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PubMed1985

Immunochemical identification of subgroups of vomeronasal nerve fibers and their segregated terminations in the accessory olfactory bulb.

Vomeronasal nerve (VNN) fibers and their terminations in the accessory olfactory bulb (AOB) were studied immunohistochemically using 3 monoclonal antibodies (MAbs). One MAb (R2D5) labeled all VNN fibers. Another MAb (R4B12) labeled a subgroup of the VNN fibers which terminated in the rostrolateral glomeruli in the AOB. The third MAb (R5A10) recognized a complementary subgroup of the VNN which terminated in the caudomedial portion of the AOB. These results for the first time show occurrence of subtypes in the VNN axons with segregated terminations in the AOB.

Animals↗
PubMed1989

Axonal redirection at the dorsoventral intraretinal boundary.

Cobaltous-lysine applied to the goldfish optic nerve backfilled retinal ganglion cells and their axons. Confined to the ventronasal and ventrotemporal retina was a small population of retinal ganglion cells whose axons traveled dorsally and parallel to the retinal margin. On reaching the boundary between dorsal and ventral retina, the axons arched, joined radially oriented bundles of axons, and traveled toward the optic disk. Control studies showed that the axons came from retinal ganglion cells rather than from retinopetal cells. The somatic area of retinal ganglion cells (RGCs) with circumferential axons was 30-50 microns, and was similar to that of average ganglion cells. The axons of these cells coursed between the optic fiber and ganglion cell layers or between the ganglion cell and inner plexiform layers. Many somata were displaced slightly toward the inner plexiform layer, but were not really displaced ganglion cells. The aberrant axonal trajectory may be related to the slightly displaced location of the cell. However, ganglion cells that are displaced to the edge of the inner nuclear layer usually have radially coursing axons. We digitized the coordinates of the bending points and the dorsoventral retinal boundary. On average, the bending points occurred within 100 microns of the dorsoventral retinal border. These findings suggest that some molecular, rather than mechanical, factor at the dorsoventral retinal boundary alters the course of the circumferential axons. Furthermore, because there are cells with circumferential axons throughout the ventral retina, the data imply that at least ventral RGC axons avoid mingling with the axons from dorsal RGCs.

Animals↗
PubMed1983

An electrophysiological study of the accessory olfactory bulb in the rabbit--I. Analysis of electrically evoked potential fields.

Following electrical stimulation of the vomeronasal nerves, the primary olfactory nerves, the lateral olfactory tract and the corticomedial amygdala, we have made a study of evoked potentials in the rabbit accessory olfactory bulb. Vomeronasal nerve stimulation evoked a complex field potential consisting of a compound action potential followed by 4 negative waves (N1, N2, N3, N4). In contrast to the field potential elicited in the main olfactory bulb following primary olfactory nerve stimulation, there was either no evoked wave or only a weak positive component of the field in the accessory bulb. Amygdala stimulation caused a long latency, long duration negative-positive dipolar field potential in the accessory olfactory bulb. Both antidromic and orthodromic field potentials showed sign reversal when the electrode penetrated the bulb at a point corresponding to the lower border of the mitral cell band. Stimulation of the lateral olfactory tract elicited a weak, short-latency wave which did not show any sign reversal when the electrode was lowered into the accessory bulb. This wave was presumably due to fibres arising in the main bulb and projecting through the accessory bulb into the lateral olfactory tract. Electrical stimulation of the primary olfactory nerves did not induce any response in the accessory bulb neither did vomeronasal nerve stimulation evoke a response in the main olfactory bulb. The origin of these potential fields is discussed and it is concluded that the synaptic organization of the accessory olfactory bulb resembles that of the main olfactory bulb in lower vertebrates. There is no detectable communication between the two olfactory systems.

Amygdala↗
PubMed2004

[The application of functional magnetic resonance imaging for the assessment of localisation and activation of cortex smell centers depending on stimulus used in normal volunteers].

