The echidna, Tachyglossus aculeatus, a monotreme mammal, is thought to possess an all-rod retina (O' Day, 1952). This study provides anatomical evidence for the presence of cone-like photoreceptors in the retina of the echidna. The cones, which constitute 10–15% of the photoreceptors, have all of the ultrastructural characteristics previously shown in the cones of placental mammals, and, like the cones of other animals (Blanks & Johnson, 1984), they bind peanut agglutinin. Unlike the cones of another monotreme, the platypus, the cones of the echidna retina do not possess oil droplets. Twin cones, pairs of cones in which there is no obvious difference in the size, shape, or ultrastructural features of the members of a pair, are common. The density of cones varies from 9000 cells/mm2 in the superior periphery to 22,000 cells/mm2 in the central retina. Nearest-neighbor analysis suggests that the cone mosaic in the echidna retina results from the presence of single and twin cones in a relatively regular array.

We have compared the responses of simple cells to laterally moving sinusoidal gratings and to stationary temporally-modulated gratings. From the amplitudes and temporal phases of the responses to stationary gratings of different spatial phases, it should be possible to predict the preferred direction of movement, the amplitudes of the responses to gratings moving in the preferred and nonpreferred directions and, thence, the degree of directional preference (Reid et al., 1987). The preferred direction can be predicted reliably. However, the magnitude of the directional preference cannot be predicted, since the measured amplitude of the response in the nonpreferred direction of movement is very much less than that predicted by a linear theory. Nonlinearities in the relationship between response amplitude and contrast may contribute to the failure of the predictions, but this contribution is small. We conclude that the magnitude of the directional preference seems to be determined predominantly by nonlinear suppression of the response in the nonpreferred direction of movement.

In the retina of a variety of vertebrate species, a monoclonal antibody against protein kinase C (PKC) has been shown to label preferentially bipolar cells. Although the functional consequences of PKC activation in these cells is yet to be revealed, the present study was motivated in part by the possibility that the antibody might be used as a selective marker for examining the development of bipolar cells in the rat retina. Here, the developmental pattern and the dynamic changes of retinal cells expressing PKC-like immunoreactivity (PKC-LI) were studied and analyzed throughout postnatal life until adulthood. Upon its initial detection by immunohistochemistry on postnatal day (PD)-10, faint PKC-LI was limited to the central region of the retina, labeling cell bodies located at the scleral margin of the inner nuclear layer (INL) adjacent to the outer plexiform layer (OPL). On subsequent days, PKC-LI spread progressively to the peripheral retina and axon terminal bulbs at the vitreal margin of theinner plexiform layer (IPL) began showing the first signs of immunoreactive labeling. Not until PD-15, the time of eye opening, did PKC-LI in these cells increase to the extent such that their thin axons were immunoreactive. Each of these axons traversed the entire thickness of the IPL and divided into two or three short branches before ending as enlarged terminal bulbs. The morphology and the location of PKC-LI cells in both the developing and adult retina observed in our study are consistent with them being rod bipolar cells. By the end of the fourth postnatal week, the rod bipolar cells appeared mature, resembling those found in the adult. Overall, more dynamic changes occurred at the axon terminal bulbs than at the cell bodies during the maturational process of rod bipolar cells. Interestingly, PKIC-LI was expressed precociusly in these cells when rat pups were reared in complete darkness starting from the day of birth.

Ganglion cell activity in response to drifting or alternating stimulus patterns was recorded in an eyecup preparation of the turtle. The stimuli were sinusoidal gratings with various spatial frequencies and spatial phases. Spike histograms of the activity were Fourier-analyzed. The results indicated that the retinal circuitry always functioned in a nonlinear manner. The average, first, and second harmonics of the activity were all a function of stimulus conditions, including spatial phase. Similar outcomes were obtained with full-field (retina-wide) and a small-field (1 mm diameter) stimulation. In the middle of the spatial-frequency range, the ratio of second to first harmonic activity was higher in response to static, contrast-reversing gratings than for drifting, constant-contrast ones. The findings suggest that, as distinct from the usual X, Y, and W segregation of ganglion cell properties in other retinas, there is a single class of cells whose responses seem to have attributes of one or another of the usual categories depending on stimulus characteristics. This may be the result of a degree of stimulus-dependent functional modification in the retinal input circuitry to the turtle ganglion cells.

Two morphologically distinct types of preganglionic endings are observed in the avian ciliary ganglion: boutonal and cap-like. Boutonal endings synapse on ciliary ganglion neurons (called choroidal neurons) innervating choroidal blood vessels, while cap-like endings synapse on ciliary ganglion neurons (called ciliary neurons) controlling the lens and pupil. Some of both types of preganglionic endings contain the neuropeptides substance P (SP) and/or leucine-enkephalin (LENK). Although both types of preganglionic terminals are also known to be cholinergic, there has been no direct evidence that SP and LENK are found in cholinergic endings in the ciliary ganglion. The present studies in pigeons, which involved the use of single- and double-label immunohistochemical techniques, were undertaken to examine this issue, as well as to (1) determine the relative percentages of the boutonal and cap-like endings that contain SP, LENK, or both SP and LENK; and (2) determine if the two different types of terminals in the ciliary ganglion arise from different subdivisions of the nucleus of Edinger-Westphal (EW).

