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PubMed1995

Differential expression of two cell surface proteins, neuropilin and plexin, in Xenopus olfactory axon subclasses.

Immunohistochemistry by using monoclonal antibodies named A5 and B2, which specifically recognize cell surface proteins the neuropilin and the plexin, respectively, revealed that olfactory axons in Xenopus tadpoles were classified into several subgroups by virtue of the expression levels of these two cell surface molecules. The vomeronasal axons expressed the plexin but not the neuropilin. The plexin-positive and neuropilin-negative vomeronasal axons form a discrete fiber bundle, even after they joined with the principal olfactory axons. However, the principal olfactory axons were divided into at least two subclasses; the neuropilin-predominant axons which expressed high levels of the neuropilin and low levels of the plexin, and the plexin-predominant axons which expressed high levels of the plexin and low levels of the neuropilin. Within the olfactory nerve the pathways for these two principal olfactory axon subclasses were initially intermingled with each other, but were gradually segregated throughout their courses from the nose to the cerebrum. Eventually, the neuropilin-predominant and the plexin-predominant principal olfactory axon subclasses projected to specified glomeruli in topographically related regions within the main olfactory bulb. Neuroanatomical tracings of the olfactory projection also confirmed the gradual segregation of the pathways for the principal olfactory axons. These results allow us to speculate that both the neuropilin and the plexin are involved in axon interactions, and play roles in the organization of the precise patterns of the olfactory pathway and projection.

Animals↗
PubMed1994

Microtubule fragmentation and partitioning in the axon during collateral branch formation.

Axons within the brain branch principally by the formation of collaterals rather than by bifurcation of the terminal growth cone (O'Leary and Terashima, 1988). This same behavior is recapitulated in cultures of embryonic hippocampal neurons (Dotti et al., 1988), rendering them ideal for studies on the cell biological mechanisms underlying collateral branch formation. In the present study, we focused on changes in the microtubule (MT) array that occur as these axons branch. In particular, we explored the mechanism by which MT number is locally increased to accommodate the need for more MTs during collateral branch formation. Serial reconstruction analyses indicate that MT number increases by severalfold and that MT length decreases correspondingly within the parent axon in the discrete region giving rise to the branch. These observations strongly suggest that MTs within the parent axon undergo a local fragmentation in this region, and hence raise the possibility that a portion of these new MTs might be destined for transport into the branch. To address this latter issue, we used quantitative immunofluorescence to compare the proportion of newly assembled to total MT polymer in different regions of the axon. As previously reported (Brown et al., 1992), the region of the axon contiguous with the terminal growth cone is particularly rich in newly assembled polymer. In contrast, there was no distinguishable difference in the proportion of newly assembled polymer in the newly formed collateral branches compared to the shaft region of the parent axon. These results indicate that the MTs within the newly formed collateral branches are on average assembled at the same time as those within the parent axon, and thus strongly suggest that the MTs in the collateral branch were assembled in the parent axon and then translocated into the branch. We conclude on the basis of these observations that collateral branch formation requires a local fragmentation of MTs within the parent axon, followed by the partitioning of a portion of the MT fragments into the branch. These short MTs presumably then resume their movement and elongation down the collateral branch as well as down the parent axon for the steady and orderly increase of both MT arrays.

Animals↗
PubMed1994

Schwann cells induce sprouting in motor and sensory axons in the adult rat spinal cord.

