The Morphology of the Granule Cells of the Olfactory Bulb

1970 ◽  
Vol 7 (1) ◽  
pp. 91-123 ◽  
Author(s):  
J. L PRICE ◽  
T. P. S POWELL

The granule cells of the olfactory bulb of the rat have been studied in material prepared by the Golgi-Kopsch method for examination with the light microscope, and in material examined with the electron microscope. With the Golgi method, the granule cells are found to have no process which can be identified as a typical axon, but from the superficial aspect of the somata stout peripheral processes arise and pass into the overlying external plexiform layer, while from the opposite side of the cell body several thinner deep dendntes extend towards the deeper parts of the bulb. Both types of processes, as well as the perikarya, have numerous spine-like appendages. On the distal portions of the peripheral processes in the external plexiform layer the appendages are much larger than on the deeper parts of the cell. The deep dendrites have localized swellings along their length which give them a varicose appearance, appendages often arise from these varicosities. The electron-microscopic features of the granule cells correspond well with the appearance of these cells in material impregnated with the Golgi method. The cell somata are small, with very little cytoplasm, and have a relatively large nucleus. The peripheral processes can be identified passing superficially from the perikarya of the granule cells; at their junction with the cell body their appearance is typically dendritic; all the cytoplasmic organelles found in the cytoplasm extend into these processes and none of the features of the initial segments of axons are found. In the external plexiform layer large spine-like appendages, which have been termed ‘gemmules’, arise from the distal portions of the peripheral processes, and participate in reciprocal synapses with the dendrites of mitral and tufted cells. The deep dendrites are much finer than the peripheral processes, and the varicosities which are seen in Golgi material may also be found with the electron microscope. Spines are found on all parts of the granule cells in the granule cell layer, including the peripheral processes, the perikarya and the deep dendrites. In addition to a spine apparatus, these spines commonly have numerous inclusions, including mitochondria, ribosomes, and vesicles which are the same size and shape as the synaptic vesicles present in the gemmules; no synapses oriented away from the spines have ever been found.

1970 ◽  
Vol 7 (3) ◽  
pp. 631-651
Author(s):  
J. L. PRICE ◽  
T. P. S. POWELL

A description is given of the mitral and short axon cells of the olfactory bulb of the rat from Golgi material examined with the light microscope and from material examined with the electron microscope. The mitral cells are large neurons with primary and secondary dendrites which both extend into the overlying external plexiform layer, although only the primary dendrite enters the glomerular formations. No predominant antero-posterior orientation of the secondary dendrites has been found. Within the glomeruli the mitral cell dendrites are in synaptic contact with the olfactory nerves and also with the periglomerular cells, but elsewhere the only synapses on the mitral cells are the ‘reciprocal synapses’ with the granule cells. Synaptic-type vesicles are found in all parts of the mitral cells, including the axon initial segments; they appear to be especially concentrated in the distal portions of the dendrites. Several types of short axon cells have been found in the granule cell layer in Golgi-impregnated material. Their cell bodies can also be distinguished with the electron microscope, and from previous work it is probable that the axons of at least some of these cells form flattened-vesicle symmetrical synapses upon the granule cells.


