Directional selectivity in hamster superior colliculus is modified by strobe-rearing but not by dark-rearing.

Visual response properties of superior collicular neurons of normal hamsters were compared with those of animals reared from birth to adulthood in either total darkness or with stroboscopic illumination. Directional selectivity was markedly reduced only in the strobe-reared animals, thus demonstrating visual plasticity in a system that develops apparently normally without visual experience.

The formation of a cortical somatotopic map.

The primary somatosensory cortex of small rodents is an isomorphic representation of the body surface. Similar representations are characteristic of the subcortical pathways, leading from the periphery to the cortex, and these representations develop in a sequence that begins at the periphery, and that ends in the cortex. Furthermore, central representations at all levels of the neural axis are altered by perinatal perturbations of the peripheral surface. This has led to the hypothesis that the periphery plays an instructional role in the formation of central neuronal structures. The morphology of this discrete organization has been examined thoroughly during the development of the thalamocortical projections. The mechanism(s) that underlies the formation of these representations remains unclear although some recent evidence suggests the involvement of activity-dependent processes that are modulated by 5-HT.

Expansion of the ipsilateral visual corticotectal projection in hamsters subjected to partial lesions of the visual cortex during infancy: anatomical experiments.

Electrophysiological methods were employed to determine whether or not partial visual cortical lesions in neonatal (7--11-day) hamster produced large scotomas in the cortical visual representation. In cases where such scotomas were present electrophoretic deposits of radioactive amino acids in the visually responsive "cortical remnant" of the damaged hemisphere resulted in labelling throughout the lower portion of the stratum griseum superficiale and the stratum opticum of the ipsilateral superior colliculus. No differential labeling of the part of the colliculus which was topographically matched with the remaining visual representation in the cortical remnant was observed. In normal hamsters relatively localized, visual cortical deposits of radioactive amino acids resulted in superficial layer labeling only in portions of the colliculus which corresponded to the locus of the cortical deposit. In a similar fashion, small lesions at physiologically defined loci in the cortical remnant produced degeneration throughout most of the superficial tectal laminae, but a more restricted "focus" of denser degeneration was also visible in these cases. The position of this focus in the colliculus for a given cortical lesion varied with the nature of the visual map in the cortical remnant. In several additional neonatally brain-damaged hamsters large lesions of the visual cortex in the intact hemisphere were combined with radioactive amino acid deposits in the cortical remnant to determine whether or not axons from the crossed corticocollicular pathway previously demonstrated in such hamsters were intermingled with fibers from the ipsilateral corticotectal projection. In alternate sections processed for autoradiography or by the Fink-Heimer ('67) method autoradiographic label and degeneration argyrophilia were both observed in the medical part of the colliculus ipsilateral to the neonatal cortical lesion.

Expansion of the ipsilateral visual corticotectal projection in hamster subjected to partial lesions of the visual cortex during infancy: electrophysiological experiments.

Single unit recording from cells in the superior colliculus ipsilateral to the damaged hemisphere in hamsters subjected to unilateral removal of a part of the posterior neocortex during infancy was combined with electrical stimulation of the cortical remnant and the visual cortex in the undamaged hemisphere. Cells activated by stimulation of the cortical remnant were recorded in all portions of the colliculus. No differences in percentages of driven cells or threshold current intensities were noted between electrode penetrations in which collicular neurons having receptive fields within the remaining visual cortical representation were recorded and tracks where units with receptive fields outside this region were isolated. In the medical part of the tectum ipsilateral to the damaged hemisphere cells driven by stimulation of either cortex were encountered. It was also demonstrated that stimulation of the ipsilateral cortical remnant and/or the contralateral cortex was capable of suppressing discharges normally elicited by optic chiasm or visual stimulation in a manner qualitatively similar to that observed for collicular cells in normal hamsters. The response properties of cells functionally influenced by the ipsilateral and/or contralateral corticles were not different from those of neurons which received no demonstrable cortical input. The receptive field characteristics of the sample of neurons recorded were, on the whole, quite similar to those of collicular neurons in hamsters subjected to lesions of the visual cortex as adults.

