Neuropeptide Y immunoreactive axons in the corpus callosum of the cat during postnatal development

Many immunocytochemical studies have identified different types of neurotransmitters localized in the corpus callosum (CC) axons in the adult mammal. Few studies have looked at the development of different neurochemically identified CC systems. Previous studies on the development of cat CC axons have indicated that a large number of transitory CC axons project to the cortex during early postnatal development. The present study focuses on the development of one neurochemically identified group of CC axons in the cat, labeled with an antibody against neuropeptide Y (NPY), to determine if this group participates in transitory CC axonal growth. Cats at specified ages from birth to adulthood were studied with a routine method of immunocytochemistry for antiserum to NPY. NPY-immunoreactive (ir) CC axons were detected at all stages examined, from newborn to adult; the peak density occurred during postnatal weeks (PNW) 3–4. During PNW 1–2, the denisty of NPY-ir CC axons increased gradually; some NPY-ir axons at this age had growth cones located within the CC bundle between the cerebral hemispheres. The density of the NPY-ir CC axons decreased gradually during PNW 5–7, and from PNW 8 to maturity only a few NPY-ir CC axons were observed. These results indicate that at least two types of NPY-ir CC axons (i.e., transitory and permanent) exist during development, and that most of these axons are eliminated or only express NPY-ir for a short period during development. The results also indicate that neurochemical subsets of CC axons participate in the extensive transitory growth observed by means of the membrane tracer DiI but they may follow unique developmental timetables.     

Ocular dominance in striate cortex is altered by neonatal section of the posterior corpus callosum in the cat

Summary In adult cats that had previously undergone surgical section of the posterior corpus callosum at 13–18 days after birth, the striate cortex was examined using extracellular single unit recordings. The receptive fields of the cells examined were located from the vertical meridian to 39 ° peripherally, and ranged from above to below the horizontal meridian. Cells were classified according to type (simple, complex), ocular dominance, receptive field size and location. Callosum sectioned cats had 53% of striate cells activated monocularly as compared to 25% for control cats. This increase in monocularly activated units primarily occurred for receptive fields in the paracentral region of the visual field, from 4–39 °. The age at which the neonatal surgery had occurred was correlated with the individual cat's proportion of monocularly activated cells.Therefore, the increase in monocular activation of striate units occurred within a large portion of the normal binocular visual field. This physiological change was partially predicted by a previous behavioral study showing a substantial loss in the extent of the binocular visual field following neonatal corpus callosum section (Elberger 1979).Support for this research was received from Training Grant No. T-32 EY 07035-02 awarded to the University of Pennsylvania. Additional support was provided by the Department of Neurobiology and Anatomy, University of Texas Medical School at Houston, No. 1-11321-215001-10     

The role of the corpus callosum in the development of interocular eye alignment and the organization of the visual field in the cat

Summary In young cats, the posterior portion of the corpus callosum was sectioned 13–29 days after birth. The animal's eyes were photographed at weekly intervals for six months using the pupil-reflex method. From the corneal reflection evident in the photographs the degree of alignment for the optical axes of each cat was estimated (Sherman, 1972). The 17 experimental cats all showed a significant tendency toward permanent divergent strabismus, as compared to six normal cats. The limits of the visual field were determined for both groups of cats using a perimetry technique similar to that of Sprague and Meikle (1965) and Sherman (1973). With one eye open normal cats responded from 90 ° ipsilateral to 45 ° past the vertical midline into the contralateral visual field. With either eye the experimental cats responded from 90 ° ipsilateral to approximately the vertical midline. The loss of visual responsiveness is within the contralateral region of the normally binocular zone. Three cats received the same operation at 9, 13, or 20 months old. Eye alignment and visual field perimetry were unaffected by the surgery. It is not known whether the observed abnormalities result from arrested development, or disruption of intrinsically determined ocular alignment.     

Postnatal shaping of callosal connections from sensory areas

Summary Horseradish peroxidase (HRP) was injected unilaterally into the first and second visual areas (VI and V2; areas 17 and 18) of 20 kittens aged between 2 and 90 days and into the second somatosensory area (S2) of 16 kittens aged between 1 and 52 days. The radial and tangential (normal and parallel to the pial surface, respectively) distributions of neurones giving origin to callosal axons (callosal neurones) were studied. In adult cats, callosal efferent zones (CZs) are defined by the distribution of callosal neurones. CZs occupy, in the visual cortices, tangentially and radially restricted parts of areas 17, 18, 19 of the lateral suprasylvian gyms and in the somatosensory cortices, parts of SI and S2. At birth, callosal neurones are distributed throughout the tangential extent of visual and somatosensory areas; they are also more widespread in depth than in the adult. During the first postnatal month, as a result of the gradual disappearence of callosal neurones from parts of the visual and somatosensory areas, the adult CZs emerge. The CZ in areas 17 and 18 undergoes a further tangential reduction during the second and third postnatal months.Supported by the Swiss National Science Foundation (3.492.075 and 3.319-0.78)Part of these experiments was carried out at the Institute of Physiology of the University of Ancona     