Circumscribed lesions were made within either the corticospinal tract or the ascending dorsal column tracts at the upper cervical level in adult rats. The responses of the tract axons were studied by orthograde transport from injections of horseradish peroxidase or biocytin. At 2 d, the ends of the cut axons were swollen, and the lesions induced en passant varicosities in the adjacent uncut axons. Although there have been reports of retraction, we found that even after several weeks, large numbers of cut axons still persisted in the central lesion area (where there was complete tissue destruction and intense macrophage infiltration), and also in the adjacent regions of the tract. The cut ends were expanded into a variety of shapes--large, complex, bulbous, and recurved--and many had profuse local branches with or without small, terminal-type varicosities. A suspension of Schwann cells cultured from neonatal sciatic nerve was injected by a minimally traumatic air pressure microinjection technique so as to form a bolus, comparable in size to the lesions, in either the corticospinal or the ascending dorsal column tracts at the upper cervical level. Despite previous findings that corticospinal axons do not elongate into peripheral nerve grafts, we found that both corticospinal and ascending dorsal column axons sprouted in response to contact with the transplanted Schwann cells. The response to the Schwann cells was much more rapid than to the lesions. By 2 d, in both descending and ascending tracts, both the axons that had been severed at the time of injection and also the adjacent uncut axons had already given rise to the branches that (unlike the localized sprouting seen after the long-term lesions) extended for considerable distances parallel to and fasciculating with each other and with the uncut tract axons. In addition, a mass of fine, tortuous, varicose branches invaded the superficial parts of the Schwann cell grafts, where they formed aborizations with small bead-like expansions resembling presynaptic boutons; as in their normal terminal fields, the aborizations formed by the corticospinal axons were smaller and finer than those formed by the ascending axons.

Animals↗
PubMed1993

A new method for expressing axonal size: rat optic nerve analysis.

Analysis of the shape of the cross sections of adult rat optic nerve axons reveals that the majority of axons do not have a true circular shape. Therefore, determination of axonal size has to utilize methods of approximation. The method presented here utilizes three calculated parameters for expression of axonal size: (i) axonal diameter, as calculated from its area, or (ii) axonal diameter, as calculated from its perimeter, both assuming axonal shape to be a perfect circle and (iii) axonal shape factor, which represents the divergence of the axon from a perfect circular shape. The use of the calculated axonal diameter, with a correction for its shape factor, provides a normalized way of expressing axonal size.

Animals↗
PubMed1994

Changes in microtubule number and length during axon differentiation.

Hippocampal neurons in culture initially extend several minor processes that are approximately 20 microns in length. The first minor process to grow approximately 10 microns longer than the others will continue to grow rapidly and become the axon (Goslin and Banker, 1989). We sought to define changes in the microtubule (MT) array that occur during axon differentiation. In theory, axon differentiation could involve an increase in MT number, MT length, or some combination of both. To address this issue, we first serially reconstructed the entire MT array of a minor process from a cell whose axon had not yet differentiated. This minor process contained 182 MTs that ranged in length from 0.14 to 20.09 microns. The average MT length was 3.87 +/- 3.83 microns, and the total MT length was 704 microns. We then reconstructed the MT arrays of a minor process and the 56 microns axon from a cell that had undergone axon differentiation. The minor process contained 157 MTs that ranged in length from 0.24 to 17.95 microns. The average MT length was 3.91 +/- 4.84 microns, and the total MT length was 600 microns. The axon contained 1430 MTs that ranged in length from 0.05 to 40.14 microns. The average MT length was 4.02 +/- 5.28 microns, and the total MT length was 5750 microns. These data indicate that a shift occurs toward shorter as well as longer MTs, but that virtually no change in average MT length occurs during axon differentiation. Thus, elongation of existing MTs cannot account for the major expansion of the MT array that occurs as a minor process becomes an axon. In contrast, the number of MTs increases by approximately 10-fold as a minor process differentiates and grows into an axon of the length we analyzed. Based on these data, we conclude that the MT array of a minor process is substantially expanded as it differentiates into an axon, and that the principal mechanism by which this expansion occurs is the copious addition of new MTs.

Animals↗
PubMed1994

Thalamocortical axons extend along a chondroitin sulfate proteoglycan-enriched pathway coincident with the neocortical subplate and distinct from the efferent path.