Cortical and spinal somatosensory input to the superior colliculus in the golden hamster: an anatomical and electrophysiological study.

The horseradish peroxidase technique was used to identify the sources of somatosensory afferent fibers to the hamster superior colliculus. These experiments demonstrated that the tectum receives axons from pyramidal cells in layer V of the ipsilateral sensorimotor cortex, contralateral lamina IV of all levels of the spinal cord, the contralateral dorsal column nuclei, lateral cervical nucleus, internal basilar nucleus, and nucleus of the spinal trigeminal tract. Electrical stimulation of the spinal cord coupled with extracellular single unit recordings concentrated, for the most part, in the posterior portion of the tectum revealed that such stimuli activated approximately 40% of the cells tested. Almost all of these units were isolated ventral to the stratum opticum and 86% were responsive only to somatosensory stimulation. Analysis of the latencies of collicular responses obtained with two point spinal stimulation in intact hamsters and in animals subjected to somatosensory cortical and/or spinal damage indicated that the initial impulse elicited from most collicular cells was mediated by a polysynaptic pathway(s) which probably synapses in the dorsal column, lateral cervical, and/or internal basilar nuclei. Damage to the dorsal spinal cord and/or somatosensory cortex altered neither the incidence nor the response characteristics of spinally driven collicular neurons. This indicated that most somatosensory collicular cells also received input from the spinotectal fibers which travel in the ventrolateral quadrant. Electrical stimulation of somatosensory cortex activated about 20% of the cells tested in the ipsilateral superior colliculus. If cortical and spinal stimulation were delivered with an interstimulus interval ranging between 50 and 80 msec the response of the tectal neuron to the latter stimulus was suppressed in most cases. This was true regardless of the order of the stimulus pairing. Concurrent somatosensory cortical shocks also suppressed responses to tactile stimuli for 21% of the cells tested.

Functional properties of the corticotectal projection in the golden hamster.

Approximately 31% of the cells recorded in the hamster's superior colliculus could be activated by stimulation of the ipsilateral primary visual cortex. While cortically activated cells were encountered in all laminae of the colliculus where visual cells were isolated, the highest probability of driving visual cells was observed in the deeper laminae, that is, those ventral to the stratum opticum. Response latency, jitter (latency variability), latency shifts as a function of shock intensity, thresholds, and spike numbers did not vary as a function of depth in the colliculus. There was a clear correspondence between the visual fields of the best cortical stimulus points and the receptive fields of cortically activated cells recorded in the superficial laminae of the colliculus. However, there was considerably less retinotopic fidelity for the cortical areas from which cells isolated in the deeper laminae could be driven. This suggests a greater degree of convergence from relatively widespread cortical regions upon visual cells of the deeper laminae. The visal organization) of the cortically activated cells did not differ appreciably from the overall sample of visual cells recorded in the colliculus. Only 3 of the 159 cells tested were driven by stimulation of the contralateral visual cortex and two of these were responsive only at very long latencies.

Conduction velocity distribution of the retinal input to the hamster's superior colliculus and a correlation with receptive field characteristics.

Cells driven reliably by shocks delivered to the optic nerve or optic chiasm were encountered throughout the depth of the colliculus. The incidence of such cells, however, decreased markedly in the laminae ventral to the stratum opticum. The distribution of conduction velocities for the retinal afferents to the tectum was quite broad (range: 1.7-25.5 m/sec) and clearly biomodal with peaks at about 6 and 12 m/sec. A small number of cells were innervated by rapidly (greater than 15 m/sec) conducting axons. No evidence of an indirect-fast pathway from the retina to the colliculus via the lateral geniculate nucleus and visual cortex was obtained. Afferent conduction velocity was not correlated with retinal eccentricity, collicular depth or speed selectivity. It was, however, clearly related to directional selectivity. Ninety percent of the tectal neurons receiving inputs from axons having conduction velocities of less than 5 m/sec were directionally selective while only 41% of those neurons innervated by more rapidly conducting fibers (greater than 5 m/sec) exhibited selectivity. One hundred and sixteen cells in the anterior portion of the colliculus were tested with shocks delivered to the ipsilateral optic nerve and photic stimulation of the ipsilateral eye. Of these, 11% exhibited some degree of binocularity and only 6% were responsive to optic nerve shocks. These electrophysiological findings were correalted with the limited nature of the retinal input to the ipsilateral superior colliculus.