PURPOSE: To determine the location and activation of brain smell centres in response to olfactory nerve-mediated and combined olfactory and trigeminal nerve-mediated stimulation using functional MRI. SUBJECTS AND METHODS: MRI brain scans were performed in 10 healthy volunteers, aged 22-36 years, right handed, non-smokers, without any CNS diseases, not taking any medication to determine and compare the activated cortex regions during stimulation by patchouli and geraniol. fMRI was performed on Siemens Magnetom Vision 1,5 T imager with a head-neck coil using SE and EPI sequences. Functional MR imaging studies were obteined in the same plane as T11 eighted images in an axial plane by using echo planar imaging. In each of 14 contiguous planes parallel to anterior-posterior commisure line images that depicted blood-oxygen-level-dependent (BOLD) effect were acquired. The subject inhaled odorized air through both nares for 30 seconds. During 30-second off period the subject received room air at the same flow rate. For each of activated cortical zones the number of pixels activated was calculated. Image reconstruction and analysis were performed using standard imager software installed in MR imager. RESULTS: The exposure to the olfactory and trigeminal nerve-mediated stimuli showed more activated regions than to the olfactory nerve-mediated stimuli. Geraniol evoked the biggest activation in orbitofrontal region, cingulate gyrus and insula. Patchouli activated orbitofrontal cortex, insula, parahippocampal gyrus, occipital cortex, and corpus callosum. CONCLUSIONS: The results of present study confirm the ability to localize patterns of cortical activity with fMRI during olfactory tasks.

Adult↗
PubMed1991

Receptor-mediated transport of human recombinant nerve growth factor from olfactory bulb to forebrain cholinergic nuclei.

Receptors for nerve growth factor are present in the olfactory bulb and in cholinergic nuclei that send projections to the olfactory bulb. The retrograde transport of 125I-labeled recombinant human nerve growth factor (rhNGF) was demonstrated in the rat 18 h following an injection of [125I]rhNGF into the left olfactory bulb. In each of six animals, [125I]rhNGF label was observed in the ipsilateral horizontal limb of the diagonal band and, in four of the 6 animals, in the vertical limb of the diagonal band. Label was not observed in any other brain region except within the injected olfactory bulb. The transport of label to the diagonal band was blocked by the injection of 170-fold greater concentration of unlabeled rhNGF. Emulsion autoradiography of hematoxylin/eosin counterstained sections revealed silver grains clustered over numerous cell profiles that resembled neurons. In contrast, cerebellar injections of [125I]rhNGF, with or without unlabeled rhNGF, did not label diagonal band neurons, nor the lateral vestibular or red nuclei, from which originate the primary cholinergic afferents to cerebellum. The receptor-dependent transport of NGF from olfactory bulb to forebrain cholinergic nuclei suggests that this projection, unlike pontomesencephalic cholinergic pathways, may be responsive to endogenous NGF or exogenously administered rhNGF.

Animals↗
PubMed1993

Nervus terminalis lesions: I. No effect on pheromonally induced testosterone surges in the male hamster.

The involvement of the nervus terminalis or terminal nerve in the pheromonally induced testosterone surge in the male hamster was investigated. Blood was collected from male hamsters not exposed to odor (baseline), and then a week later from the same hamsters exposed to the odor of vaginal discharge from an estrous female. Terminal nerve lesions, forebrain lesions, or sham surgeries were performed, and blood was collected again with and without odor stimulation. Serum testosterone levels were assessed by radioimmunoassay. None of the surgical procedures interrupted the ability of the male hamsters to demonstrate an increase in serum testosterone following exposure to the odor of an estrous female. We conclude that the terminal nerve is not necessary for this pheromonally mediated neuroendocrine reflex.

Animals↗
PubMed2000

Nerve growth factor applied onto the olfactory epithelium alleviates degenerative changes of the olfactory receptor neurons following axotomy.

The olfactory neuroepithelium of the mammalian nervous system manifests continuous neurogenesis throughout life. Recent studies suggest that neurotrophic factors and their receptors may play a role in the regulation of development and regeneration in the olfactory system. However, there have been very few in vivo studies investigating the effect of exogenous neurotrophic factors in the olfactory system. In the present study, nerve growth factor (NGF) and brain-derived neurotrophic factor (BDNF) were administered into the rat olfactory mucosa for 5 days just after the transection of the olfactory nerve. We then examined the effect of exogenous neurotrophic factors on the degenerative changes in axotomized olfactory receptor neurons (ORNs). Further, we examined the location of their receptors, Trk A and Trk B. We found that both mature and immature ORNs expressed more intense signals for olfactory marker protein and beta-tubulin mRNAs, respectively, when NGF was applied to the axotomized olfactory neuroepithelium for 5 days, compared to the ORNs of saline-treated controls. BDNF at a 10 microg total dose did not show this effect. The effect of NGF applied onto the olfactory epithelium is consistent with the immunohistochemical finding that Trk A was present in the dendrites and axon bundles in normal and axotomized ORNs. These results suggest that NGF may protect the degenerative changes in mature and immature ORNs following axotomy through the binding to the Trk A receptor located on the surface of the olfactory epithelium.

