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PubMed1979

The guidance of optic axons in the developing and adult mouse retina.

Previous studies in chick embryos (Goldberg, '77) indicated that unidirectional guidance of retinal axons toward the optic nerve is restricted to the vitread portion of the ganglion cell fiber layer (GCFL) of the retina; random fiber growth was noted after deflection of the optic axons sclerad to the GCFL. The present study on mice confirms these observations. Silver-stained flat mounts of retinal colobomas were examined. Many optic axons in colobomas do not exit normally from the eye, but travel randomly when deflected sclerad to the GCFL. Newborn mouse axons grew around retinal lesions in a highly directed manner. Such axons were always situated in the vitread portion of the GCFL. The unidirectional guidance found in newborn mice was absent in adults. Deflected adult axons traveled randomly regardless of their level within the GCFL. We propose that defective guidance largely accounts for failure of axonal regeneration in the adult mouse retina. The inability of the adult axons to fasciculate (adhere to one another and form fiber bundles) suggests that impaired cellular adhesivity may be part of the mechanism of regenerative failure.

Animals↗
PubMed1984

The radioscapholunate ligament: a gross and histologic description.

The anatomic relationships of the carpal radioscapholunate ligament to its contiguous structures were analyzed by studying 1) 12 grossly dissected fresh adult wrists, and 2) multiple histologic sections from six adult wrists. Observations indicate that the radioscapholunate ligament originates from the prominence between the scaphoid and lunate articular facets on the distal articular surface of the radius, and from the palmar margin of the distal radius, deep and medial to the origin of the radiotriquetral and radiocapitate ligaments. The primary insertion of the radioscapholunate ligament is the medial margin of the proximal pole of the scaphoid. The ligament secondarily inserts into the lateral margin of the lunate and significantly contributes to the proximal portion of the scapholunate interosseous ligament. The radioscapholunate ligament is distinguished morphologically from the other palmar radiocarpal ligaments by its loosely organized collagen fibers and relatively high degree of vascularity. The radiotriquetral and radiocapitate ligaments are composed of densely fasciculated collagen fibers surrounded by perpendicularly oriented perifascicular and epiligamentous fibers. A fibrous capsular layer covers the most superficial aspect of each carpal ligament. On the deep surfaces of these ligaments, a condensation of epiligamentous fibers forms a synovial capsular layer. The palmar radiocarpal ligaments are truly intracapsular structures, as they are interposed between the fibrous and synovial capsular layers. No histologic evidence of elastin is present within the substance of these ligaments.

Aged↗
PubMed2006

GAP-43 heterozygous mice show delayed barrel patterning, differentiation of radial glia, and downregulation of GAP-43.

GAP-43 heterozygous (HZ) mice exhibit abnormal thalamocortical pathfinding, fasciculation, and terminal arborization at postnatal day 7 (P7). Here we tested whether these defects are correlated with delayed development of HZ cortical patterns. We assessed the rate of barrel segregation and radial glia differentiation in wild-type (WT) and HZ cortices. Since GAP-43 is involved in some forms of neural plasticity, we also compared the duration of the critical period for lesion-induced plasticity in both genotypes. Cytochrome oxidase histochemistry revealed a delay of approximately 1 day in barrel pattern formation in GAP-43 HZ mice. GAP-43 WT barrels showed complete segregation between P2-P3, while HZ barrels did not reach the same level of segregation until P3-P4. We found a similar delay in the transformation of radial glia from monopolar to multipolar phenotypes, from P5 in WT to P7 in HZ cortex. Radial glial cells represent many of the neuronal progenitors in developing cortex and aid in cell migration. Thus, the delay in radial glial differentiation may contribute to the delay in HZ barrel segregation. Interestingly, we found no change in the extent of the critical period for HZ cortical responsiveness to early peripheral damage or in the time course of the cortical response. As expected, GAP-43 expression in HZ cortex is significantly reduced early in development. However, HZ GAP-43 expression remains at maximum levels after P9, when it is normally downregulated. As a result, HZ GAP-43 expression is near-normal by P26, by which time near-normal barrel dimensions have been restored. Our findings indicate that GAP-43 deficiency leads to early delays in barrel development and suggest that these failures are followed by homeostatic responses, including prolonged GAP-43 expression. These compensatory mechanisms may rescue normal cortical reorganization in neonates and near-normal barrel morphology and GAP-43 expression in adulthood.

