On-off units in the first optic chiasm of the blowfly II. Spatial properties

We recorded from the spiking on-off unit in the first optic chiasm (between lamina and medulla) in the blowfly Calliphora vicina, and investigated its spatial properties. The receptive field extends over (11.4±0.9)° horizontally and (8.7±0.6)° vertically, i.e. about 7 by 5 interommatidial angles. The line spread function of the on-off unit — calculated from its response to moving sinusoidal gratings — has a half-width of (2.3±0.2)°. This half-width is slightly broader than that of the photoreceptor. Lateral inhibition occurs when two different areas of the receptive field are stimulated simultaneously. Fast temporal adaptation (i.e. adaptation to trains of short light pulses) takes place independently in different areas of the receptive field.     

On spiking units in the first optic chiasm of the blowfly

We recorded from the spiking sustaining unit in the optic chiasm between lamina and medulla in the brain of the blowfly Calliphora vicina, and investigated both temporal and spatial properties of the light-adapted cell. The sustaining unit fails to follow the highest temporal frequencies followed by the photoreceptor, but its temporal resolution is substantially better than that of the on-off unit. The sustaining unit does not display the fast temporal adaptation as previously described in the on-off unit. As compared with the on-off unit, the sustaining unit has a high sensitivity to small contrasts. Although the sustaining unit continues spiking as long as the light is on, its response is also transient as it adapts rapidly after a change of intensity. The receptive field and the line spread function of the sustaining unit have a similar size and profile: a central lobe with a half-width of approximately 2° surrounded by a circular inhibitory zone located at about 3° off-axis.     

The temporal frequency response function of pattern ERG and VEP: changes in optic neuritis

Steady-state pattern electroretinograms (PERGs) and visual evoked potentials (VEPs) in response to sinusoidal gratings (2 c/deg), sinusoidally counterphased at closely spaced temporal frequencies (TFs) between 4 and 28 Hz, were recorded from 11 patients with unilateral optic neuritis (ON; 11 affected eyes and 10 healthy fellow eyes) and 7 age-matched normal subjects (7 eyes). Amplitude and phase of responses' second harmonics were measured. Responses' apparent latencies were estimated from the rate at which phase lagged with TF. When compared to control values, mean PERG and VEP amplitudes of ON eyes were reduced (by about 0.4 log units) at both low (5–10 Hz) and high (16–20 Hz) TFs. Mean PERG amplitudes of fellow eyes were selectively reduced at low TFs (by about 0.3 log units). Mean PERG apparent latencies of both ON and fellow eyes were delayed (by 15 and 9 ms, respectively). Mean VEP apparent latency of ON eyes was delayed at both low and high TFs (by 24 and 30 ms, respectively), while that of fellow eyes was selectively delayed at high TFs (by 28 ms). The results in ON eyes indicate non-selective abnormalities of PERG and VEP generators responding at both low and high TFs. VEP TF losses may be in part accounted for by corresponding PERG losses. In the fellow eyes of patients, more selective PERG and VEP TF abnormalities may suggest differential impairment of retinal and postretinal subsystems responding better to low and high TFs (i.e. parvo-and magnocellular streams).     

A route for direct retinal input to the preoptic hypothalamus: dendritic projections into the optic chiasm.

With the use of Golgi, horseradish peroxidase, and electron microscopic techniques, neurons within a broad region of the preoptic hypothalamus of the mouse were shown to have dendrites that projected well into the depths of the optic chiasm. Further experimental and ultrastructural investigation demonstrated synapses between these dendrites and retinal axonal boutons within the chiasm. All synapses located in the chiasm were classified as Gray's type I. The possible function of these dendritic projections is discussed.     

Responses of visual interneurons in the fly optic lobe during stimulation by motion in depth

Summary A high-order wide-field neuron in the optic lobe of the fly,Phormia terraenovae (Calliphoridae), has been investigated by stimulation with simple or multiple black stripes on a white background moving sinusoidally towards and away from the head. The variation in spike frequency in correlation with the optical stimulation provides evidence that the unit detects angular velocity as well as more complex features of the stimulus. It responds preferentially to movement towards the head. Spike frequency turned out to be a two-valued function of relative angular velocity across the eye. A simple formula predicts the highly reliable responses to visual stimuli in different parts of the visual field.The author wants to thank Professor G. A. Horridge for his valuable support and clarifying discussions and to Dr. M. Srinivasan for his comments. The project has been supported by grants from the Swedish Council for Natural Science Research.     