An autoradiographic study of the retinotectal projection in the golden hamster.

Intraocular injections of tritiated leucine and proline were used to examine the retinotectal projection of the golden hamster. In the contralateral superior colliculus intense and complete label was seen in the stratum zonale, stratum griseum superficiale and the upper portion of the stratum opticum, with relatively less dense label in the lower part of the optic layer. On the ipsilateral side no label was found in the most rostral portion of the tectum. This area comprised about 10% of the rostro-caudal extent of the colliculus, and most likely, it receives a crossed input from the temporal retina (as demonstrated in the cat by Harting and Guillery, '76). Very sparse label was observed in an anterior segment of the ipsilateral colliculus. In coronal sections it appeared as discrete clumps or patches which were confined to the stratum opticum. Within this layer there was a tendency for the clumps to be located more dorsally with increasing laterality. There was considerable variability between and within animals in the size of the clumps as well as the distance between clumps. Reconstruction of coronal sections showed that the ipsilateral label forms discontinuous ribbons which extend up to 180 microns in the rostro-caudal dimension. No label was seen on the ipsilateral side in the remaining tectum (caudal 60%).

Representation of whisker follicle intrinsic musculature in the facial motor nucleus of the rat.

Retrograde transport of wheatgerm-agglutinin horseradish peroxidase (WGA-HRP) and fluorescent tracers (true blue-TB, nuclear yellow-NY, and diamidino yellow-DY) from isolated whisker follicles was used to define the somatotopic organization of the facial (VII) motoneurons which innervate the intrinsic follicle muscles. Motoneurons supplying these muscles were restricted almost completely to the lateral (Martin and Lodge, '77) facial subnucleus and the motoneurons which innervated a given follicle were distributed over the entire length of this subnucleus. Cells projecting to dorsal (A-row) follicles were located in the most lateral part of the lateral subnucleus, while those supplying ventral (E-row) follicles were restricted to the medial part of the subnucleus. Injections of different tracers into rostral and caudal follicles within a given row revealed no somatotopic representation of the rostrocaudal axis of the whiskerpad. Additional control experiments demonstrated that some of the labelling obtained with WGA-HRP resulted from spread of this tracer to extrinsic muscles. This was not the case with the fluorescent tracers. The results of the control experiments suggested further that a significant percentage of the motoneurons in the lateral facial subnucleus innervate only intrinsic follicle muscles.

Effects of neonatal infraorbital lesions upon central trigeminal primary afferent projections in rat and hamster.

Transganglionic and anterograde horseradish peroxidase transport was used to evaluate the central projections of undamaged trigeminal (V) nerve branches in adult rats and hamsters subjected to transection of the infraorbital nerve and to cauterization of the vibrissae follicles at birth. In rats, deafferented regions of the V brainstem nuclear complex did not receive abnormal projections from undamaged mandibular sensory afferents. Undamaged ophthalmic-maxillary fibers also failed to terminate heavily in the region deafferented by the neonatal infraorbital lesions. In the hamster, on the other hand, neonatal infraorbital nerve lesions were associated with statistically significant increases in mandibular terminal fields in the principalis, subnucleus interpolaris, and subnucleus caudalis. Tracing experiments were also carried out in neonatal rats and hamsters to determine whether the above-described differences in the response to infraorbital nerve damage reflected a difference in the maturity of the V primary afferent projections to the brainstem at the time of our neonatal lesions. In neonatal rats, the infraorbital and mandibular projections to the V brainstem nuclear complex were quite adultlike, both in their pattern and in the extent of their overlap, which was minimal. Overlap between mandibular and infraorbital terminal fields was also minimal in the newborn hamsters.

Receptive field characteristics of superior colliculus neurons and visually guided behavior in dark-reared hamsters.

