The Neuropil of the Periglomerular Region of the Olfactory Bulb

1971 ◽  
Vol 9 (2) ◽  
pp. 379-409
Author(s):  
A. J. PINCHING ◽  
T. P. S. POWELL

The periglomerular region of the olfactory bulb, apart from containing the somata and stem dendrites of the cells contributing to the glomeruli, is the sole region of distribution of the periglomerular cell thin dendrites and the short-axon cell dendrites. It is also the major site of termination of all axons to the glomerular layer except the olfactory axons - i.e. tufted cell collaterals, periglomerular cell and short-axon cell axons and centrifugal fibres. Its characteristic neuropil has been studied with the electron microscope to define the cells of origin of the types of neuronal process and their synaptic relationships. Three types of axon terminals have been found: those with spherical, large flattened and small flattened vesicles, which are deduced to derive from tufted cell collaterals or centrifugal fibres, periglomerular cell and short axon-cell axons respectively; those with spherical vesicles are consistently associated with asymmetrical membrane thickenings and those with either type of flattened vesicles with symmetrical thickenings. The thin periglomerular cell dendrites are very irregular and often have a somewhat dense cytoplasm, rich in ribosomes; they may become extended into very attenuated glia-like sheets that surround the mitral or tufted cell stem dendrites, from which they may receive synaptic contacts. Such dendrites also receive some synapses from all 3 types of axon in the periglomerular region. The short-axon cell dendrites are thick and varicose and show no sign of synaptic specialization oriented from them; they have few spines but receive many asymmetrical-type synapses on their shafts. Both axon terminal types synapsing with symmetrical thickenings are also found on the shafts. The evidence obtained from the study of normal material is summarized and the various cellular roles considered. In the light of observations on the olfactory bulb, it is suggested that dendrites may be divided into 2 major classes: those that only receive synapses (Class A) and those that make synaptic contacts as well as receiving them (Class B). Further comparisons with the deep layers of the bulb and physiological implications are discussed.

1971 ◽  
Vol 9 (2) ◽  
pp. 347-377
Author(s):  
A. J. PINCHING ◽  
T. P. S. POWELL

The neuropil of the glomeruli of the rat olfactory bulb has been studied with the electron microscope with a view to elucidating the type of processes involved - dendrites, appendages and axons - their cellular identity, and the synaptic relationships they establish. The problems encountered in defining these are considered and criteria based on the previous study of neuron types and on examination of serial sections are put forward. The glomeruli are large structures containing many thousands of processes and are the sole site of termination of the olfactory receptor axons. The terminals of the latter are characteristically electron-dense, allowing identification in normal material; they run through the glomeruli making many synapses by means of spherical vesicles and asymmetrical thickenings on to all types of dendritic profile. The glomerular dendritic arborizations of mitral and tufted cells, which are indistinguishable from each other, start as large, fairly regular, pale profiles but become increasingly varicose as they branch and diminish in size. They regularly show groups of spherical vesicles, often in association with asymmetrical synaptic thickenings directed from the dendrite; these are typically associated with return, reciprocal, synapses of the symmetrical type from profiles containing large flattened vesicles. These latter profiles are those of the dendrites and gemmules of periglomerular cells; the dendrites are of irregular outline and give rise to many appendages, mostly gemmules making synaptic contact with mitral or tufted cell dendrites. A small number of pale axon terminals containing either small or large flattened vesicles, derived from short-axon and periglomerular cells respectively, synapse with symmetrical thickenings on to the periglomerular cell dendritic processes. Close associations of particular types of axo-dendritic and dendro-dendritic synapses on interconnecting processes, termed synaptic patterns, are described and their significance considered. The nature of the glomerular interactions is discussed and then placed in the context of other, smaller glomeruli in the central nervous system; certain common principles of glomeruli are suggested.


Author(s):  
Thomas P. Eiting ◽  
Matt Wachowiak

AbstractSniffing—the active control of breathing beyond passive respiration—is used by mammals to modulate olfactory sampling. Sniffing allows animals to make odor-guided decisions within ~200 ms, but animals routinely engage in bouts of high-frequency sniffing spanning several seconds; the impact of such repeated odorant sampling on odor representations remains unclear. We investigated this question in the mouse olfactory bulb, where mitral and tufted cells (MTCs) form parallel output streams of odor information processing. To test the impact of repeated odorant sampling on MTC responses, we used two-photon imaging in anesthetized male and female mice to record activation of MTCs while precisely varying inhalation frequency. A combination of genetic targeting and viral expression of GCaMP6 reporters allowed us to access mitral (MC) and superficial tufted cell (sTC) subpopulations separately. We found that repeated odorant sampling differentially affected responses in MCs and sTCs, with MCs showing more diversity than sTCs over the same time period. Impacts of repeated sampling among MCs included both increases and decreases in excitation, as well as changes in response polarity. Response patterns across ensembles of simultaneously-imaged MCs reformatted over time, with representations of different odorants becoming more distinct. MCs also responded differentially to changes in inhalation frequency, whereas sTC responses were more uniform over time and across frequency. Our results support the idea that MCs and TCs comprise functionally distinct pathways for odor information processing, and suggest that the reformatting of MC odor representations by high-frequency sniffing may serve to enhance the discrimination of similar odors.