Fast temporal adaptation of on-off units in the first optic chiasm of the blowfly

1. We recorded from spiking units in the first optic chiasm between lamina and medulla in the brain of the blowfly (Calliphora vicina). Both previously characterized neuron types, on-off units and sustaining units, were encountered. On-off units had a temporal frequency response with a lower cut-off frequency than blowfly photoreceptors. This low cut-off frequency is related to a fast temporal adaptation of the on-off units to trains of short light pulses. Temporal adaptation occurred independently for short on- and off-pulses. 2. On-off units only responded to stimuli of relatively large contrast. Contrasts of less than 10% gave little or no response.     

Optic chiasm formation in planarian I: Cooperative netrin- and robo-mediated signals are required for the early stage of optic chiasm formation

Freshwater planarians can regenerate a brain, including eyes, from the anterior blastema, and coordinately form an optic chiasm during eye and brain regeneration. To investigate the role of the netrin- and slit-signaling systems during optic chiasm formation, we cloned three receptor genes (Djunc5A, Djdcc and DjroboA) expressed in visual neurons and their ligand genes (DjnetB and Djslit) and analyzed their functions by RNA interference (RNAi). Although each of DjroboA(RNAi), Djunc5A(RNAi) and DjnetB(RNAi) showed a weak phenotype and Djslit(RNAi) showed a severe defect of eye formation, we did not observe any defect of crossing of visual axons over the midline among single knockdown planarians. However, among double knockdown planarians, some of DjnetB(RNAi);DjroboA(RNAi) and Djunc5A(RNAi);DjroboA(RNAi) showed complete disconnection between the visual axons from the two sides, suggesting that some combination of netrin- and robo-mediated signals may be required for crossing over the midline. Finally, we carefully investigated the distribution patterns of cells expressing DjNetB protein, DjnetB, and Djslit at the early stage of regeneration, and found that visual axons projected along a path sandwiched between DjNetB protein and Djslit-positive cells. These results suggest that two different collaborative or combinatory signals may be required for midline crossing at the early stage of chiasm formation during eye and brain regeneration.     

Retinal axon pathfinding in the optic chiasm: divergence of crossed and uncrossed fibers

In the developing mammalian visual system, retinal fibers grow through the optic chiasm, where one population crosses to the opposite side of the brain and the other does not. Evidence from labeling growing retinal axons with the carbocyanine dye Dil in mouse embryos indicates that the two subpopulations diverge at a zone along the midline of the optic chiasm. At the border of this zone, crossed fibers grow directly across, whereas uncrossed fibers turn back, developing highly complex terminations with bifurcating and wide-ranging growth cones. When one eye is removed at early stages, uncrossed fibers from the remaining eye stall at the chiasm midline. These results suggest that crossed and uncrossed retinal fibers respond differently to cues along the midline of the chiasm and that the uncrossed fibers from one eye grow along crossed fibers from the other eye, both guidance mechanisms contributing to the establishment of the bilateral pattern of visual projections in mammalian brain.     

Axon routing at the optic chiasm after enzymatic removal of chondroitin sulfate in mouse embryos

The effects of removing chondroitin sulfate from chondroitin sulfate proteoglycan molecules on guidance of retinal ganglion cell axons at the optic chiasm were investigated in a brain slice preparation of mouse embryos of embryonic day 13 to 15. Slices were grown for 5 hours and growth of dye-labeled axons was traced through the chiasm. After continuous enzymatic digestion of the chondroitin sulfate proteoglycans with chondroitinase ABC, which removes the glycosaminoglycan chains, navigation of retinal axons was disrupted. At embryonic day 13, before the uncrossed projection forms in normal development, many axons deviated from their normal course, crossing the midline at aberrant positions and invading the ventral diencephalon. In slices from embryonic day 14 embryos, axons that would normally form the uncrossed projection at this stage failed to turn into the ipsilateral optic tract. In embryonic day 15 slices, enzyme treatment caused a reduction of the uncrossed projection that develops at this stage. Growth cones in enzyme-treated slices showed a significant increase in the size both before and after they crossed the midline. This indicates that responses of retinal axons to guidance signals at the chiasm have changed after removal of the chondroitin sulfate epitope. We concluded that the chondroitin sulfate moieties of the proteoglycans are involved in patterning the early phase of axonal growth across the midline and at a later stage controlling the axon divergence at the chiasm.     