The distinct axonal tracts of the mature nervous system are defined during development by sets of substrate-bound and diffusible molecular signals that promote or restrict axonal elongation. In the adult cerebral cortex, efferent and afferent axons are segregated within the white matter. To define the relationship of growing efferent and afferent axons in the developing murine cortex to chondroitin sulfate proteoglycans (CSPGs) in the pericellular and extracellular matrix, we used the fluorescent tracer Dil to determine axonal trajectories and immunolabeling to disclose the distribution of CSPGs. Axons of neurons in the preplate are the first to leave the cortex; they arise in the CSPG-rich preplate and extend obliquely across it to enter the CSPG-poor intermediate zone. Slightly later, axons of cortical plate neurons extend directly across the CSPG-rich subplate, and then turn abruptly to run in the upper intermediate zone. In contrast, once afferent axons from the thalamus reach the developing cortical wall, their intracortical trajectory is centered on the CSPG-rich subplate, above the path taken by efferent axons. Our findings demonstrate a molecular difference between the adjacent but distinct efferent and afferent pathways in developing neocortex. Early efferents cross the subplate and follow a pathway that contains very little CSPG, while afferents preferentially travel more superficially within the CSPG-rich subplate. Thus, CSPGs and associated extracellular matrix (ECM) components in the preplate/subplate do not form a barrier to axonal initiation or outgrowth in the neocortex as they may in other locations. Instead, their distribution suggests a role in defining discrete axonal pathways during early cortical development.

Animals↗
PubMed1995

Spinohypothalamic tract neurons in the cervical enlargement of rats: descending axons in the ipsilateral brain.

Spinohypothalamic tract (SHT) cells are spinal cord neurons with axons that project directly to or through the contralateral hypothalamus. Frequently, SHT axons decussate in the posterior optic chiasm, turn posteriorly and descend to unknown locations in the ipsilateral brain. We attempted to determine the course and the termination of these descending axons. Sixty neurons in the cervical enlargement of rats were antidromically activated initially from the contralateral hypothalamus and then from multiple anterior-posterior levels in the ipsilateral brain. Fifty-three (88%) were backfired with low currents at increased latencies from the ipsilateral brain. The axons of 35 neurons were surrounded with electrode penetrations from which high currents could not activate the neuron antidromically, suggesting the examined axons terminated in the surrounded areas. Seven SHT axons that were surrounded (20%) appeared to terminate in the contralateral hypothalamus, 5 (14%) in the ipsilateral hypothalamus, and 9 (26%) in the ipsilateral thalamus. Fourteen SHT axons (40%) ended in the ipsilateral midbrain mainly in the superior colliculus, cuneiform nucleus or nucleus brachium inferior colliculus. An additional 11 axons were followed even further posteriorly into the ventral pons or rostral medulla. Each of the 26 neurons that could be physiologically classified responded either preferentially or specifically to noxious mechanical stimuli. These results indicate that SHT axons course through a surprisingly long and complex path. After decussating in the hypothalamus, the axons of many SHT neurons descend into the ipsilateral posterior thalamus, midbrain, pons, or even rostral medulla. These axons may provide nociceptive information to a variety of nuclei throughout the diencephalon and brainstem bilaterally.

Animals↗
PubMed1996

Spatially restricted increase in polysialic acid enhances corticospinal axon branching related to target recognition and innervation.

The polysialic acid (PSA) modification of the neural cell adhesion molecule (NCAM) has been shown to alter the responses of developing axons to their environment. We have studied the potential role of PSA in regulating the innervation of the spinal cord by corticospinal axons, which occurs by a delayed formation of collateral branches from the parent axons. Developmental changes in the distribution of PSA were examined immuno-histochemically using light and electron microscopy. Whereas NCAM is distributed along the entire pathway of rat corticospinal axons as they grow from the cortex to the spinal cord, PSA-modified NCAM does not become evident until later. When PSA becomes evident, it is restricted to the distal segment of these axons from the caudal hindbrain through the spinal cord. The increase in PSA on corticospinal axons coincides with the time that they begin to form collateral branches in the spinal cord. This unique spatiotemporal distribution of PSA suggests its involvement in corticospinal axon branching. To test this hypothesis, PSA was selectively removed by an in vivo injection of endoneuraminidase N. This treatment did not seem to interfere with the pathfinding of corticospinal axons; however, PSA removal delayed the onset of collateral branching by corticospinal axons within the spinal cord and later diminished the magnitude of branching. These findings indicate a role for PSA in the regulation of interstitial axon branching, a crucial step in the process of target recognition and innervation by corticospinal axons.