1972 ◽  
Vol 10 (3) ◽  
pp. 585-619
Author(s):  
A. J. PINCHING ◽  
T. P. S. POWELL

An ultrastructural study of degeneration in axon terminals of the glomerular layer of the rat olfactory bulb is described, concentrating particularly on the sequence of degeneration in the olfactory nerve terminals and the long-term events in the degeneration process in several terminal types. Olfactory nerve terminal degeneration is divided into 5 stages, representing parts of the sequential changes taking place in the terminal after fibre section. The main features in the sequence are as follows: I. Swelling of the terminal and its vesicles. II. Initial shrinkage of the terminal, while vesicles remain swollen and some are distorted. III. Further shrinkage and darkening of the terminal, distortion of many of the vesicles and some mitochondrial swelling. IV. Extreme shrinkage of the terminal and loss of cytoplasmic detail; synaptic contact still intact or partially separated. V. Disappearance of the presynaptic terminal and persistence of the post-synaptic thickening. The validity of the observation of the persistence of post-synaptic membrane thickenings is considered and evidence adduced in its favour, both qualitative and quantitative. Characteristics of the newly apposed profiles are described, including cisternae and alveolate vesicles; the extracellular material of the synaptic cleft is considered in the light of its association with persisting thickenings. The relevance of these associated Structures is discussed in terms of function and development. Spontaneous degeneration of olfactory nerves and the degeneration of the vomeronasal nerves in the accessory olfactory bulb are described, as well as Stage V in the degeneration of other terminal types of the glomeruli, following various lesions; persistence of post-synaptic thickenings after the degeneration of terminals showing symmetrical membrane thickenings is included. Observations on the persistence of post-synaptic thickenings in various sites in the central and peripheral nervous systems are reviewed in the context of the present study; the problems arising out of the morphological identification of apposition or reinnervations are discussed.


2014 ◽  
pp. 133-160 ◽  
Author(s):  
Shin Nagayama ◽  
Kei M. Igarashi ◽  
Hiroyuki Manabe ◽  
Kensaku Mori

Microscopy ◽  
2019 ◽  
Vol 68 (4) ◽  
pp. 316-329
Author(s):  
Eiji Notsu ◽  
Kazunori Toida

Abstract The olfactory bulb (OB) contains various interneuron types that play key roles in processing olfactory information via synaptic contacts. Many previous studies have reported synaptic connections of heterogeneous interneurons in superficial OB layers. In contrast, few studies have examined synaptic connections in deep layers because of the lack of a selective marker for intrinsic neurons located in the deeper layers, including the mitral cell layer, internal plexiform layer (IPL) and granule cell layer. However, neural circuits in the deep layers are likely to have a strong effect on the output of the OB because of the cellular composition of these regions. Here, we analyzed the calbindin-immunoreactive neurons in the IPL, one of the clearly neurochemically defined interneuron types in the deep layers, using multiple immunolabeling and confocal laser scanning microscopy combined with electron microscopic three-dimensional serial-section reconstruction, enabling correlated laser and volume electron microscopy (EM). Despite a resemblance to the morphological features of deep short axon cells, IPL calbindin-immunoreactive (IPL-CB-ir) neurons lacked axons. Furthermore, multiple immunolabeling for plural neurochemicals indicated that IPL-CB-ir neurons differed from any interneuron types reported previously. We identified symmetrical synapses formed by IPL-CB-ir neurons on granule cells (GCs) using correlated laser and volume EM. These synapses might inhibit GCs and thus disinhibit mitral and tufted cells. Our present findings indicate, for the first time, that IPL-CB-ir neurons are involved in regulating the activities of projection neurons, further suggesting their involvement in synaptic circuitry for output from the deeper layers of the OB, which has not previously been clarified.


Physiology ◽  
2012 ◽  
Vol 27 (4) ◽  
pp. 200-212 ◽  
Author(s):  
Claudia Lodovichi ◽  
Leonardo Belluscio

In mammals, smell is mediated by odorant receptors expressed by sensory neurons in the nose. These specialized receptors are found both on olfactory sensory neurons' cilia and axon terminals. Although the primary function of ciliary odorant receptors is to detect odorants, their axonal role remains unclear but is thought to involve axon guidance. This review discusses findings that show axonal odorant receptors are indeed functional and capable of modulating neural connectivity.


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