Visual response properties of single neurons in the superior colliculus of golden hamsters reared from birth to adulthood in total darkness were compared to those of normal hamsters. Directional selectivity, speed preferences, and receptive field organization in dark-reared hamsters were essentially the same as those found in normally reared animals. Subtle neurophysiological effects of visual deprivation were indicated by the longer latencies of "on" responses to flashed spots of light in the dark-reared animals. Also, in the visually deprived animals three cells were encountered which changed their responses from phasic to tonic as the size of the visual stimulus was increased. In normally reared animals all cells responding to stationary stimuli showed only phasic responses regardless of spot size. Behaviorally, dark-reared animals could not be distinguished from normal animals on the basis of visual orienting and following tests. It was concluded that in the golden hamster visual experience during development has a minimal role in the induction or maintenance of the normal functional development of the superior colliculus.

An electronmicroscopic analysis of the optic nerve in the golden hamster.

The electronmicroscopic examination of sections taken from the hamster's optic nerve 5 mm behind the globe indicated that the nerve contains 110,165 +/- 4,177 (p less than 0.05) fibres of which 96.4% are myelinated. The fibre diameter distribution is unimodal with a peak at 1.2 micrometer and axon diameters ranging from 0.20 micrometer to 3.93 micrometer. Fibres of all sizes are distributed uniformly throughout the cross section of the nerve. The thickness of the myelin sheath surrounding a given axon is highly (0.80) correlated with axonal diameter and the degree of myelination for a fibre of a given size is nearly constant throughout the nerve's cross section. In nerve sections taken just posterior to the globe most (64%) of the fibres counted are unmyelinated and the percentage of unmyelinated axons is highest near the peripheral boundary of the nerve. The process of myelination is essentially complete in sections taken 3.5 mm behind the eye. These differences in the myelination of the proximal and distal nerve most probably account for the discrepancy between the results reported here and those provided by a previous study (Tiao and Blakemore, '76) concerned with the structure of the optic nerve in this species.

Structure-function relationships in rat brainstem subnucleus interpolaris: IV. Projection neurons.

In a companion paper (Jacquin et al., '89), the structure and function of local circuit (LC) neurons in spinal trigeminal (V) subnucleus interpolaris (Sp Vi) were described. The present report provides similar data for 44 projection neurons in Sp Vi. Of these, 25 thalamic, 16 cerebellar, 2 superior collicular, and 1 inferior olivary projecting neurons were studied. The majority responded to vibrissa(e) deflection, and all except 4 of these had multivibrissae receptive fields. The remainder were responsive to either guard hair deflection or indentation of glabrous skin. Latencies to V ganglion shocks were suggestive of monosynaptic activation from the periphery. Sp Vi projection neurons were topographically organized in a manner consistent with that of their primary afferent inputs. Nonvibrissa sensitive cells had diverse morphologies. Morphometric analyses of the more heavily sampled thalamic and cerebellar projecting, vibrissa(e)-sensitive cells indicated the following. (1) As compared to LC neurons, projection neurons had bigger receptive fields, cell bodies, dendritic trees, and axons; less circular dendritic trees; a greater preponderance of spiny dendrites and fewer axon collaterals in Sp Vi. (2) Dendritic tree extent correlated significantly with receptive field size, thus suggesting that dendritic tree size is one mechanism contributing to receptive field size in vibrissae-sensitive projection neurons. (3) V thalamic cells had significantly bigger receptive fields and dendritic trees, and also give off more local axon collaterals, than V cerebellar neurons. Collicular and inferior olivary projecting neurons shared structural and functional attributes with other Sp Vi long-range projecting cells. Structure-function relationships exist for vibrissa-sensitive projection neurons in Sp Vi. The relevant parameters correlating with projection neuron morphology are receptive field size and projection status, whereas for Sp Vi LC neurons the relevant correlative parameter is peripheral receptor association.

Structure-function relationships in rat brainstem subnucleus interpolaris: III. Local circuit neurons.