Animals↗
PubMed1989

The adult rat olfactory system expresses microtubule-associated proteins found in the developing brain.

We have compared the expression and localization of neuronal microtubule-associated proteins (MAPs) MAP5, MAP2, and tau in the adult rat olfactory system and cerebral cortex. Each of these MAPS is known to exist as distinct "early" and "late" forms in the developing and adult brain, respectively. Because axonal growth and dendritic reinnervation continue in the adult olfactory system, it can serve as a test of whether expression of early MAP forms is necessary for neuronal growth and plasticity. We found that for all three MAPs, the early forms continue to be expressed in the adult olfactory system, whereas the cerebral cortex switches to the late forms during neuronal maturation (between 10 and 20 days of age in the rat). For MAP2, the 6 kb mRNA and low-molecular weight MAP2c, both of which are typically found in embryonic tissue, persist in the adult olfactory bulb. For MAP5, an early highly phosphorylated form, MAP5a, is present throughout the brain and disappears during maturation simultaneously with a several-fold drop in the overall level of MAP5. However, in the olfactory bulb, MAP5 levels do not fall, and MAP5a persists in the adult. The early form of tau is also prominent in the adult olfactory bulb and, by immunohistochemistry, is mainly confined to a subset of olfactory axons, the vomeronasal nerve. Thus, in the adult olfactory bulb, both MAP protein synthesis and phosphorylation conform to a pattern associated with the developing brain. Immunohistochemistry also showed that MAP5 is concentrated in the olfactory nerve axons and the mitral cell dendrites of the olfactory bulb, i.e., exactly those elements that are involved in the olfactory nerve innervation that takes place in the adult. These results suggest that the expression of the early MAP forms is closely associated with neurite outgrowth and plasticity.

Animals↗
PubMed1982

Regeneration of olfactory sensory axons into transplanted segments of peripheral nerve.

Isolated segments of cervical sympathetic trunk were transplanted onto the olfactory mucosa and bulb in adult rats, and olfactory nerve fascicles were sectioned in the region of the transplant. Olfactory axons, presumably arising from newly-formed sensory neurons, were observed to grow into the transplanted segments of peripheral nerve and were ensheathed by Schwann cells of the transplant. The axons were ensheathed as large bundles containing many axons in direct contact, an arrangement characteristic of the normal olfactory nerves, and not of the normal sympathetic trunk. However, single Schwann cell 'units' were each surrounded by basal lamina, an arrangement typical of the sympathetic trunk, not of olfactory nerves. The results indicate that olfactory axons, like other peripheral axons, are capable of directing some aspects of the manner in which they are ensheathed by Schwann cells.

Animals↗
PubMed1981

Regeneration of vomeronasal nerves into the main olfactory bulb in the mouse.

After surgical section of the vomeronasal nerves the neurosensory cells in the vomeronasal epithelium die. Electron microscopy has been used to demonstrate that their axons, and synaptic terminals in the accessory olfactory bulb degenerate and are removed by phagocytic astroglia. The vacated postsynaptic sites in the accessory bulb persist, and are not re-innervated, either by vomeronasal or olfactory axons, as long as 150 days post-operatively. However, new neurosensory cells which are produced in the vomeronasal epithelium after vomeronasal nerve section do form axons. Light and electron microscope autoradiography of axonally transported material has been used to show that some of these axons grow back into the cranial cavity and form glomeruli in the main olfactory bulb, in regions where it is damaged or de-afferented. The regenerated vomeronasal glomeruli contain synapses between vomeronasal nerve terminals and dendrites of main bulb neurons.

Animals↗
PubMed2000

Expression of neuropeptide Y in olfactory ensheathing cells during prenatal development.