Animals↗
PubMed2003

Median nerve penetration by a muscular branch of the brachial artery.

Arterial variations in the distal half of the arm are common (Anagnostopoulou and Venieratos 1999, J. Anat. 195:471-476). We observed a rare variant of a muscular branch of the brachial artery that penetrated the median nerve in the lower part of the arm. After its origin from the posteromedial surface of the brachial artery, the muscular branch passed through a nervous loop in the median nerve to enter and supply the brachialis muscle. At the site of arterial penetration, the nerve displayed perineurial thickening and increased fasciculation and interfascicular connective tissue. Three small branches of the artery supplied the nerve fascicles. We did not observe nerve compression or muscular changes, but the possibility of altered clinical symptoms produced by irritation from arterial pulsation cannot be ruled out. This variation may be clinically important because symptoms of median nerve compression arising from similar variations are often confused with more common causes, such as radiculopathy and carpal tunnel syndrome.

Arm↗
PubMed2002

Expression of specific glycoconjugates in both primary and secondary olfactory pathways in BALB/C mice.

Binding of cell surface carbohydrates to their receptors specifically promotes axon growth and synaptogenesis in select regions of the developing nervous system. In some cases these interactions depend upon cell-cell adhesion mediated by the same glycoconjugates present on the surface of apposing cells or their processes. We have previously shown that the plant lectin Dolichos biflorus agglutinin (DBA) binds to a subpopulation of mouse primary olfactory neurons whose axons selectively fasciculate prior to terminating in the olfactory bulb. In the present study, we investigated whether these glycoconjugates were also expressed by postsynaptic olfactory neurons specifically within the olfactory pathway. We show here for the first time that DBA ligands were expressed both by a subset of primary olfactory neurons as well as by the postsynaptic mitral/tufted cells in BALB/C mice. These glycoconjugates were first detected on mitral/tufted cell axons during the early postnatal period, at a time when there is considerable synaptogenesis and synaptic remodelling in the primary olfactory cortex. This is one of the few examples of the selective expression of molecules in contiguous axon tracts in the mammalian nervous system. These results suggest that glycoconjugates recognized by DBA may have a specific role in the formation and maintenance of neural connections within a select functional pathway in the brain.

Animals↗
PubMed2003

Developmental relationship between cholinergic amacrine cell processes and ganglion cell dendrites of the mouse retina.

Ganglion cells of the mammalian retina undergo structural remodeling before their dendrites are confined to functionally distinct laminas within the inner plexiform layer. It has been proposed that cholinergic amacrine cells provide laminar cues that remodel ganglion cell dendrites, because their processes stratify before those of the ganglion cells. To address this possibility, it is necessary to know whether cholinergic cells contact all or only some classes of ganglion cells during development. We, therefore, used two-photon microscopy to simultaneously reconstruct the dendritic arbors of different classes of ganglion cells and terminal processes of cholinergic cells in neonatal mouse retina. We determined that, after birth, cholinergic cells contacted only a subset of ganglion cells. Large bistratified cells (LBCs), resembling direction selective ganglion cells in other species, had dendrites that fasciculated with the cholinergic plexuses. The LBCs received numerous presynaptic cholinergic contacts shortly after birth. In contrast, large monostratified cells (LMCs), ramifying outside the cholinergic plexuses at maturity, received few, if any, cholinergic contacts even at early stages when their dendrites overlapped with the cholinergic processes. These observations suggest that cholinergic cells provide laminar cues for only subsets of ganglion cells. They also indicate that the synaptic organization between amacrine and ganglion cells may be specified early in development.

Acetylcholine↗
PubMed2003

Abnormal thalamocortical pathfinding and terminal arbors lead to enlarged barrels in neonatal GAP-43 heterozygous mice.