The postnatal development of visual callosal connections in the absence of visual experience or of the eyes

Summary Counts of callosal neurons retrogradely labeled by horseradish peroxidase (visualized using multiple substrates) were obtained in areas 17 and 18 of five kittens reared with their eyelids bilaterally sutured and of three kittens which had undergone bilateral enucleation on postnatal days 1–4. These counts were compared with those obtained in normal adult cats.The normal adult distribution of the callosal neurons results from the gradual postnatal reduction of a more widespread juvenile population. Binocular visual deprivation by lid suturing dramatically decreases the final number of callosal neurons and narrows their region of distribution (callosal zone) in areas 17 and 18. A less severe reduction in the final number of callosal neurons is caused by bilateral enucleation, which also increases the width of the callosal zone compared to that of normal cats. Thus, visual experience is necessary for the normal stabilization of juvenile callosal connections. However, since some callosal neurons form connections in the absence of vision, other influences capable of stabilizing juvenile callosal neurons also exist. These influences are probably antagonized by destabilizing influences or inhibited, when the eyes are intact.This work was supported by Swiss National Science Foundation grant 2.219.0.78 to Dr. G.M. Innocenti; Dr. D.O. Frost received a fellowship from the American-Swiss Foundation for Scientific Exchange     

Differential dynamics of transient neuronal assemblies in visual compared to auditory cortex

Large-scale, coherent, but highly transient networks of neurons, ‘neuronal assemblies’, operate over a sub-second time frame. Such assemblies of brain cells need not necessarily respect well-defined anatomical compartmentalisation, but represent an intermediate level of brain organisation between identified brain regions and individual neurons dependent on the activity status of the synaptic connections and axonal projections. To study neuronal assemblies both in slices and in the living brain, optical imaging using voltage-sensitive dyes (VSDI) offers the highest spatial and temporal resolution in real-time. Applying VSDI technique to compare assemblies in visual versus auditory cortices under standardised experimental protocols, we observed no significant variations in the basic parameters of fluorescence signal and assembly size: such results might be predicted from the canonical invariance of cortical structures across modalities. However, further analysis revealed less obvious yet significant differences in the assembly dynamics of the two regions. The neural assemblies spread widely across layers in the two cortices following paired-pulse stimulation of putative layer 4. The respective patterns of activity started to differentiate within a specific time frame (250–300 ms). The signal was predominant near the point of stimulation in the visual cortex, whereas in the auditory cortex the signal was stronger in the superficial layers. This modality-specific divergence in assembly dynamics highlights a previously under-appreciated level of neuronal processing. Additionally, these findings could prompt a new approach to the understanding of how information from different senses, transmitted as action potentials with identical electrochemical characteristics across different cortices, be it visual or auditory, can eventually yield, nonetheless, the qualitatively distinct experiences of seeing or hearing.     

Emergence of callosally projecting neurons with stellate morphology in the visual cortex of the kitten

Summary Callosally projecting neurons in areas 17 and 18 of the adult cat can be classified into two types on the basis of their dendritic morphology: pyramidal and stellate cells. The latter are nearly exclusively of the spinous type and are predominantly located in upper layer IV. Retrograde transport of the carbocyanine dye DiI, applied to the corpus callosum, showed that, up to P6, all callosally projecting neurons resemble pyramids in the possession of an apical dendrite reaching layer I. At P10, however, callosally projecting neurons with stellate morphology were found. A study was designed to distinguish whether these neurons are late in extending their axons to the corpus callosum or, alternatively, have transient apical dendrites. To this end, callosally projecting neurons were retrogradely labeled by fluorescent beads injected in areas 17 and 18 at P1–P3 and then either relabeled with DiI applied to the corpus callosum at P10 or intracellularly injected with Lucifer Yellow at P57. Double-labeled stellate and pyramidal cells were found in similar proportions to those found for the total, single-labeled population of callosally projecting neurons. It is therefore concluded that callosally projecting spiny stellate cells initially possess an apical dendrite and a pyramidal morphology. At P6, i.e. close to the time when stellate cells appear, layer IV neurons with an atrophic apical dendrite were found, suggestive of an apical dendrite in the process of being eliminated.     

