PubMed HealthSearch

Biomedical subjects

G Rager

Publications and source records attributed to G Rager.

18 recordsLinked to original sources

Postnatal development of area 17 callosal connections in Tupaia.

The goal of the present study was to investigate the pattern of maturation of callosal projecting neurons in a well-studied mammalian visual system with unique structural and functional properties. Studies of the distribution pattern of interhemispheric connections in the adult tree shrew primary visual cortex reveal not only a high concentration of labeled neurons along the area 17/18 border, as in standard experimental animals such as the cat and monkey, but also numerous callosal projecting neurons in the adjacent dorsal part of area 17, which largely corresponds to the binocular visual field (Kretz and Rager, Exp. Brain Res. 82:271, '90). Callosal projections were anatomically traced in 11 tree shrews (Tupaia belangeri) at various ages between postnatal day 7 (7, 9, 10, 13, 15, 17, 19, and 26 days old) and adulthood (107 days old). In each animal, four injections of wheat germ agglutinin conjugated to horseradish peroxidase were made in a standard configuration into the striate cortex of one hemisphere. In young tree shrews only 7 and 9 days old, heavily labeled terminal axon structures could be seen in the white matter and in layer VI of the opposite hemisphere. Only a few labeled neurons, however, were detected in layer III. The small number of labeled neurons indicated that early in postnatal development, only a few callosal axons had invaded the upper cortical layers. By 10 days of age, the number of supragranular neurons was increasing and the maximal value was counted in a 13-day-old tree shrew. A sharp decline in the number of labeled supragranular neurons was noticed--about 94% in our case--between days 13 and 15. In animals more than 15 days old, the distribution pattern and the density of the neurons looked like the pattern seen in the adult Tupaia brain. The labeled cells were mostly concentrated in layers II and III. The majority of neurons resembled typical pyramidal cells. However, some of the neurons in sublayer IIIc had elongated cell bodies oriented parallel to the laminar boundaries. In contrast to the supragranular cells found in all stages investigated, small populations of labeled cells in layer VI were observed in 9- to 17-day-old tree shrews only. In young postnatal animals 7 to 13 days old, a peculiar cell type was labeled on the ipsilateral side. In coronal sections these cell bodies formed a continuous band that extended from the ventricular wall to the subcortical white matter. These cells might belong to a population of cells still in migration.

Animals

Organization of the optic chiasm in the hatched chick.

In the hatched chick the fibers of the two optic nerves segregate into clearly defined bundles when they cross to the other side. These bundles run in horizontally oriented tiers. The tiers are demarcated by blood vessels and pial tissue. The organization of these tiers was investigated qualitatively and quantitatively using light and electron microscopy as well as tracer techniques. The fibers within the tiers cross to the contralateral optic tract without leaving their respective tier. The mean total number of tiers is 34 with a great individual variation. A preference in the superposition of one side over the other could not be observed. Comparing these data with our earlier study (Rager et al.: Anat. Embryol., 179:135-148, 1988) it can be concluded that neither the segregation of fibers into discrete bundles nor the variability in the number of alternating tiers seem to disturb the topography of fibers as it is achieved in the optic nerve. The pattern of vascularization correlates with the order of crossing axon bundles and contributes to the demarcation of the tiers. The chiasm is vascularized by the Aa. preopticae and the A. infundibularis.

Animals

Synaptogenesis in the primary visual cortex of the tree shrew (Tupaia belangeri).