Neuropeptide Y (NPY) is expressed in a special type of glial cell, the olfactory ensheathing cells, that surround the axons of olfactory sensory neurons on their way from the olfactory epithelium to the glomeruli in the olfactory bulb. The expression of NPY in ensheathing cells was examined during prenatal development of the olfactory system by using immunohistochemistry and in situ hybridization. NPY expression was compared with the expression of growth associated protein-43, olfactory marker protein, the low-affinity nerve growth factor receptor (p75) and S-100, factors expressed in the olfactory system at known stages of development. NPY-like immunoreactivity (NPY-LI) and NPY mRNA expression was first detected in the olfactory nerve layer of the olfactory bulb at embryonic day 15. From embryonic day 16 and onward, a clear segregation could be observed in the intensity of both NPY-LI and NPY mRNA expression within the olfactory nerve layer. NPY expression was most intense in the inner part of the olfactory nerve layer. In the outer olfactory nerve layer, a clear decrease in NPY expression was observed. The inner olfactory nerve layer, showing high NPY expression, did not stain for S-100 or p75. However, NPY-LI was found to coexist with S-100-LI from the outer olfactory nerve layer until the olfactory epithelium and with p75-LI in cells surrounding the olfactory nerve. These results show that NPY is expressed in ensheathing cells before olfactory sensory neurons mature and the formation of the glomerular layer starts. NPY might be involved in the guidance, growth, or both, of olfactory sensory axons toward their target glomeruli in the olfactory bulb or have a function in the maturation of the olfactory sensory neurons.

Animals↗
PubMed1992

Drainage of brain extracellular fluid into blood and deep cervical lymph and its immunological significance.

Cerebral extracellular fluids drain from brain to blood across the arachnoid villi and to lymph along certain cranial nerves (primarily olfactory) and spinal nerve root ganglia. Quantification of the connection to lymph in rabbit, cat and sheep, using radiolabelled albumin as a marker of flow, indicates that a minimum of 14 to 47% of protein injected into different regions of brain or cerebrospinal fluid passes through lymph. The magnitude of the outflow to lymph is at variance with the general assumption that the absence of conventional lymphatics from the brain interrupts the afferent arm of the immune response to brain antigens. The immune response to antigens (albumin or myelin basic protein) introduced into the central nervous system (CNS) has been analysed using a rat model with normal brain barrier permeability. The micro-injection of antigen into brain or cerebrospinal fluid elicits a humoral immune response, with antibody production in cervical lymph nodes and spleen, and also affects cell-mediated immunity. Furthermore, antigen may be more immunogenic when administered into the CNS than into conventional extracerebral sites. Clearly, the afferent arm of the immune response to antigens, within the CNS, is intact. Modern studies suggest that the efferent arm is also intact with passage of activated lymphocytes into the brain. Results support a new view of CNS immunology which incorporates continuous and highly regulated communication between the brain and the immune system in both health and disease.

Animals↗
PubMed2005

Experimental study of early olfactory neuron differentiation and nerve formation using quail-chick chimeras.

For the formation of a functional olfactory system, the key processes are neuronal differentiation, including the expression of one or the other olfactory receptors, the correct formation of the nerve and organization of periphero-central connections. These processes take place during embryonic development starting from early stages. Consequently, avian embryos afford an attractive model to study these mechanisms. Taking advantage of species-specific equipment of olfactory receptors genes in different bird species, interspecific avian chimeras were set up by grafting early chick olfactory placodes in same stage quail embryos. Their analysis was performed using different complementary approaches. In situ hybridisation using probes to different chick olfactory receptor (COR) genes indicated that the choice of expression of an olfactory receptor by a neuron is independent of the environment of the olfactory placode and of interactions with the central nervous system. Futhermore, a chick olfactory receptor gene subgroup (COR3 ), absent in the host genome, was expressed by neurons from the graft. The question was then raised of the consequences of such heterospecific differentiation on axonal projections and fiber convergence. The DiI labeling of olfactory fibres in chimeras revealed anomalies in the formation of the nerve from the chick graft. In agreement with the hypothesis of olfactory receptor (OR) involvement in axonal guidance and periphero-central synapse organisation, the presence of migrating cells and axonal fibres from the graft, expressing foreign ORs and having different interactions with the host environment than the host fibres and migrating cells, might explain these anomalies.

Animals↗