Animals↗
PubMed1997

Stop and branch behaviors of geniculocortical axons: a time-lapse study in organotypic cocultures.

The behavior of growing thalamic axons was studied in an organotypic coculture of the lateral geniculate nucleus (LGN) with the visual cortex (VC) to reveal cellular interactions that underlie the formation of lamina-specific thalamocortical connections. The LGN explant was placed at the ventral side, pial surface, or lateral edge of the VC explant, and fluorescent dye-labeled LGN axons were observed by confocal microscopy in fixed and living tissue. The axonal projection pattern in fixed cocultures after 1 week in vitro demonstrated that, in all three configurations, LGN axons formed primitive branches mainly in layer 4. A time-lapse study further examined axonal growth and branch formation in the living cortical explant. The majority of branches emerged within layer 4 behind the axonal tip, regardless of the direction of axonal entry. In addition, most axons entering from the ventral or pial side of the VC exhibited a transient or persistent stop of axonal growth in and around layer 4, whereas those entering from the lateral edge of the VC traveled along layer 4 without exhibiting stop behavior. The axonal stop often was accompanied by growth cone collapse and a slight retraction. These results suggest the existence of branch and stop cues in layer 4 of the cortex that are recognized by LGN axons.

Animals↗
PubMed1997

Conduction in segmentally demyelinated mammalian central axons.

The prominent symptoms associated with central demyelinating diseases such as multiple sclerosis (MS) are primarily caused by conduction deficits in affected axons. The symptoms may go into remission, but the mechanisms underlying remissions are uncertain. One factor that could be important is the restoration of conduction to affected axons, but it is not known whether demyelinated central axons resemble their peripheral counterparts in being able to conduct in the absence of repair by remyelination. In the present study we have made intra-axonal recordings from central axons affected by a demyelinating lesion, and then the axons have been labeled ionophoretically to permit their subsequent identification. Ultrastructural examination of 23 labeled preparations has established that some segmentally demyelinated central axons can conduct, and that they can do so over continuous lengths of demyelination exceeding several internodes (2500 micron). Such segmentally demyelinated central axons were found to conduct with the anticipated reduction in velocity and a refractory period of transmission (RPT) as much as 34 times the value obtained from the nondemyelinated portion of the same axon; the RPT was typically prolonged to 2-5 times the normal value. We conclude that some segmentally demyelinated central axons can conduct, and we propose that the restoration of conduction to such axons is likely to contribute to the remissions commonly observed in diseases such as MS.

Action Potentials↗
PubMed1998

Postnatal development of olfactory receptor cell axonal arbors.

The mechanisms that subserve the distribution of the terminal arbors of olfactory receptor cell axons remain unknown. Elsewhere in the central nervous system, a common theme is early axonal exuberance followed by activity-dependent pruning to achieve the mature distribution. This led to the hypothesis that the orderly morphology of afferent axons in the olfactory glomerulus may follow a similar developmental scheme of exuberance followed by pruning. To test this hypothesis, we studied morphological features of olfactory receptor neuron (ORN) axonal arbors on postnatal days 0, 3, 6, 9, 12, and 21. The olfactory bulbs from Sprague-Dawley rats were processed using a Golgi technique that impregnated ORN axons. Axons from each age group were reconstructed by using camera lucida at x100, oil immersion, and morphometrically characterized. In the adult, the percent glomerular area occupied by a single ORN axon was 14%, whereas the mean length of branches was 169.67 microm, the sum of branches and varicosities was 27, and the distance to first branch point in glomeruli was 21.98 microm. The values from the younger age groups were not statistically different from those in the adult. Because there was no evidence of early exuberance, our data suggest that ORN axons must innervate single glomeruli and arborize in a highly specific manner to achieve the adult pattern. Because our data suggest that ORN axons do not follow the hypothesized scheme, it is plausible to suggest that as ORN axons innervate a glomerulus during development they arborize to their adult levels but not beyond. This argues strongly that specific cell surface and trophic factors are used by the ORN axon to guide glomerular targeting and innervation.