Single- and double-label immunohistochemical studies revealed that all neurons of EW, regardless of whether they contained immunohistochemically detectible amounts of SP or LENK, are cholinergic. In the medial subdivision of EW (EWM), which was found to contain approximately 700 neurons, 20.2% of these neurons were observed to contain both SP and LENK, while 11.6% were observed to contain SP only and 10.7% were observed to contain LENK only. In contrast, in lateral EW (EWL), which was found to contain approximately 500 neurons, 16.2% of the neurons were observed to contain both SP and LENK, while 19.2% of the neurons were observed to contain SP only and 12.6% were observed to contain LENK only. Retrograde-labeling studies involving horseradish peroxidase injections into the ciliary ganglion revealed that EW was the sole source of input to the ciliary ganglion and all, or nearly all, neurons in EW innervate the ciliary ganglion.

Immunohistochemical labeling of the ciliary ganglion neurons with an antiserum against choline acetyltransferase revealed that approximately 900 choroidal neurons and approximately 600 ciliary neurons are present in the ganglion, all of which receive cholinergic preganglionic endings. Of the choroidal neurons, 94% receive butonal terminals containing both SP and LENK, while only 2% receive SP+ only boutonal endings and 2% receive LENK+ only butonal endings. Of the ciliary neurons, 25% receive cap-like endings containing both SP and LENK, 30% receive cap-like endings containing only SP and 3% receive cap-like endings containing only LENK. Total unilateral lesions of EW resulted in the loss of all SP+ or LENK+ terminals in the ipsilateral ganglion. Subtotal EW lesions that spared either part of EWM or part of EWL revealed that boutonal endings arise from EWM neurons and cap-like endings from EWL neurons.

The present results suggest that the choroidal neurons, which regulate choroidal blood flow, may be relatively uniform in their functional properties since they nearly all receive boutonal endings from EWM that co-contain SP, LENK, and acetylcholine. In contrast, the ciliary neurons, which receive their preganglionic input from EWL, may consist of at least three major functionally distinct subgroups: (1) those receiving SP/LENK/acetylcholine-containing cap-like endings; (2) those receiving SP/acetylcholine-containing cap-like endings; and (3) those receiving acetylcholine-containing cap-like endings. The functional diversity of ciliary neurons may in part be related to the fact that some ciliary neurons innervate the iris and others the ciliary body.

Larvae of the tobacco hornworm moth Manduca sexta were reared on either a carotenoid-supplemented or a carotenoid-deficient diet. The former yields fortified adults with normal visual function, whereas visual sensitivity and rhodopsin content are reduced by 2−4 log units in the compound eyes of the deprived moths reared on the latter. We characterized the retinoids of fortified retinas and investigated the recovery of visual function in deprived moths that were provided with retinaldehyde as a source of photopigment chromophore. Retinoids were identified and measured by high-performance liquid chromatography (HPLC). Fortified retinas contained mainly 3-hydroxyretinaldehyde (R3); 11-cis R3 predominated in dark-adaptation, all-trans in light-adaptation, indicating that R3 is the photopigment chromophore. No retinoids could be measured in deprived eyes. Retinaldehyde (R1) was delivered to the retinas of deprived moths by “painting” solutions of 11-cis or all-trans R1 in dimethylsulfoxide (DMSO) on the corneal surfaces of the compound eyes or on the head capsule between the eyes. 11-cis R1 induced rapid recovery: during 3 days, sensitivity rose to within a log unit of that measured from fortified animals. By 7 days, sensitivity was close to normal. Although rhodopsin and P-face particle densities of photoreceptor membranes increased, neither rose to the levels found in fortified animals. All-trans R1 induced only a slight increase in sensitivity that could have resulted from some nonspecific isomerization of the all-trans to the 11-cis isomer; we found no evidence for a retinal isomerase that functions in darkness. Small amounts of R3 were measured in recovering retinas, indicating some conversion of R1 to R3. However, the chromophore of most of the rhodopsin that was synthesized must have been R1. It is possible that rhodopsin did not reach normal levels in the retina even after a week of recovery because the normal chromophore R3 was not provided. Although the rhodopsin that initially formed in recovering moths may have resulted from the association of the chromophore with pre-existing opsin, the extent of eventual recovery indicates that opsin synthesis was stimulated by 11-cis R1.

Circadian rhythms affect the anatomy, physiology, and biochemistry of the visual cells in the eyes of the horseshoe crab (Limulus polyphemus). These rhythms are mediated by the activity of efferent neurons that project from the central nervous system to all of the eyes. In this study, the optic nerves of Limulus were backfilled with Neurobiotin revealing the location of efferent cell bodies and their projections through the central nervous system. We propose that this efferent system mediates the circadian changes in visual functions in Limulus. Whether these cells are the circadian pacemaker neurons is unknown.