The primary visual cortex of the tree shrew is characterized by the lack of ocular dominance columns. The two eyes are represented in sublayers of laminae 3 and 4. In an earlier study using the transneuronal transport we observed that the geniculate afferents from the two eyes do not initially overlap and then segregate into their appropriate sublaminae. The final distribution pattern can already be observed during the early postnatal period. Since segregation and elimination of afferent terminal branches do not seem to take place, we wanted to investigate whether or not an overproduction of synapses can be observed as in several other animals. We examined layers 3B, 3C, 4A, and 4B, which receive afferents from the retina via the lateral geniculate nucleus, from P5 to maturity by using the electron microscope. The brain tissue was excised in the region where the central vision is represented in adult animals. Then we determined the density of synapses per 100 microns 2 neuropil for each of the four sublayers at the ages P5, P15, P19, P23, P31, and P42 and in the adult animal (AD). In determining the neuropil we measured the size of two additional compartments, i.e., the compartments consisting of perikarya and of blood vessels. At a higher resolution we determined the fraction of Gray type I and type II synapses in each sublamina and in each developmental stage. The size of the neuropil increases from 57% at P5 to 81% in AD whereas the compartment of perikarya decreases from 42% to 15% and the compartment of blood vessels increases from 1.3% to 3.9%. The synaptic density starts with very low values (3.5/100 microns 2) at P5. Then it increases rapidly and attains a maximal rate of increase during the period of eyelid opening. After this period the increase is slowed down and approaches the adult value (12.5/100 microns 2) slowly. An overproduction of synapses could not be observed. The percentage of type I and type II synapses also changes during this period. The fraction of type I synapses amounts to 73% at P5 and increases to 92% in AD. The increase in density of type I synapses is continuous and does not show any sign of overproduction. The density of type II synapses rapidly reaches it final value and then remains constant. Possibly there is a slight overproduction during the period of eyelid opening.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

[The visual cortex of Tupaia: an alternative model?].

The tree shrew has become an interesting animal for research in neurobiology because of its unique structural and functional properties, in particular in its visual system. Some of these properties are described and discussed here as far as they are related to the functional architecture of the primary visual cortex: The representation of the ocularity, the ON and OFF responses, the development of the retino-geniculo-cortical projection, and synaptogenesis. From these data it appears that the tree shrew forms an important exception to the functional organization as it is found in standard experimental animals like cats and monkeys. It challenges a deeper thought on a common basis for both types of organization and development.

Animals

Reciprocal heterotopic callosal connections between the two striate areas in Tupaia.

WGA-HRP injections were placed into area 17 close to the border with area 18 of Tupaia belangeri in order to study the callosal connections of the striate area in this animal. Most callosal neurons were found in the striate cortex (57.6-86.9%), some in the extrastriate area 18 (10.6-28.1%), and a few in even more temporal regions (2.5-14.3%). Concerning only the area 17, reciprocal homotopic connections could be observed as a strip along the area 17/18 border. Additionally, heterotopic callosal connections could be seen in regions representing the binocular visual field, especially the lower part. The area 17 cells were mostly located in the supragranular layers II and III (94.1-97.2%). But neurons could also be found in the infragranular layers, especially layer VI (2.6-5.2%) and in layer IV (0.2-1.1%). Homotopic projections were mostly seen in layers IIIc and V. The majority of the supragranular and infragranular neurons are pyramidal cells. However, a newly defined subpopulation of neurons, most probably stellate cells, were discovered forming a band in sublayer IIIc, very close to the layer III/IV border.

Animals

The cellular origin of the b-wave in the electroretinogram -- a developmental approach.

Retinal ganglion cells and retinotectal synapses of chick embryos can be activated by electrical stimulation at early stages of development (Rager, '76a,b), whereas light evoked responses occur only towards the end of the incubation period. Thus, photoreceptors seem to be the last cells to mature in the chain of elements necessary to enable transmission of visual information to tectal neurons. In the present study the development of light evoked activity in the retina was investigated and compared with the structural maturation of retinal cells. This ontogenetic approach offers a solution to the problem of the cellular origin of the b-wave called in question by recent records of the potassioretinogram (KRG). Lammellar structures in the developing outer segments of photoreceptors can first be observed on incubation day 17. Late on the same day a corneal electroretinogram (ERG) and a visual evoked response on the optic tectum (VER) can be recorded. The response properties of the developing b-wave and VER were tested using various stimulus parameters. From the latencies of the b-wave and of the VER it is concluded that the b-wave is not generated directly by the activity of neurons involved in intraretinal signal transmission. Thus it is necessary to consider secondary processes triggered by neuronal activity such as depolarization of glial cells. In the chick retina, Müller cells are virtually the only glial cells. They fulfill all structural requirements necessary to explain the current which spreads through the retina during the b-wave. Electronmicroscopic analysis reveals that Müller cells undergo drastic changes during the early phase of b-wave development (incubation day 18). In particular, the number of microtubules per unit volume and the surface area of Müller cell processes in the outer plexiform layer increase considerably. It is, therefore, suggested that the b-wave originates in the depolarization of Müller cells secondary to synaptic activity in the outer plexiform layer.i

Animals

Ingrowth and ramification of retinal fibers in the developing optic tectum of the chick embryo.