Intracellular recording, electrical stimulation, receptive field mapping, and intracellular injection of horseradish peroxidase were used to assess the response properties, collateral projections, and morphology of 44 local circuit (LC) neurons in the subnucleus interpolaris (Sp Vi) of the trigeminal brainstem complex of the rat. LC neurons were defined as those with axons restricted to brainstem areas receiving trigeminal primary afferent fibers. Thus, none were antidromically activated from the thalamus, tectum, or cerebellum, and their axons could be seen terminating exclusively within the trigeminal brainstem complex or reticular formation. All neurons sampled were discharged by innocuous or noxious mechanical stimulation of a restricted portion of the face or mouth. They were classified functionally as sensitive to vibrissae (N = 22), nociceptors (N = 9), guard hairs (N = 7), hairy skin (N = 3), or periodontia (N = 3). Fifty percent of the stained neurons were vibrissa sensitive. Twenty-one of these 22 responded to deflection of only one vibrissa. The remaining functional groups also had small receptive fields. Intracellular staining revealed a consistency in vibrissa-sensitive LC morphology. Somata were small to medium in size and multipolar. Their axons had an initial transverse trajectory and gave off recurrent collaterals which arborized extensively in the region of the soma. The parent axon then bifurcated. One branch traveled rostrally to subnucleus principalis while the other branch traveled caudally to subnucleus caudalis. The branches periodically sent collaterals into regions of the trigeminal complex corresponding to the transverse position of the soma. Dendrites extended 440 +/- 140 microns rostrocaudally, forming a tree with a transverse perimeter of 459 +/- 226 microns. Distal dendrites were thin and sinuous, had few spines, and extensively arborized adjacent to the soma. They ended in multiple swellings connected by slender processes. The stereotyped morphology of vibrissa-sensitive LC neurons differed from the variable morphologies of LC neurons activated by nociceptors, guard hairs, hairy skin, or periodontia. Although no group of neurons in one of these categories displayed a distinguishing morphological characteristic, they collectively had features which distinguished them from the vibrissa-sensitive neurons. Non-vibrissa-responsive neurons generally had more expansive, but less circular, dendritic and recurrent axonal arbors; dendrites had more spines, and axons often sent endings into the reticular formation.(ABSTRACT TRUNCATED AT 400 WORDS)

Effects of cortical and thalamic lesions upon primary afferent terminations, distributions of projection neurons, and the cytochrome oxidase pattern in the trigeminal brainstem complex.

Early postnatal lesions of the primary somatosensory cortex alter the vibrissa-related cytochrome oxidase (CO) pattern in nucleus principalis (PrV) of the rat's trigeminal (V) brainstem complex (Erzurumlu and Ebner, '88: Dev. Brain Res. 44:302-308). At present, the reason for this change is not clear. It may be that the corticotrigeminal projection is necessary for the maintenance of vibrissa-related patterns in PrV. However, it is also possible that the loss of the normal pattern of CO activity reflects a change in the organization of brainstem cells resulting from transneuronal retrograde degeneration. In order to address this question, we made lesions of either the primary somatosensory cortex (S-I) or ventrobasal thalamus (VB) in newborn rats and directly assayed distribution of V primary afferents by transganglionic transport of horseradish peroxidase and V-thalamic neurons by retrograde transport of either fluorogold or true blue. Neonatal cortical and thalamic lesions produced no qualitative change in the distribution of primary afferent terminals in either PrV or V subnucleus interpolaris (SpI) beyond that which could be attributed to shrinkage of the brainstem resulting from retrograde degeneration. Most importantly, the "patchy" pattern of terminations observed in normal rats remained apparent in the brain-damaged animals. The normal distribution of V-thalamic neurons in PrV was disrupted by both cortical and thalamic lesions. These cells are normally patterned in a way that matches the distribution of primary afferent terminals and thus that of the mystacial vibrissae. This was not the case in the neonatally brain-damaged rats. Taken together, these results are consistent with the conclusion that neonatal cortical and thalamic lesions disrupt the normal CO pattern in PrV primarily because of their effects upon the patterning of brainstem cells. The present findings demonstrate further that clustering of primary afferents does not require a normal complement of postsynaptic neurons.

Structure-function relationships in rat brainstem subnucleus interpolaris. XI. Effects of chronic whisker trimming from birth.