Animals↗
PubMed1997

[Immunohistochemical methods in the diagnosis of diffuse axonal injury].

Diffuse axonal injury (DAI) was diagnoses according to axonal reaction proved as balls by Palmgren's silver impregnation and axonal lesions by immunohistochemical investigation of ubiquitin and low molecular weight neurofilaments (68 kD). Diffuse axonal injury was found in 16 cases from a group of 36 persons deceased of craniocerebral trauma (44%). Diagnosis was based on the presence of axonal retraction balls in 15 cases; the balls showed also a striking positivity with antibodies against ubiquitin and low molecular neurofilaments as well. In the last patient who died 10 hours after a head injury, too early for axonal retraction balls to be formed, diagnosis of DAI could be settled because of the presence of axonal swellings immunohistochemically positive for ubiquitin and neurofilaments. A control groups of 6 persons after sudden death showed both axonal widenings and swellings but their immunohistochemistry was negative. Immunohistochemical methods of proving ubiquitin and low molecular weight neurofilaments represent substantial contribution to diagnosis of diffuse axonal injury. Earlier phase of axonal lesions than in simple histology can be observed and histological diagnosis of axonal injury independently confirmed.

Adolescent↗
PubMed1998

Carbon disulfide neurotoxicity in rats: V. Morphology of axonal swelling in the muscular branch of the posterior tibial nerve and spinal cord.

The study objectives were to examine the morphological progression and dose response of carbon disulfide (CS2) distal axonopathy in the muscular branch of the posterior tibial nerve (MBPTN) and spinal cord. Male and female F344 rats were exposed to 0, 50, 500 or 800 ppm CS2 by inhalation, 6 hours/day, 5 days per week, for 2, 4, 8 or 13 weeks. At 8 weeks, in the MBPTN, single fascicles contained individual swollen axons. By 13 weeks, multiple fascicles had giant swollen axons with thin myelin sheaths and occasional degenerated and regenerated axons. At 8 weeks, in the spinal cord, white matter changes in cervical segments 1 and 2 consisted of prominent multifocal axonal swelling in the fasciculus gracilis nerve tracts. In lumbar segments 1 and 2, multifocal axonal swelling was first present at 8 weeks in the lateral and ventro-medial funiculus. By 13 weeks, axonal swelling was diffuse in the fasciculus gracilis nerve tracts of the cervical spinal cord and the lateral and ventral funiculus nerve tracts in the lumbar spinal cord. Compared to the spinal cord, where axonal swelling was present in rats exposed to 800 and 500 ppm, in the muscular branch of the posterior tibial nerve, axonal swelling was only present at 800 ppm at both 8 and 13 weeks. Electron microscopic examination demonstrated marked accumulations of neurofilaments in swollen axons in the spinal cord and MBPTN. Axonal swelling was not present in the spinal cord at 50 ppm, or in the MBPT at 50 and 500 ppm. Axonal swelling was not present at earlier time points of 2 and 4 weeks in either the spinal cord or MBPTN.

Administration, Inhalation↗
PubMed1998

Axon projections of reciprocal inhibitory interneurons in the spinal cord of young Xenopus tadpoles and implications for the pattern of inhibition during swimming and struggling.