Onset, temporal sequence, and pattern of ingrowth of retinal fibers into the developing optic tectum of the chicken were investigated with histological procedures including the Golgi technique. Invading fibers could first be detected by stage 34 (eight days of incubation) at a specific locus which is the central area of the optic tectum. Compared to other tectal regions the central area is distinguished at this time by its advanced cytoarchitectural development and by the maturation of dendrites of radial cells located within superficial laminae. Immediately after their arrival at the central area some fibers can be observed invading the outer tectal layers and forming side branches. These observations permit the conclusion that fibers do not wait at their termination site for several days, as has been suggested earlier. Retinal axons start to invade the tectum at the site which is most advanced in its structural development. This early maturation of neurons in a specific tectal region might be a sufficient explanation for the central retinal fibers connecting to neurons of this area, which, propter hoc, is called the central tectal area.

Age Factors

An improved silver stain for developing nervous tissue.

A reduced silver technique using physical development to stain embryonic nervous tissue is described. Brains are fixed in Bodian's fixative. Paraffin sections are pretreated with 1% chromic acid or 5% formol. They are impregnated with 0.01% silver nitrate dissolved in 0.1 M boric acid/sodium tetraborate buffer of pH 8 or with silver proteinate. Finally they are developed in a special physical developer which contains 0.1% silver nitrate, 0.01-0.1% formol as reducing agent, 2.5% sodium carbonate to buffer the solution at pH 10.3, 0.1% ammonium nitrate to prevent precipitation of silver hydroxide, and 5% tungstosilicic acid as a protective colloid. The development takes several minutes in this solution, thus the intensity of staining can be controlled easily. The method yields uniform, complete and reproducible staining of axons at all developmental stages of the nervous tissue and is easy to handle.

Animals

Systems-matching by degeneration. I. A quantitative electron microscopic study of the generation and degeneration of retinal ganglion cells in the chicken.

The total number of optic nerve fibers of the chicken was determined at twenty sequential developmental stages from incubation day 5 to 104 days after hatching. It was found that the total number of optic nerve fibers increases from 4400 on incubation day 5 to about 4.0 million on incubation days 10 and 11. Thereafter, it decreases to a final value of about 2.4 million by incubation day 18 and remains constant from that time on until adulthood. Thus, 40% of optic fibers degenerate. Degenerating ganglion cells in the retina are first detectable by incubation day 9. Initially degenerating cells are located mainly in the central retina, but on subsequent days they can be found predominantly in peripheral zones. It is postulated that cell death occurs because of competition for adequate arborization space. If more retinal afferent fibers arrive than tectal termination sites are available, supernumerary fibers may degenerate. By degeneration the two systems retina and optic tectum, are matched in size.

Age Factors

Systems-matching by degeneration. II. Interpretation of the generation and degeneration of retinal ganglion cells in the chicken by a mathematical model.

Quantitative data on generation and degeneration of retinal ganglion cells during development (Rager and Rager, 1978) are interpreted in terms of a mathematical model which consists of a system of differential equations. By these equations we attempt to describe the formation of retinal ganglion cells and their termination domains in the tectum. Since ganglion cells seem not to degenerate before their axons have arrived at their termination site and start branching, from the arrival time on they may become competent either to continue to mature or to die. Therefore, to find the actual number of competent cells the extension of the fiber pathway between the retina and the optic tectum had also to be measured and computed. The differential equations are united by the principle that at any given time cells in excess of the number of termination domains have to die. By this model the mathematical function was determined. Several parameter values of this function were optimized with the Gauss-Newton method by which the curve was fitted to the measured values. The high correlation obtained by this method allows to conclude that, to a first approximation, the model may be satisfactory. The evidence of competition for termination sites and of systems-matching by cell death is discussed.

Age Factors