Whisker trimming from birth reduces activity and alters receptive fields (RFs) in the barrel cortex and thalamus. To assess whether or not this reflects deprivation effects on trigeminal (V) first- and second-order neurons, 59 primary afferents and 343 cells in V brainstem subnucleus interpolaris (SpVi) were studied in rats whose whiskers were trimmed daily for 6-9 weeks from birth. Deprivation did not effect brainstem somatotopy or primary afferent RFs. However, many SpVi cells had abnormal RFs and higher-order inputs, resembling the changes caused by infraorbital nerve injury. For example, in controls, only 3% of whisker-sensitive local circuit neurons responded to more than one whisker, whereas 35% of the deprived and 41% of the infraorbital nerve cut samples had multiwhisker. RFs. Deprived rats also had higher than normal incidences of cells with split or absent RFs, RFs spanning more than one V division, intermodality convergence, and directional or high-velocity sensitivity. Because these changes mimic those caused by nerve section, deprivation may underlie some nerve injury effects on V brainstem RF size and character. Insofar as cytochrome oxidase, anterograde labeling, and unit recordings revealed normal topography in deprived primary afferents and SpVi cells, RF changes in SpVi cells may reflect altered SpVi circuitry. To test this hypothesis, we assessed the morphology of 32 similarly deprived V primary afferents. In SpVi, deprived fibers had normal numbers of collaterals with normal shapes, transverse arbor areas, and topography. However, the total number of boutons per collateral was significantly reduced. Thus, deprivation effects on V higher-order RFs reflect quantitative changes in V afferent terminals.

Evidence for prenatal competition among the central arbors of trigeminal primary afferent neurons: single axon analysis.

Previous studies from this laboratory have demonstrated that prenatal damage to vibrissae follicles results in significant increases in the brainstem representations of the remaining vibrissae as demonstrated by staining for the mitochondrial enzyme cytochrome oxidase (CO). Because CO is primarily a postsynaptic marker, these results do not directly address the question of whether there were changes in the projections of primary afferent fibers. To address this issue, we made intra-axonal recordings from individual vibrissa-related primary afferents in rats that sustained damage to vibrissae follicles on embryonic day 17, and then injected horseradish peroxidase (HRP) into these axons to visualize their terminal arbors in the brainstem at the level of trigeminal subnucleus interpolaris (SpI). All vibrissae-related primary afferents responded to deflection of one and only one vibrissa, and the terminal arbors of axons (N = 47) recovered from animals that sustained fetal peripheral lesions were significantly larger than those (N = 23) from normal rats. Fibers from fetally damaged animals had increased total fiber lengths and numbers of branch points. These results indicate that reduced competition among primary afferent axons results in increases in the terminal arbors that remain. These increases occur without any significant alteration in their peripheral receptive fields.

Transection of the infraorbital nerve in newborn hamsters alters the somatosensory but not the visual representation in the superior colliculus.

The experiments described in this report were designed to determine whether changing the somatosensory representation in the deep laminae of the hamster's superior colliculus would result in a corresponding reorganization in the visual map in the overlying superficial layers. The somatosensory representation was altered by transecting the infraorbital (IO) nerve on the day of birth. This trigeminal branch supplies, among other targets, the snout and mystacial vibrissa follicles. These peripheral structures compose a major portion of the somatosensory representation in the deep collicular laminae. The effects of these lesions were assessed in anatomical and physiological experiments when the animals reached adulthood. Retrograde tracing with true blue demonstrated a 48% reduction in the number of trigeminocollicular neurons in the partially deafferented subnucleus interpolaris (the main source of trigeminal input to the rodent colliculus--e.g., Killackey and Erzurumlu: J. Comp. Neurol. 201:221-242, '81), but anterograde tracing with horseradish peroxidase (HRP) and wheat germ agglutinin-conjugated HRP showed that the terminal field of the trigeminocollicular projection from the deafferented subnucleus was essentially normal. This pathway terminated as a series of patches along the border between the stratum griseum intermediale and stratum album intermedium and encompassed approximately 75% of the rostrocaudal extent of the colliculus. The electrophysiological experiments revealed a marked change in somatosensory collicular topography. There was an under-representation of the vibrissae and the entire IO peripheral field in the deep laminae of the nerve-damaged animals, and neurons with trigeminal ophthalmic and mandibular receptive fields were recorded from portions of the colliculus in which only whisker-sensitive neurons would be normally isolated. There were no corresponding changes in the organization of the visual representation. These results support the conclusion that the organization of the visual representation in the superficial laminae and the organization of the somatosensory map in the deep layers of the mammalian superior colliculus follow independent developmental programs.