1970 ◽  
Vol 7 (1) ◽  
pp. 157-187
Author(s):  
J. L. PRICE ◽  
T. P. S. POWELL

An experimental investigation has been made of the site and mode of termination of the 3 groups of afferent fibres to the olfactory bulb which come from more caudal parts of the cerebral hemisphere. Lesions have been placed in the relevant parts of the brain of the rat and the resulting degeneration of axon terminals in the olfactory bulb studied with the electron microscope. All 3 groups of these extrinsic afferent fibres end in asymmetrical synapses upon the granule cells, and they have a differential termination upon its various processes. The possibility that these fibres also end upon other cells in the bulb (particularly the short-axon and periglomerular cells) cannot be excluded. The centrifugal fibres end upon gemmules in the deep half of the external plexiform layer only; no degenerating terminals were found in relation to the glomeruli although degenerating centrifugal axons are present here. The fibres of the anterior commissure terminate upon spines and varicosities of the deep dendrites and upon somatic spines of the granule cells. After lesions of the anterior olfactory nucleus, degenerating terminals were found in the ipsilateral olfactory bulb, which could not be ascribed to the centrifugal fibres or to the fibres of the anterior commissure, as they ended upon the spines of peripheral processes in the granule cell layer, and upon gemmules in the superficial as well as in the deep half of the external plexiform layer. It is proposed that these terminals are those of the axon collaterals from the ipsilateral anterior olfactory nucleus. The axons which form symmetrical synapses, and many which form asymmetrical synapses, do not degenerate even after a lesion immediately behind the olfactory bulb, and are therefore intrinsic to the bulb. It is suggested that the axons which are associated with symmetrical synapses are those of the short-axon cells, and the asymmetrical synapses are formed by the axon collaterals of the mitral and tufted cells.


2021 ◽  
Vol 15 ◽  
Author(s):  
Fu-Wen Zhou ◽  
Adam C. Puche

Olfactory bulb and higher processing areas are synaptically interconnected, providing rapid regulation of olfactory bulb circuit dynamics and sensory processing. Short-term plasticity changes at any of these synapses could modulate sensory processing and potentially short-term sensory memory. A key olfactory bulb circuit for mediating cortical feedback modulation is granule cells, which are targeted by multiple cortical regions including both glutamatergic excitatory inputs and GABAergic inhibitory inputs. There is robust endocannabinoid modulation of excitatory inputs to granule cells and here we explored whether there was also endocannabinoid modulation of the inhibitory cortical inputs to granule cells. We expressed light-gated cation channel channelrhodopsin-2 (ChR2) in GABAergic neurons in the horizontal limb of the diagonal band of Broca (HDB) and their projections to granule cells in olfactory bulb. Selective optical activation of ChR2 positive axons/terminals generated strong, frequency-dependent short-term depression of GABAA-mediated-IPSC in granule cells. As cannabinoid type 1 (CB1) receptor is heavily expressed in olfactory bulb granule cell layer (GCL) and there is endogenous endocannabinoid release in GCL, we investigated whether activation of CB1 receptor modulated the HDB IPSC and short-term depression at the HDB→granule cell synapse. Activation of the CB1 receptor by the exogenous agonist Win 55,212-2 significantly decreased the peak amplitude of individual IPSC and decreased short-term depression, while blockade of the CB1 receptor by AM 251 slightly increased individual IPSCs and increased short-term depression. Thus, we conclude that there is tonic endocannabinoid activation of the GABAergic projections of the HDB to granule cells, similar to the modulation observed with glutamatergic projections to granule cells. Modulation of inhibitory synaptic currents and frequency-dependent short-term depression could regulate the precise balance of cortical feedback excitation and inhibition of granule cells leading to changes in granule cell mediated inhibition of olfactory bulb output to higher processing areas.


2005 ◽  
Vol 37 (4) ◽  
pp. 276-282 ◽  
Author(s):  
Zhao-Ping Qin ◽  
Shu-Ming Ye ◽  
Ji-Zeng Du ◽  
Gong-Yu Shen

Abstract The distribution of calbindin, calretinin and parvalbumin during the development of the mouse main olfactory bulb (MOB) was studied using immunohistochemistry techniques. The results are as follows: (1) calbindin-immunoreactive profiles were mainly located in the glomerular layer, and few large calbindin-immunoreactive cells were found in the subependymal layer of postnatal day 10 (P1 0) to postnatal day 40 (P40) mice; (2) no calbindin was detected in the mitral cell layer at any stage; (3) calretinin-immunoreactive profiles were present in all layers of the main olfactory bulb at all stages, especially in the olfactory nerve layer, glomerular layer and granule cell layer; (4) parvalbumin-immunoreactive profiles were mainly located in the external plexiform layer (except for P10 mice); (5) weakly stained parvalbumin-immunoreactive profiles were present in the glomerular layer at all stages; and (6) no parvalbumin was detected in the mitral cell layer at any stage.