GAP-43 has been implicated in axonal pathfinding and sprouting, synaptic plasticity, and neurotransmitter release. However, its effect on cortical development in vivo is poorly understood. We have previously shown that GAP-43 knockout (-/-) mice fail to develop whisker-related barrels or an ordered whisker map in the cortex. Here we used cytochrome oxidase (CO) histochemistry to demonstrate that GAP-43 heterozygous (+/-) mice develop larger than normal barrels at postnatal day 7 (P7), despite normal body and brain weight. Using serotonin transporter (5HT-T) histochemistry to label thalamocortical afferents (TCAs), we found no obvious abnormalities in other somatosensory areas or primary visual cortex of GAP-43 (+/-) mice. However, TCA projections to (+/-) primary auditory cortex were not as clearly defined. To clarify the mechanism underlying the large-barrel phenotype, we used lipophilic (DiI) axon labeling. We found evidence for multiple pathfinding abnormalities among GAP-43 (+/-) TCAs. These axons show increased fasciculation within the internal capsule, as well as abnormal turning and branching in the subcortical white matter. These pathfinding errors most likely reflect failures of signal recognition and/or transduction by ingrowing TCAs. In addition, many DiI-labeled (+/-) TCAs exhibit widespread, sparsely branched terminal arbors in layer IV, reflecting the large-barrel phenotype. They also resemble those found in rat barrel cortex deprived of whisker inputs from birth, suggesting a failure of activity-dependent synaptogenesis and/or synaptic stabilization in (+/-) cortex. Our findings suggest that reduced GAP-43 expression can alter the fine-tuning of a cortical map through a combination of pathfinding and synaptic plasticity mechanisms.

Animals↗
PubMed2003

Innervation of the heart of the adult fruit fly, Drosophila melanogaster.

The innervation of the adult abdominal heart of Drosophila melanogaster was studied by neuronal staining with green fluorescent protein and immunocytochemical techniques. The investigation was undertaken to determine whether the adult heart receives neuronal input or whether its complex activity must be considered independent from the nervous system. The larval heart lacks innervation, suggesting that the cardiac impulse is totally myogenic. At metamorphosis, segmental neural processes grow onto the myocardium. A pair of transverse nerves innervates bilaterally each cardiac chamber and its alary muscles. These nerve terminals are immunoreactive to glutamate and form unique synaptic structures on the ventral layer of longitudinal cardiac muscles of the conical chamber. This characteristic cardiac synapse may represent part of the neural mechanism controlling the retrograde heartbeat, and, thus, the cardiac reversal that is characteristic of adults. In addition, crustacean cardioactive peptide-immunoreactive fibers originating from peripheral, bipolar neurons (BpNs) fasciculate with the transverse nerve projections and terminate segmentally throughout the abdominal heart. An additional cluster composed of four large, CCAP-positive neurons innervates the terminal chamber. The cardioacceleratory effect of CCAP release at this location may modulate the properties of a pacemaker producing the anterograde heartbeat.

Animals↗
PubMed2003

Expression of L1 decreases during postnatal development of rat spinal cord.

L1 is a cell adhesion molecule that is highly expressed on developing axons and is associated with neurite outgrowth, guidance, and fasciculation. In this study we systematically examined L1 expression at all spinal levels across eight postnatal ages to detect regional and developmental differences. We observed striking changes in the developmental pattern of L1 expression between birth (P0) and adult ages, with intense L1-immunopositive axons prevalent throughout the funiculi at P0 compared with predominantly L1-immunonegative funicular axons in adults. At all ages and spinal levels examined, some L1-positive dorsal root afferents entered the spinal cord, coursed in Lissauer's tract, and projected into the superficial dorsal horn and the dorsal columns, as well as across the dorsal commissure. Additional L1-positive axons were detected consistently around the perimeter of the spinal cord, in the dorsolateral funiculus, and adjacent to the central canal. While specific L1-labeled axons were detected at all ages, a pattern of segmental variation was observed within animals, with the highest levels of L1 expression detected in lumbar and sacral segments and the lowest in cervical spinal cord. The pattern of L1 immunoreactivity was compared to that of the growth-associated protein GAP-43 and the results indicated colabeling of most axons. These observations demonstrate that L1 is expressed on immature axons well into postnatal development, possibly until they have completed their differentiation. Furthermore, the L1-positive axons that continue to be detected in adults are likely to be either unmyelinated or sprouting axons.