We have examined the morphology and longitudinal axon projections of a population of spinal commissural interneurons in young Xenopus tadpoles. We aimed to define how the distribution of axons of the whole population constrains the longitudinal distribution of the inhibition they mediate. Forty-three neurons at different positions were filled intracellularly with biocytin and processed with avidin-conjugated horseradish peroxidase. Soma size did not vary longitudinally and only one ipsilateral axon was found. Contralateral axons ascended, descended, or usually branched to do both. Total axon length and the extent of dendritic arborisation decreased caudally. The distributions of ascending and descending axon lengths were different; there were more long ascending (mean 737 +/- standard deviation 365 microm) than long descending (447 +/- 431 microm) axons. We used the axon length distribution data with existing data on the distribution of commissural interneuron somata to calculate the overall longitudinal density of these inhibitory axons. Axon numbers showed a clear rostrocaudal gradient. Axon length distributions were then incorporated into a simple spatiotemporal model of the forms of inhibition during swimming and struggling motor patterns. The model predicts that the peak of inhibition on each cycle will decrease from head to tail in both motor patterns, a feature already confirmed physiologically for swimming. It also supports a previous proposal that ascending inhibition during struggling shortens cycle period by shortening rostral motor bursts, whereas descending inhibition could delay subsequent burst onset.

Animals↗
PubMed1998

Acute inactivation of tau has no effect on dynamics of microtubules in growing axons of cultured sympathetic neurons.

Tau is a developmentally regulated microtubule (MT)-associated protein in neurons that has been implicated in neuronal morphogenesis. On the basis of test tube studies, tau has been proposed to function in axon growth by stabilizing MTs and thereby promoting MT assembly. We have tested this hypothesis by examining the effects of acute inactivation of tau on axonal MTs. Tau was inactivated by microinjecting purified antibodies against recombinant tau into neurons before they extended axons. The injected antibodies quantitatively precipitated tau into aggregates in the soma. With these conditions the neurons elaborate normal-appearing axons, and MTs extend throughout the axons and into the growth cones, but the axons and their MTs are depleted of tau. The immunodepletion of tau had no detectable effect on several parameters of the dynamics of axonal MTs. Depletion of tau also was not accompanied by a reorganization of other major MT-associated proteins or actin filaments in these neurons. Thus, neurons effectively depleted of tau can extend axons that resemble those of control cells, and the axons contain normal-appearing MT arrays with normal dynamic behavior. These observations are exactly the opposite of those expected on the basis of the hypothesis that the stability of axonal MTs is a direct function of their content of tau, indicating that tau in growing axons of cultured sympathetic neurons is not specialized to promote microtubule assembly and stability.

Adrenergic Fibers↗
PubMed2001

Thalamic terminal morphology and distribution of single corticothalamic axons originating from layers 5 and 6 of the cat motor cortex.

We investigated the axonal morphology of single corticothalamic (CT) neurons of the motor cortex (Mx) in the cat thalamus, using a neuronal tracer, biotinylated dextran amine (BDA). After localized injection of BDA into the Mx, labeled CT axons were found ipsilaterally in the thalamic reticular nucleus (TRN), the ventroanterior-ventrolateral complex (VA-VL), the central lateral nucleus (CL), the central medial nucleus, and the centromedian nucleus, but with the primary focus in the VA-VL. The terminals in the VA-VL formed a large laminar cluster, which extended approximately in parallel with the internal medullary lamina. The laminar organization mirrored morphologic features of single CT axons. We reconstructed the trajectories of 25 single CT axons that arose from layer V (16 axons) or layer VI (9 axons) and terminated in the VA-VL. Terminals of single CT axons that originated from both layer V and layer VI were confined within a laminar structure about 700 microm thick, suggesting the existence of laminar input organization in the VA-VL. Otherwise, the two groups of the CT axons showed contrasting features. All of the CT axons derived from layer VI gave rise to a few short collaterals to the TRN and then formed extensive arborization with numerous small, drumstick-like terminals in the VA-VL. On the other hand, the CT axons arising from layer V gave rise to collaterals whose main axons descended into the cerebral peduncle. Each collateral projected to the VA-VL or CL without projection to the TRN and formed a few small clusters of giant terminals. The two groups of CT neurons in the same cortical column had convergent rather than segregated termination in the VA-VL. However, the terminals of layer VI CT neurons were distributed diffusely and widely in the VA-VL, whereas the terminals of layer V CT neurons were much more focused and surrounded by the terminals of the former group. These contrasting features of the two types of CT projections appear to represent their different functional roles in the generation of motor commands and control of movements in the Mx.