2003 ◽  
Vol 90 (2) ◽  
pp. 644-654 ◽  
Author(s):  
Brian Halabisky ◽  
Ben W. Strowbridge

Recurrent and lateral inhibition play a prominent role in patterning the odor-evoked discharges in mitral cells, the output neurons of the olfactory bulb. Inhibitory responses in this brain region are mediated through reciprocal synaptic connections made between the dendrites of mitral cells and GABAergic interneurons. Previous studies have demonstrated that N-methyl-d-aspartate (NMDA) receptors on interneurons play a critical role in eliciting GABA release at reciprocal dendrodendritic synapses. In acute olfactory bulb slices, these receptors are tonically blocked by extracellular Mg2+, and recurrent inhibition is disabled. In the present study, we examined the mechanisms by which this tonic blockade could be reversed. We demonstrate that near-coincident activation of an excitatory pathway to the proximal dendrites of GABAergic interneurons relieves the Mg2+ blockade of NMDA receptors at reciprocal dendrodendritic synapses and greatly facilitates recurrent inhibition onto mitral cells. Gating of recurrent and lateral inhibition in the presence of extracellular Mg2+ requires γ-frequency stimulation of glutamatergic axons in the granule cell layer. Long-range excitatory axon connections from mitral cells innervated by different subpopulations of olfactory receptor neurons may provide a gating input to granule cells, thereby facilitating the mitral cell lateral inhibition that contributes to odorant encoding.


2019 ◽  
Vol 45 (1) ◽  
pp. 3-13
Author(s):  
Brittnee McDole ◽  
Rachel Berger ◽  
Kathleen Guthrie

Abstract Adult-born neurons produced in the dentate gyrus subgranular zone (SGZ) develop as excitatory hippocampal granule cells (GCs), while those from the subventricular zone (SVZ) migrate to the olfactory bulb (OB), where most develop as GABAergic olfactory GCs. Both types of neurons express TrkB as they mature. Normally ~50% of new olfactory GCs survive, but survival declines if sensory drive is reduced. Increases in endogenous brain-derived neurotrophic factor (BDNF) in hippocampus, particularly with wheel running, enhance dentate GC survival. Whether survival of new olfactory GCs is impacted by augmenting BDNF in the OB, where they mature and integrate, is not known. Here, we determined if increasing OB BDNF expression enhances survival of new GCs, and if it counters their loss under conditions of reduced sensory activity. Neurogenesis was assessed under normal conditions, and following unilateral naris occlusion, in mice overexpressing BDNF in the granule cell layer (GCL). OB BDNF levels were significantly higher in transgenic mice compared to controls, and this was maintained following sensory deprivation. Bromodeoxyuridine (BrdU) cell birth dating showed that at 12–14 days post-BrdU, numbers of new GCs did not differ between genotypes, indicating normal recruitment to the OB. At later intervals, transgenic and control mice showed levels of GC loss in deprived and nondeprived animals that were indistinguishable, as was the incidence of apoptotic cells in the GCL. These results demonstrate that, in contrast to new dentate GCs, elevations in endogenous BDNF do not enhance survival of adult-born olfactory GCs.


Author(s):  
R. A. Waugh ◽  
J. R. Sommer

Cardiac sarcoplasmic reticulum (SR) is a complex system of intracellular tubules that, due to their small size and juxtaposition to such electron-dense structures as mitochondria and myofibrils, are often inconspicuous in conventionally prepared electron microscopic material. This study reports a method with which the SR is selectively “stained” which facilitates visualizationwith the transmission electron microscope.


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