Animals↗
PubMed2001

A role for cingulate pioneering axons in the development of the corpus callosum.

In many vertebrate and invertebrate systems, pioneering axons play a crucial role in establishing large axon tracts. Previous studies have addressed whether the first axons to cross the midline to from the corpus callosum arise from neurons in either the cingulate cortex (Koester and O'Leary [1994] J. Neurosci. 11:6608-6620) or the rostrolateral neocortex (Ozaki and Wahlsten [1998] J. Comp. Neurol. 400:197-206). However, these studies have not provided a consensus on which populations pioneer the corpus callosum. We have found that neurons within the cingulate cortex project axons that cross the midline and enter the contralateral hemisphere at E15.5. By using different carbocyanine dyes injected into either the cingulate cortex or the neocortex of the same brain, we found that cingulate axons crossed the midline before neocortical axons and projected into the contralateral cortex. Furthermore, the first neocortical axons to reach the midline crossed within the tract formed by these cingulate callosal axons, and appeared to fasciculate with them as they crossed the midline. These data indicate that axons from the cingulate cortex might pioneer a pathway for later arriving neocortical axons that form the corpus callosum. We also found that a small number of cingulate axons project to the septum as well as to the ipsilateral hippocampus via the fornix. In addition, we found that neurons in the cingulate cortex projected laterally to the rostrolateral neocortex at least 1 day before the neocortical axons reach the midline. Because the rostrolateral neocortex is the first neocortical region to develop, it sends the first neocortical axons to the midline to form the corpus callosum. We postulate that, together, both laterally and medially projecting cingulate axons may pioneer a path for the medially directed neocortical axons, thus helping to guide these axons toward and across the midline during the formation of the corpus callosum.

Age Factors↗
PubMed2001

Identification and characterization of a cell surface marker for embryonic rat spinal accessory motor neurons.

The developing mammalian spinal cord contains distinct populations of motor neurons that can be distinguished by their cell body positions, by the expression of specific combinations of regulatory genes, and by the paths that their axons take to exit the central nervous system (CNS). Subclasses of spinal motor neurons are also thought to express specific cell surface proteins that function as receptors which control the guidance of their axons. We identified monoclonal antibody (mAb) SAC1 in a screen aimed at generating markers for specific subsets of neurons/axons in the developing rat spinal cord. During early embryogenesis, mAb SAC1 selectively labels a small subset of Isl1-positive motor neurons located exclusively within cervical segments of the spinal cord. Strikingly, these neurons extend mAb SAC1-positive axons along a dorsally directed trajectory toward the lateral exit points. Consistent with the finding that mAb SAC1 also labels spinal accessory nerves, these observations identify mAb SAC1 as a specific marker of spinal accessory motor neurons/axons. During later stages of embryogenesis, mAb SAC1 is transiently expressed on both dorsally and ventrally projecting spinal motor neurons/axons. Interestingly, mAb SAC1 also labels the notochord and floor plate during most stages of spinal cord development. The mAb SAC1 antigen is a 100-kD glycoprotein that is likely to be the rat homolog of SC1/BEN/DM-GRASP, a homophilic adhesion molecule that mediates axon outgrowth and fasciculation.

Accessory Nerve↗
PubMed2005

L1 CAM expression in the superficial dorsal horn is derived from the dorsal root ganglion.