Transcallosally evoked responses in the visual cortex of normal and monocularly enucleated rabbits

Summary In visual cortex of normal adult rabbits, callosal projections are restricted to a 2 mm wide band at the area 17/18 border. In adult rabbits which are monocularly enucleated (ME) on the day of birth, the callosal zone extends 4 mm into the medial region of area 17 in the cortex ipsilateral to the remaining eye. In this study, the function of these anomalous callosal projections in ME rabbits was investigated using electrophysiological techniques. A microelectrode was placed in the visual cortex ipsilateral to the enucleated eye at the 17/18 border, bipolar stimulating electrodes were placed in a homotopic location in the contralateral cortex, and averaged evoked responses (AERs) to stimulation were recorded. The stimulating electrodes were then moved mediolaterally in 1 mm steps, and the AERs were recorded for each location of the stimulating electrodes. In the normal rabbit, a maximal short latency evoked response was recorded when the stimulating electrodes were at a location homotopic to the recording electrode. When the stimulating electrodes were moved a distance of 1 mm or more from this optimal position, this short latency response was either absent or dramatically decreased in amplitude, reflecting the precise topographic pattern of the normal callosal projection. In contrast, in ME rabbits, a consistent response was evoked at the 17/18 border when the stimulating electrodes were moved as much as 3 mm medial to the homotopic position. Since antidromically activated responses and both pre- and post-synaptic orthodromically activated responses contribute to the AER, recordings were also made from single cells in some animals. Orthodromically activated single cell responses were evoked by electrical stimulation in the abnormal medial callosal zone of ME rabbits. The data indicate that abnormal callosal projections in ME rabbits can mediate functional interactions between nonhomotopic areas of the primary visual cortices.     

Binocularity and single cell acuity are related in striate cortex of corpus callosum sectioned and normal cats

Summary Cats with corpus callosum section at 4–37 postnatal days underwent electrophysiological recording in striate cortex after they reached adulthood. Single cells were examined to determine both their ocular dominance and spatial frequency threshold (acuity). Data were analyzed for each cat according to the extent of binocular interaction (binocularity) and the mean striate acuity. Both visual functions were found to be significantly related to the age at which the corpus callosum section occurred, with the greatest deficits in visual function resulting from callosum section at the younger ages. There was a significant relationship between striate binocularity and acuity in the callosum sectioned, as well as in normal, cats. This suggests that visual resolution is at least partially determined by the ability to integrate information from both eyes.     

Spatial frequency thresholds of single striate cortical cells in neonatal corpus callosum sectioned cats

Summary Following section of the corpus callosum at 1–6 postnatal weeks in cats, behavioral visual acuity was measured binocularly and monocularly from 6–29 postnatal weeks; physiological determination of spatial frequency thresholds of single striate cortical cells was performed when the cats were at least 8 months old. Results were compared between cats with callosum section at each postnatal week, as well as with normal cats. Cats with callosotomy at 1–3 postnatal weeks had deficits in behavioral visual acuity, and the deficits were greatest in the youngest operated cats. Cats with callosotomy at 1–2 postnatal weeks failed to resolve as high spatial frequencies as did normal cats, and the resolution of the 1 week operated cats was lower than the resolution of the 2 week operated cats. Cats with callosotomy at 3–6 postnatal weeks had spatial frequency thresholds that were equivalent to those of normal cats. To determine what kinds of striate cells had reduced spatial resolution following neonatal corpus callosum section, cells were categorized according to class (Simple, Complex), receptive field location (Central, Peripheral), and monocular behavioral acuity eye performance (Better Eye, Worse Eye). Cats with corpus callosum section during postnatal week 1 had the lowest spatial resolution for all cell categories compared to all groups tested. However, cats with callosum section during postnatal week 2 had normal spatial frequency thresholds for Simple, Central and Better Eye categories. The cats with callosum section in postnatal weeks 3–6 had normal spatial frequency thresholds for all cell categories. For corpus callosum sectioned cats with and without visual deficits, and for normal cats, visual acuity measured behaviorally is significantly related to visual acuity measured physiologically. The results show that neonatal corpus callosum section in cats can affect behavioral visual acuity, as well as the spatial frequency thresholds of many categories of striate cortical cells. However, callosum section at different ages affects different populations of cortical cells. Furthermore, the results suggest that neonatal corpus callosum section may directly affect a single fundamental property of cells in primary visual cortex with a resulting disruption of many visual functions.     