Intracellular recording and injection study of corticospinal neurons in the rat somatosensory cortex: effect of prenatal exposure to ethanol.

The effects of prenatal exposure to ethanol on the structure and function of corticospinal neurons was investigated. The subjects were the 3-4-month-old offspring of hooded rats fed a nutritionally balanced liquid diet containing 6.7% (v/v) ethanol (Et), pair-fed a nutritionally matched isocaloric diet (Ct), or fed chow and water (Ch). Corticospinal neurons in primary somatosensory cortex were examined by intracellularly recording and filling cells that were driven by antidromic stimulation of the pyramidal decussation. In the control rats, corticospinal neurons comprised a homogeneous morphophysiological population. Morphologically, all of the antidromically driven cells examined were pyramidal neurons with cell bodies in layer Vb. The dendrites of these neurons were spinous and branched within layers I, IV, and V. Their axons arborized within layers IV, V, and VI and some collaterals extended laterally for distances up to 2.6 mm from the cell body. The mean conduction latency was 3.6 and 3.4 msec for Ch- and Ct-treated rats, respectively. In Et-treated rats, corticospinal neurons constituted a heterogeneous population. The laminar distribution of the corticospinal neurons in Et-treated rats was broad; the cell bodies of labeled neurons were in layers II, IV, V, and VI. The dendrites of layer Vb neurons were spinous; however, many of the spines appeared dysmorphic and the density of spines was significantly greater (32%) in Et-treated rats than in Ct-treated rats. Although the dendritic branching pattern for layer Vb neurons was similar to that described for the controls, a Sholl analysis showed that the complexity and extent of their dendritic trees were significantly greater in Et-treated rats. The axons of all layer Vb neurons in Et-treated rats had long horizontal processes that arborized in layers IV-VI, and some neurons also had an array of collaterals that ascended to layer I. The mean conduction latency for layer Vb neurons was 3.9 msec. The structure and function of ectopic neurons (those in layers II, IV, Va, Vc, and VI) in Et-treated rats differed markedly from those of the layer Vb neurons. Morphologically, the dendritic and axonal fields of these neurons were narrower than for the layer Vb neurons, and the ectopic neurons had a mean conduction latency of 7.1 msec. The heterogeneity of the population of corticospinal neurons in Et-treated rats may result from the effects of ethanol on early events in neuronal development such as neuronal generation and migration.

Neonatally elevated serotonin levels alter terminal arbors of individual retinal ganglion cells in superior colliculus of hamsters.

Previous studies from this laboratory showed that sprouting of serotoninergic (5-HT) axons in the hamster's superior colliculus (SC), induced by a single subcutaneous injection of 5,7-dihydroxytryptamine (5,7-DHT) at birth (postnatal day 0 [P-0]), resulted in an increased terminal distribution of the uncrossed retinocollicular projection that was not associated with any changes in the number or distribution of ipsilaterally projecting retinal ganglion cells. The present study was undertaken to determine what effect this manipulation had on the terminal arbors of such axons. Retinocollicular axons of normal and 5,7-DHT-treated animals were anterogradely labeled with small intraretinal injections of the lipophilic dye 1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate (DiI) on P-16. After tissue processing on P-19, single retinocollicular axon arbors were reconstructed by using confocal microscopy. Quantitative analysis indicated that arbors from 5,7-DHT-treated hamsters had significantly greater total fiber lengths, areas, and volumes than those from normal animals. There were no differences between axons from the two groups in number of branch points, distribution of relative branch lengths, and numbers of bouton-like swellings. These results support the hypothesis that increased SC concentrations of 5-HT alter development of the uncrossed retinocollicular pathway such that a greater territory is covered by individual terminal arbors but that the number of synaptic contacts per arbor remains constant. This may explain, at least in part, the abnormally widespread distribution of the aggregate ipsilateral projection.