The cell adhesion molecule L1 is highly expressed on embryonic axons and may play a role in axonal outgrowth and fasciculation. Generally only low levels of L1 are found in adult spinal cord except for intense labeling in Lissauer's tract, in laminae I-II, and on dorsolateral funicular axons. In this study we determine the source of L1 immunoreactivity in the dorsal spinal cord, the presence of L1 expression on sprouting axons, and the effect of exercise on L1 expression. We determined the source of L1 immunoreactivity in the superficial dorsal horn by performing acute unilateral rhizotomies (T12-L4) in adult rats. This resulted in a marked decrease in L1 expression in Lissauer's tract and laminae I-II on the deafferented side. The peptidergic and nonpeptidergic small-diameter primary afferent markers, calcitonin gene-related peptide (CGRP) and the lectin IB4 respectively, closely correlated with L1 expression and also decreased dramatically after rhizotomy. Considering its developmental role, we asked whether L1 was expressed on sprouting axons following chronic rhizotomy. L1 and CGRP, but not IB4, were detected on sprouting axons. Lastly, we investigated the effect of exercise on L1 expression by giving animals with chronic rhizotomies free access to an exercise wheel. After extensive exercise, L1, CGRP, and IB4 expression levels were unchanged compared with those of sedentary chronic animals. Combined, these data demonstrate that the dorsal root ganglia is a major source of L1-positive axons in the superficial dorsal horn, that both L1 and CGRP identify sprouting axons following rhizotomy, and that exercise does not upregulate L1 expression.

Animals↗
PubMed2006

Septal organ of Grüneberg is part of the olfactory system.

The olfactory system in rodents and many other mammals is classically divided into two anatomically separate, and morphologically distinct, sensory systems: the main olfactory system and the accessory olfactory system. We have now identified a novel third population of olfactory marker protein-expressing sensory neurons that is located in a discrete pocket of the rostral nasal septum, which we refer to as the septal organ of Grüneberg (SOG). Neurons in this region of the septum are located in the submucosa, in small grape-like clusters, rather than in a pseudostratified neuroepithelium, as seen in both the olfactory and vomeronasal neuroepithelia. Despite their unusual location, axons projecting from the SOG neurons fasciculate into several discrete bundles and terminate in a subset of main olfactory bulb glomeruli. These glomeruli most likely represent a subset of atypical glomeruli that are spatially restricted to the caudal main olfactory bulb. The unique rostral position of the SOG suggests that the SOG may be functionally specialized for the early detection of biologically relevant odorants.

Animals↗
PubMed1977

An autoradiographic study of the development of the entorhinal and commissural afferents to the dentate gyrus of the rat.

The development of the entorhinal, ipsilateral associational, and commissural afferents to the dentate gyrus have been studied autoradiographically, following the injection of small amounts of tritiated proline into the medial and lateral parts of the entorhinal cortex, and into fields CA3c and CA4 of the hippocampus, in a series of rats, on the third, sixth, and twelfth postnatal days. Clear labeling of the entorhinal afferents were found at the third postnatal day, and from the earliest stage studied the afferents from the two parts of the entorhinal cortex appear to be spatially segregated within the stratum moleculare of the dentate gyrus: the fibers from the lateral entorhinal area occupying the outermost one-third, or so, of this stratum, while those from the medial entorhinal cortex occupy its middle zone. The ipsilateral hippocampo-dentate associational pathway is present at the third postnatal day, but the commissural projection (which shares with it the inner part of the stratum moleculare) could not be labeled until the sixth postnatal day. By the twelfth day the characteristic adult pattern of distribution of the terminals of the two hippocampo-dentate pathways is established. Although this pattern is best accounted for on the basis of a temporal competition for the available synaptic sites on the proximal parts of the dendrites of the granule cells, the spatial segregation of these two fiber systems from those arising in the entorhinal cortex, is probably due to the selective fasciculation of fibers in each group of afferents and to their early cytochemical specificity.

Afferent Pathways↗
PubMed1978

The cochlear nerve in the cat: topography, cochleotopy, and fiber spectrum.

The topographical and cytological features of the cochlear nerve in normal adult cats were studied by microdissection, light microscopy, transmission and scanning electron microscopy. The 2-mm long cochlear nerve trunk is situated within the internal acoustic meatus. The nerve is wrapped like a roll with the more basal fibers situated peripheral to the more apical ones. The fibers are fasciculated according to their target in the cochlea and follow a helical course with the sharpest spiralling in the distal portion of the fiber. Apical and basal coil fibers are of approximately the same length measured from the spiral lamina to the rostral border of the anteroventral cochlear nucleus. The nerve contains about 90% thick, myelinated fibers, about 10% unmyelinated axons, and a few thin, but heavily myelinated axons. The first group shows a unimodal distribution of axonal diameters, with a gradual increase in average diameter from basal to apical coil fibers. The number of axonal microtubules and neurofilaments have a high linear correlation to axonal circumference without principal cochleotopic differences. The myelin sheaths are thicker and the internodal segments shorter than expected from similarly sized peripheral fibers in rodents. The possible nature and origin of the three fiber categories are discussed.