A direct pathway from thalamus to visual callosal neurons in cat

Summary Horseradish peroxidase was injected in the right visual cortex and a large electrolytic lesion made in the left lateral geniculate nucleus of an adult cat. Neurons of origin of the callosal projection to the injected cortex were identified by retrograde labelling and selected for electron microscopic study. Degenerating thalamo-cortical axon terminals were found to contact a labelled stellate cell in layer IV and a labelled pyramidal cell in layer III at the border region of areas 17 and 18. We conclude that there is a monosynaptic pathway from lateral geniculate nucleus to the cells of origin of callosal axons to the contralateral visual cortex.Supported by the Swiss National Science Foundation (3.0950.77)     

Types of callosally projecting nonpyramidal neurons in rat visual cortex identified by lysosomal HRP retrograde labeling

Summary Callosally projecting neurons, labeled following injection of horseradish peroxidase (HRP) into the 17/18a border of the contralateral hemisphere, have been examined by light and electron microscopy. These neurons exhibit two types of horseradish peroxidase labeling: either a diffuse, Golgi-like labeling, or a granular, punctate labeling. The punctate type of HRP-labeling is the predominant form in nonpyramidal neurons, while pyramidal neurons frequently display either diffuse or punctate labeling. Only punctately labeled neurons have been examined in this study. Light microscopic analyses of 1-m sections show that in the heavily labeled zone at the area 17/18a border approximately 9% of all of the cells in layer II/III are callosally projecting nonpyramidal cells, and 70% of them are callosally projecting pyramidal cells. Light and electron microscopic examinations indicate that the nonpyramidal neurons are a heterogeneous group which consists of small multipolar neurons, large multipolar neurons, small bipolar neurons, and large bipolar neurons. To investigate the ultrastructural appearance of the punctate HRP labeling, selected neurons have been examined in thin sections. In the electron microscope, the tetramethylbenzidine (TMB) reaction product appears as electron-dense crystals, while the diaminobenzidine (DAB) reaction product appears as dark, electron-dense material which fills the lysosomes. These lysosomes occasionally have a halo of reaction product, but often they are not morphologically distinguishable from dark lysosomes present within neurons from control animals in which the darkening results from staining the thin sections with lead citrate and uranyl acetate. However, labeled neurons possess more dark lysosomes than neurons from control animals. These additional dark lysosomes presumably contain the HRP reaction product visible by light microscopy.     

Cross-modal innervation of primary visual cortex by auditory fibers in congenitally anophthalmic mice

Auditory-visual cross-modal innervation was examined in control (sighted, ZRDCT-N) and congenitally anophthalmic (eyeless, ZRDCT-AN) mice using electrophysiological recording and pathway tracing with carbocyanine dyes. Electrophysiological data demonstrate that the primary visual cortex of congenitally eyeless, blind, mice receives auditory stimuli. Neuroanatomical data demonstrate a direct connection between the inferior colliculus (IC) and visual cortex. Our experiments provide new information about how the brain adapts to the loss of sight.     

Distribution of the cells of origin of the corpus callosum and anterior commissure in the marmoset monkey

Summary The neurons of origin of the great cerebral commissures of the marmoset monkey were identified by horseradish peroxidase histochemistry and their distribution was studied. Six adult marmosets were used. Three were normal: the others were subjected to section of the corpus callosum (CC), sparing the anterior commissure (AC). All six were injected with horseradish peroxidase throughout one cerebral hemisphere. The three normals provide information on the origins of both the CC and AC, whereas the three callosotomized monkeys allow study of the origins of the AC alone. All CC and AC neurons in the marmoset are pyramidal cells. Except for layer I, all cortical layers possess commissural cells; their laminar organization varies according to cortical area. There exists a progression in predominance from supra- to infragranular commissural neurons proceeding from temporal through occipital to parietal and finally to frontal cortex. Major acallosal zones are found in the primary visual cortex and the fore- and hindlimb representations of the somatosensory cortex. Correlations between commissural neuron distribution and cytoarchitectonic areas are not always obvious. Commissural neurons were not organized in columnar fashion.     