Invariance and selectivity in the ventral visual pathway.

Pattern recognition systems that are invariant to shape, pose, lighting and texture are never sufficiently selective; they suffer a high rate of "false alarms". How are biological vision systems both invariant and selective? Specifically, how are proper arrangements of sub-patterns distinguished from the chance arrangements that defeat selectivity in artificial systems? The answer may lie in the nonlinear dynamics that characterize complex and other invariant cell types: these cells are temporarily more receptive to some inputs than to others (functional connectivity). One consequence is that pairs of such cells with overlapping receptive fields will possess a related property that might be termed functional common input. Functional common input would induce high correlation exactly when there is a match in the sub-patterns appearing in the overlapping receptive fields. These correlations, possibly expressed as a partial and highly local synchrony, would preserve the selectivity otherwise lost to invariance.     

Calcium-binding proteins and cytochrome oxidase activity in the turtle optic tectum with special reference to the tectofugal visual pathway

Using immunohistochemistry and a tracer technique we investigated the distribution in the optic tectum of turtles (Emys orbicularis and Testudo horsfieldi) of the calcium-binding proteins (CaBPr) parvalbumin (PV), calbindin (CB) and calretinin (CR) before and after labeling of the nucleus rotundus (Rot) with horseradish peroxidase. The optic tectum activity of the cytochrome oxidase (CO) was studied in parallel. In the principal link of the tectofugal visual pathway (central gray layer, SGC) in both chelonian species, the sparse PV-ir as well as CB- and CR-ir neurons were found significantly varying both in number and the intensity of immunoreactivity of their bodies and dendrites. In contrast, the superficial (SGFS) and deeper periventricular (SGP) tectal layers comprised numerous cells immunoreactive to all three CaBPr in different proportions. Only few retrogradely labeled tectorotundal SGC neurons expressed PV, CB or CR. The very large PV-ir neurons in SGC and SAC were not retrogradely labeled; morphologically they matched the efferent neurons with descending projections. SGC neurons of two chelonian species differed in the level of CO activity. Intense immunoreactivity to all three CaBPr and high CO activity were detected in both species in SGFS neuropil with some differences in sublaminar distribution patterns. The peculiarities of the CaBPr and CO activity distribution patterns in different segments of SGC neurons are discussed as related to the laminar organization of the turtle tectum and its retinal innervation. It is suggested that in the projection tectorotundal SGC neurons the CaBPr are concentrated mainly in their distal dendrites that contact retinal afferents in the superficial retinorecipient tectal layer.     

Cysteine sulfinate carboxylase in the visual pathway of adult chicken.

Cysteine sulfinate (CSA) carboxylyase, the enzyme which synthesizes taurine through hypotaurine, shows a higher activity in the inner plexiform and nuclear layer of adult chick retina compared to the outer plexiform and nuclear layers whereas the outer segments of photoreceptors do not show any activity of this enzyme. These observations suggest an endogenous synthesis of taurine preferentially in certain layers of retina. Therefore, taurine fulfills one more criteria which is required by a substance to be accepted as a neurotransmitter in an organ. Studies on the distribution of CSA-carboxylyase in the visual pathway and other brain areas show a very high activity of this enzyme in optic tectum followed by cerebral cortex, cerebellum, retina, lateral geniculate body and optic nerve, taken with chiasma and tract in decreasing order. On the other hand, analysis of the free amino acid pool reveals a very high content of taurine in retina as compared to optic tectum. Cysteine sulfinate carboxylyase activity and the content of taurine therefore do not seem to bear a good correlation and other mechanisms of release, uptake and degradation might be involved in regulating the taurine content in these tissues.