The cell bodies of the efferent neurons are ovoid and have a diameter of 40−80 μm. They lie within the cheliceral ganglion of the tritocerebrum, just posterior to the protocerebrum. This ganglion is on the lateral edge of the circumesophageal ring, near the middle of the dorsal-ventral axis of the ring. Each optic nerve contains axons from both ipsilateral and contralateral efferent cells, and some, possibly all, of them project bilaterally and to more than one type of optic nerve.

The efferent axons form a tract that projects anteriorly from the cell bodies to the protocerebrum, and bifurcates just lateral to the protocerebral bridge. One branch crosses the midline and projects anteriorly to the optic tract and medulla on the side contralateral to the cell of origin; the other branch follows a symmetric pathway on the ipsilateral side. Small branches arising from the major efferent axons in the optic tract project through the ocellar ganglia to the median optic nerves. The efferent axons branch again in the medulla, and some of these branches innervate the ventral optic nerves. The major branches of the efferent axons continue through the lamina and enter the lateral optic nerve.

Pigeons were trained on four different visual discrimination tasks: (1) concept of natural stimuli (food vs. non-food object discrimination); (2) arbitrary classification of natural stimuli (pseudoconcept); (3) concept of artificial stimuli (triangles generated by computer graphics); and (4) discrimination of one pair of artificial stimuli. Then, lesions of the ectostriatum were carried out. The ectostriatal lesions impaired the arbitrary classification of natural stimuli and the concept of artificial pattern but did not impair the natural concept or the simple discrimination of fixed two stimuli. Lesions in the neostriatum did not cause deficits in any discrimination task. The birds had to learn individual stimuli for the arbitrary classification of stimuli and the stimulus generalization test after the artificial pattern concept discrimination indicated that the pigeons formed a concept more complicated than “triangle” in human language. These results suggest that the ectostriatum plays a role in task discrimination that requires much visual processing to classify stimuli.

The effects of a variable-phase synaptic modification rule on the experience-dependent development of a simulated visual cortex were investigated. Of interest were the process of optimization of the internal representation with respect to orientation, through which the weakly tuned neurons of visually inexperienced animals attain their tightly tuned adult characteristics, and the process of association by which identical stimuli presented to either eye come to evoke identical cortical representations. In its general form, the synaptic modification rule was Hebbian. However, it was not assumed that positive correlation of presynaptic and postsynaptic activity would lead to an increase in synaptic weight. The relative phase of presynaptic vs. postsynaptic activity that would effect an increase in synaptic weight was a parameter of the modification rule. When this parameter was zero, synaptic modification conformed to the standard Hebbian type. With a value of 180 deg, or the reverse-Hebb condition, negative correlation between presynaptic and postsynaptic activity led to increased synaptic weight. It was found that a synaptic modification rule of the reverse-Hebb type not only optimized the cortical representation, and associated the representations from the two eyes, but was quite stable with respect to retaining the optimized state for long periods of learning.

When negotiating a narrow gap, honeybees tend to fly through the middle of the gap, balancing the distances to the boundary on either side. To investigate the basis of this “centering response,” bees were trained to fly through a tunnel on their way to a feeding site and back, while their flight trajectories were filmed from above. The wall on either side carried a visual pattern. When the patterns were stationary vertical gratings, bees tended to fly through the middle of the tunnel, i.e. along its longitudinal axis. However, when one of the gratings was in motion, bees flying in the same direction as the moving grating tended to fly closer to while bees flying in the opposite direction tended to fly closer to the stationary grating. This demonstrates, directly and unequivocally, that flying bees estimate the distances of surfaces in terms of the apparent motion of their images. A series of further experiments revealed that the distance to the gratings is gauged in terms of their apparent angular speeds, and that the visual system of the bee is capable of measuring angular speed largely independently of the spatial period, intensity profile, or contrast of the grating. Thus, the motion-sensitive mechanisms mediating range perception appear to be qualitatively different from those that mediate the well-known optomotor response in insects, or those involved in motion detection and ocular tracking in man.

We have examined the time course of branch formation in neurites of retinal ganglion cells isolated from adult goldfish (Carassius auratus). These neurites elongate at approximately 13 μm/h, and usually branch by bifurcation of growth cones at their tips. The times elapsed between branchings in different neurites of single cells can be described by a Poisson distribution with a mean interval of approximately 2 h. As predicted by this distribution, a relatively large number of branchings occur simultaneously in different neurites of individual cells. Simultaneous branchings of neurites elongating at a common rate generate branch points that lay equidistant from their soma. Since similar branching patterns can be seen in dendrites of retinal ganglion and amacrine cells in situ, these results are consistent with the possibility that dendrites of individual neurons branch synchronously and grow at common rates during development.