新疆地区正常成人脑干,胼胝体,小脑的MRI正中矢状径线测量

目的 本文应用ＲＭＩ测量了６８例新疆地区正常成人的正中矢状面的正常脑干、胼胝体、小脑各径线数值,以确定中国人正常值范围,为诊断颅脑疾病提供正常数据。方法 正常成人６８例,用ＤｉａｓｏｎｉｃｓＭＴ／Ｓ超导磁共振成像系统ＳＥ序列,５ｍｍ导厚,正中矢状面Ｔ１Ｗ１图像,直接光标测量统计分析。结果 胼胝体前后最大距离男６３．２４ｍｍ,女６４．２０ｍｍ;胼胝体水平的大脑前后径国国３３８．１３ｍｍ,女１３７．９     

Postnatal development of the visual cortex of the mouse after enucleation at birth

Summary Quantitative data in the neocortex up to the age of 180 days (neuronal densities, number of neurones, glial cells, dendritic intersections and spines) were compared in normal mice and mice enucleated at birth.Bilateral enucleation induced an increase of neuronal density in all cortical layers of areas 17, 18a, and 41, the supragranular layers II–III being more affected than layers IV–VI. This was noticed in layer II 10 days after the operation and was maximal in all layers between 30 and 60 days; at 180 days there was some return to normal of the neuronal density in all layers. The total number of neurones and glial cells were the same in the bilaterally enucleated mice as in the controls. No reaction in dendritic branching was evident for pyramids of layers III and V in areas 17 and 41 after bilateral enucleation. In contrast the number of spines was reduced on the apical dendrites of pyramids from layers III and V in area 17, but not in area 41.After unilateral enucleation the reaction was less severe and delayed compared with bilateral enucleation, the first signs of increase of neuronal density appearing 30 days after the lesion in the contralateral hemisphere. The contralateral areas 17 and 18a were more affected than the ipsilateral ones and area 41 showed no change compared to the control. As after bilateral enucleation, layers IV and V were least affected by unilateral enucleation in both ipsi- and contralateral cortices.These results suggest that deafferentation in an immature system affects the development of all cortical layers but with a greatest intensity in supragranular layers, which are not the main direct targets of thalamo-cortical input.Supported by the Swiss National Research Foundation, Grants no. 3-641-71 and 3-434-74     

Postnatal development of zinc-containing cells and neuropil in the visual cortex of the mouse

Summary The postnatal development of zinc-containing synaptic boutons and their cells of origin in the visual cortex of a pigmented mouse is described. Two phases can be distinguished. During the early phase zinc-containing neuropil is first apparent by postnatal day 3. By day 7 a light, but distinct neuropil staining sketches the primary and secondary visual cortices. The primary visual area contains light precipitate in layers V and VI as well as the monocular portion of layer II/III. The secondary visual areas contain slightly denser precipitate in layers II/III through VI. The transition to the second phase is marked by a large increase in precipitate density by day 11. Thereafter, the intensity of the neuropil staining increases to day 28, first in layer II/III and then in layer V, as the adult pattern of neuropil staining gradually develops. In the primary visual cortex precipitate is dense in layers II/III and V, moderate in layer VI, and sparse in layers I and IV. In the secondary visual areas the precipitate is dense in layers II/III and V and moderate in the lower portion of layer I and in layers IV and VI. Cells of origin of zinc-containing boutons are visible by the end of the second postnatal week in layer II/III of the secondary visual cortex. By 21 days of age the pattern of staining in the mature mouse is established, and cells in layers II/III and VI are labeled in both the primary and secondary visual cortices. The developmental sequence of zinc-containing cells and neuropil does not preclude an involvement of zinc in the postnatal regulation of NMDA receptor function.     

Identification and localisation of auditory areas in guinea pig cortex

The organisation of guinea pig auditory cortex was studied by combining histological methods with microelectrode mapping. This allowed the location of seven auditory areas to be determined in relation to the visual and primary somatosensory areas. The auditory areas were identified by single-unit recordings and their borders defined by evoked potential mapping. The visual areas were identified by their relatively high densities of myelinated fibres, while the primary somatosensory cortex was identified by its characteristic barrels of high cytochrome oxidase (CYO) activity in layer IV. The auditory region had moderate levels of CYO and myelin staining. When staining was optimal, there was a clear edge to the moderate CYO activity, which apparently corresponds to the dorsal border of the primary auditory area (AI) and the other core field that lies dorsocaudal to it (DC). Thus the primary somatosensory area and the visual and auditory regions were separated from each other by a region with lower levels of CYO and myelin staining. The ventral borders of AI and DC could not be determined histologically as there were no sharp transitions in the levels of CYO or myelin staining. The two core areas were partially surrounded by belt areas. The dorsorostral belt and most of the belt around DC responded more strongly to broad-band stimuli than pure tones, while the ventrorostral belt, small field and a belt zone ventral to the rostral part of DC responded better to pure tones. Units in the small field (S) typically had higher thresholds and broader tuning to pure tones than AI, while units in the ventrorostral belt typically had longer onset latencies and gave more sustained responses than units in AI.