Animals↗
PubMed1979

The environment of axonal migration in the developing chick retina: a scanning electron microscopic (SEM) study.

We have made a SEM study of the basal intercellular spaces of the retina in chick embryos of different developmental stages. Since this is the environment where optic axons grow, the structural characteristics of this region might play some role in the orientation of axonal migration towards the choroid fissure. The basal region of undifferentiated retinas is formed by the vitreal expansions of neuroepithelial cells. In pre-axonal stages, the intercellular spaces between these expansions do not show any preferential orientation towards the fissure. The growth cones of ganglion cell axons appear in an apicobasal direction and turn towards the fissure immediately beneath the vitreal surface. Fasciculation is an early event during development and, in the more advanced stages, the vitreal expansions from retinal cells are placed in rows following the same orientation as the axon bundles. These observations are discussed in relationship to current hypotheses on axonal migration and orientation.

Animals↗
PubMed1980

Ontogeny of the retina and optic nerve in Xenopus laevis. II. Ontogeny of the optic fiber pattern in the retina.

The ontogeny of optic fiber patterns within the Xenopus laevis retina has been studied in silver-stained serial sections and whole mounts. Ganglion cell axons ("pioneers") appear much earlier than previously recognized, at Nieuwkoop-Faber ('56) Stage 28. Fibers fasciculate into bundles as they arise in the retina and reach the chiasma by NF Stage 32. Because the retina grows in rings at its margins, ganglion cell axons differentiate at the periphery and join the nearest fiber bundles (fascicles) to reach the optical disc. A radial fascicle pattern develops by Stage X (Grant et al., '80) within the optic fiber layer of the retina and acts as a template of pathways that lead all later appearing fibers out of the retina. Since the retina grows asymmetrically along its dorso-ventral axis, the fascicle pattern develops asymmetrically, with dorsal quadrant fascicles forming an arcuate pattern in contrast to shorter, thicker ventral quadrant fascicles. The growth pattern of the retina and the "principle" of fiber following suggest that "positional information" is acquired by ganglion cells according to a system of polar coordinates (MacDonald, '77). A radial distance "r" from the center is determined temporarily by ganglion cell birthday and an angular value "theta" is given by the angular position of the fascicle followed by its axons to the optical disc. Wedges of ganglion cells send their axons into a common fascicle which retains its integrity as it leaves the retina at the optic disc. This pattern of optic fiber development suggests a model for retino-topic ordering of the optic fiber projection in Xenopus, with the fascicle acting as the "pattern-forming unit of connectivity."

Animals↗
PubMed1980

Pattern of development of ascending and descending fibers in embryonic spinal cord of chick: I. Role of position information.

Previous studies on the rigid time and position pattern of embryonic spinal cord development are extended to include the pattern of growth of axonal processes. This study was designed to analyze the position of axons in the marginal zone as they course (fasciculate) in the ascending and descending direction from a single spinal segment. Injections of 3H-proline were made into L1 or T6 of chick embryos of stages 21--29 (3.5--6 days of incubation). The location of radioactive label in the marginal zone was analyzed autoradiographically to show the location of labeled axons originating from the injected segment. In the early stages of development (stage 21), the labeled descending axons were located in the most ventromedial aspect of the marginal zone. The labeled ascending axons were located more laterally and to the perimeter of this zone. In stages 27--29, the labeled axons near the site of injection were located in the inner two-thirds of the marginal zone, and in the progressively more rostral levels the label shifted laterally and to the perimeter in a laminar pattern. It was concluded that the labeled ascending axons from an individual spinal segment layer upon the axons which originate from the more rostral segments. How this rigid time and position pattern of axonal growth can impart a topographic order into fiber tracts of the spinal cord was discussed.

Afferent Pathways↗