Animals↗
PubMed1986

Mesencephalic trigeminal sensory neurons of cat: axon pathways and structure of mechanoreceptive endings in periodontal ligament.

We injected 3H-proline into cat brainstem in order to label the entire mesencephalic trigeminal nucleus (Mes-V) for autoradiographic analysis of the size and pathways of Mes-V sensory axons and for microscopic study of Mes-V receptor structure in dental tissue. Labeled sensory axons were found in the trigeminal motor and sensory tracts and roots; approximately equal numbers of axons were found in both roots. The sensory root and all three divisions of the trigeminal nerve contained larger Mes-V axons than the motor root. Labeled Mes-V axons were found at the ganglion in the dorsomedial (infratrochlear) branch of the ophthalmic nerve but not in the ventrolateral branch. The mean diameter of Mes-V axons in periodontal ligament was 4.0 +/- 1.9 micron compared to 7.3 +/- 2.1 micron in maxillary and mandibular nerve, suggesting axonal arborization prior to innervation of ligament. Mes-V receptors in dental tissue were confined to ipsilateral periodontal ligament close to the root apex, with greater innervation on the posterior side. Receptor incidence was moderate for most teeth; however, maxillary first and second incisors and maxillary and mandibular canines had focal areas with remarkably dense innervation. No labeled axons were found in pulp of any ipsilateral teeth, and none was found in any contralateral dental tissue. EM-autoradiography demonstrated that Mes-V axons form unencapsulated Ruffini-like mechanoreceptors in periodontal ligament. The preterminal axons were small and myelinated. Neighboring bundles of unmyelinated axons and rare encapsulated endings were not labeled. The labeled mechanoreceptors branched to varying degrees among the ligament fibers; they contained numerous mitochondria and glycogen particles, as well as some vesicles and rare multivesicular bodies. They were surrounded by special Schwann cells that formed one or several layers around the ending. The endings were exposed to the basal lamina at numerous sites and occasionally extended fingers beyond the lamellar Schwann cells to contact ligament collagen.

Animals↗
PubMed1987

Axon trajectories and pattern of terminal arborization during the prenatal development of the cat's retinogeniculate pathway.

In this study we have examined the trajectories taken by populations of ganglion cell axons and the spatial gradients of terminal arbor maturity within the lateral geniculate nucleus (LGN) during the prenatal development of the cat's visual system. To do so, an in vitro method of labeling optic tract axons from fetal brains between embryonic day 37 (E37) and postnatal day 2 (P2) with horseradish peroxidase (HRP) was used. At the earliest ages studied (E37-E53), optic axons leave the optic tract to run across the LGN toward their sites of termination in straight trajectories parallel to each other. At later ages (E57-P2), however, axons with abrupt changes in their course across the nucleus can be clearly identified. When the detailed terminal arbor morphology of the set of retinogeniculate axons filled with HRP at a given age was examined, two different spatial gradients of maturation could be detected. The terminal arbors of axons within LGN layer A are always more mature than those ending in layer A1, an observation consistent with previous findings that axons from the contralateral eye arrive within the LGN several days before those from the ipsilateral eye. Moreover, the terminal arbors of axons projecting to the medial portions of each layer are always more mature than their more lateral counterparts. These gradients are likely to be a direct reflection of the central-first, peripheral-last gradient associated with the neurogenesis of the retinal ganglion cells themselves. In the oldest animals studied (E58-P2), a remarkable periodic pattern of terminal arbor labeling was seen following a localized HRP injection into the optic tract. Within the labeled portions of the LGN, densely filled axon terminal arbors are separated by unlabeled gaps of similar width. This pattern of labeling could reflect local topographic disorder within the optic tract or could arise if axons of different classes of retinal ganglion cells run in separate portions of the optic tract. Taken together, all of these observations suggest that there may be a fair degree of topographic order in the retinogeniculate projection within the cat's LGN early on in development. However, when topographic errors are present, some can be corrected by minor readjustments in axonal trajectories.

Animals↗