 2. Dendrodendritic reciprocal synapse. Dendrites of mitral, tufted, granule and juxtaglomerular cells are multitasking-signalling devices, doubling as presynaptic and postsynaptic elements. Excitatory-inhibitory reciprocal synapses implement feedback and lateral inhibition between principal neurons (M/T and ET cells) and interneurons (PG and granule cells). The schematic shows operation of a mitral–granule reciprocal synapse: an action potential back-propagating into a mitral cell secondary dendrite (blue) opens voltage-gated calcium channels, and calcium entry releases glutamate (green arrow) from which activates AMPA and NMDA receptors on granule cell spines (yellow). Spine depolarization opens granule cell calcium channels, releasing GABA (red arrow) which inhibits the mitral cell by activating dendritic GABAA receptors.
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 References
    Adrian ED (1942) Olfactory reactions in the brain of the hedgehog. The Journal of Physiology (London) 100: 459–473.
    Aungst JL, Heyward PM, Puche AC et al. (2003) Centre-surround inhibition among olfactory bulb glomeruli. Nature 426: 623–629.
    Castillo PE, Carleton A, Vincent JD and Lledo PM (1999) Multiple and opposing roles of cholinergic transmission in the main olfactory bulb. The Journal of Neuroscience 19: 9180–9191.
    Christie JM, Bark C, Hormuzdi SG et al. (2005) Connexin36 mediates spike synchrony in olfactory bulb glomeruli. Neuron 46: 761–772.
    Gutierrez-Mecinas M, Crespo C, Blasco-Ibanez JM et al. (2005) Soluble guanylyl cyclase appears in a specific subset of periglomerular cells in the olfactory bulb. The European Journal of Neuroscience 21: 1443–1448.
    Hardy A, Palouzier-Paulignan B, Duchamp A, Royet JP and Duchamp-Viret P (2005) 5-Hydroxytryptamine action in the rat olfactory bulb: in vitro electrophysiological patch-clamp recordings of juxtaglomerular and mitral cells. Neuroscience 131: 717–731.
    Kosaka T and Kosaka K (2005) Structural organization of the glomerulus in the main olfactory bulb. Chemical Senses 30(Suppl. 1): i107–i108.
    Murphy GJ, Glickfeld LL, Balsen Z and Isaacson JS (2004) Sensory neuron signaling to the brain: properties of transmitter release from olfactory nerve terminals. The Journal of Neuroscience 24: 3023–3030.
    Pressler RT and Strowbridge BW (2006) Blanes cells mediate persistent feed forward inhibition onto granule cells in the olfactory bulb. Neuron 49: 889–904.
    Rall W, Shepherd GM, Reese TS and Brightman MW (1966) Dendrodendritic synaptic pathway for inhibition in the olfactory bulb. Experimental Neurology 14: 44–56.
    Schneider SP and Macrides F (1978) Laminar distributions of interneurons in the main olfactory bulb of the adult hamster. Brain Research Bulletin 3: 73–82.
    Schoppa NE (2006) Synchronization of olfactory bulb mitral cells by precisely timed inhibitory inputs. Neuron 49: 271–283.
 Further Reading
    Dulac C and Wagner S (2006) Genetic analysis of brain circuits underlying pheromone signaling. Annual Review of Genetics 40: 449–467.
    Egger V and Urban NN (2006) Dynamic connectivity in the mitral cell–granule cell microcircuit. Seminars in Cell and Developmental Biology 17: 424–432.
    Isaacson JS and Strowbridge BW (1998) Olfactory reciprocal synapses: dendritic signaling in the CNS. Neuron 20: 749–761.
    Lledo PM, Gheusi G and Vincent JD (2005) Information processing in the mammalian olfactory system. Physiological Reviews 85: 281–317.
    Mori K, Nagao H and Yoshihara Y (1999) The olfactory bulb: coding and processing of odour molecule information. Science 286: 711–715.
    Schoppa NE and Urban NN (2003) Dendritic processing within olfactory bulb circuits. Trends in Neurosciences 26: 501–506.
    book Shepherd GM, Chen WR and Greer CA (2004) "Olfactory bulb". In: Shepherd GM (ed.) The Synaptic Organization of the Brain, 5th edn, pp. 165–216. New York, NY: Oxford University Press.
    book Shipley MT, McLean JH and Ennis M (1995) "Olfactory system". In: Paxinos G (ed.) The Rat Nervous System, 2nd edn, pp. 899–926. San Diego, CA: Academic Press, Inc.
    Wachowiak M and Shipley MT (2006) Coding and synaptic processing of sensory information in the glomerular layer of the olfactory bulb. Seminars in Cell & Developmental Biology 17: 411–423.
    Wilson RI and Mainen ZF (2006) Early events in olfactory processing. Annual Review of Neuroscience 29: 163–201.
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Lowe, Graeme(Dec 2007) Olfactory Bulb: Synaptic Organization. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0020289]