Inhibition in the eye of Limulus

In the compound lateral eye of Limulus each ommatidium functions as a single receptor unit in the discharge of impulses in the optic nerve. Impulses originate in the eccentric cell of each ommatidium and are conducted in its axon, which runs without interruption through an extensive plexus of nerve fibers to become a fiber of the optic nerve. The plexus makes interconnections among the ommatidia, but its exact organization is not understood. The ability of an ommatidium to discharge impulses in the axon of its eccentric cell is reduced by illumination of other ommatidia in its neighborhood: the threshold to light is raised, the number of impulses discharged in response to a suprathreshold flash of light is diminished, and the frequency with which impulses are discharged during steady illumination is decreased. Also, the activity that can be elicited under certain conditions when an ommatidium is in darkness can be inhibited similarly. There is no evidence for the spread of excitatory influences in the eye of Limulus. The inhibitory influence exerted upon an ommatidium that is discharging impulses at a steady rate begins, shortly after the onset of the illumination on neighboring ommatidia, with a sudden deep minimum in the frequency of discharge. After partial recovery, the frequency is maintained at a depressed level until the illumination on the neighboring receptors is turned off, following which there is prompt, though not instantaneous recovery to the original frequency. The inhibition is exerted directly upon the sensitive structure within the ommatidium: it has been observed when the impulses were recorded by a microelectrode thrust into an ommatidium, as well as when they were recorded more proximally in single fibers dissected from the optic nerve. Receptor units of the eye often inhibit one another mutually. This has been observed by recording the activity of two optic nerve fibers simultaneously. The mediation of the inhibitory influence appears to depend upon the integrity of nervous interconnections in the plexus: cutting the lateral connections to an ommatidium abolishes the inhibition exerted upon it. The nature of the influence that is mediated by the plexus and the mechanism whereby it exerts its inhibitory action on the receptor units are not known. The depression of the frequency of the discharge of nerve impulses from an ommatidium increases approximately linearly with the logarithm of the intensity of illumination on receptors in its vicinity. Inhibition of the discharge from an ommatidium is greater the larger the area of the eye illuminated in its vicinity. However, equal increments of area become less effective as the total area is increased. The response of an ommatidium is most effectively inhibited by the illumination of ommatidia that are close to it; the effectiveness diminishes with increasing distance, but may extend for several millimeters. Illumination of a fixed region of the eye at constant intensity produces a depression of the frequency of discharge of impulses from a nearby ommatidium that is approximately constant, irrespective of the level of excitation of the ommatidium. The inhibitory interaction in the eye of Limulus is an integrative process that is important in determining the patterns of nervous activity in the visual system. It is analogous to the inhibitory component of the interaction that takes place in the vertebrate retina. Inhibitory interaction results in the exaggeration of differences in sensory activity from different regions of the eye illuminated at different intensities, thus enhancing visual contrast.     

Inhibitory interaction of receptor units in the eye of Limulus

The inhibition that is exerted mutually among the receptor units (ommatidia) in the lateral eye of Limulus has been analyzed by recording oscillographically the discharge of nerve impulses in single optic nerve fibers. The discharges from two ommatidia were recorded simultaneously by connecting the bundles containing their optic nerve fibers to separate amplifiers and recording systems. Ommatidia were chosen that were separated by no more than a few millimeters in the eye; they were illuminated independently by separate optical systems. The frequency of the maintained discharge of impulses from each of two ommatidia illuminated steadily is lower when both are illuminated together than when each is illuminated by itself. When only two ommatidia are illuminated, the magnitude of the inhibition of each one depends only on the degree of activity of the other; the activity of each, in turn, is the resultant of the excitation from its respective light stimulus and the inhibition exerted on it by the other. When additional receptors are illuminated in the vicinity of an interacting pair too far from one ommatidium to affect it directly, but near enough to the second to inhibit it, the frequency of discharge of the first increases as it is partially released from the inhibition exerted on it by the second (disinhibition). Disinhibition simulates facilitation; it is an example of indirect effects of interaction taking place over greater distances in the eye than are covered by direct inhibitory interconnections. When only two interacting ommatidia are illuminated, the inhibition exerted on each (decrease of its frequency of discharge) is a linear function of the degree of activity (frequency of discharge) of the other. Below a certain frequency (often different for different receptors) no inhibition is exerted by a receptor. Above this threshold, the rate of increase of inhibition of one receptor with increasing frequency of discharge of the other is constant, and may be at least as high as 0.2 impulse inhibited in one receptor per impulse discharged by the other. For a given pair of interacting receptors, the inhibitory coefficients are not always the same in the two directions of action. The responses to steady illumination of two receptor units that inhibit each other mutually are described quantitatively by two simultaneous linear equations that express concisely all the features discussed above. These equations may be extended and their number supplemented to describe the responses of more than two interacting elements.     

Inhibitory Fields in the Limulus Lateral Eye

The inhibition that is exerted mutually among receptor units (ommatidia) of the lateral eye of Limulus does not diminish uniformly with increasing distance between units. Instead the response of a receptor unit is most effectively inhibited by other units separated from it by approximately 1 mm (three to five receptor diameters); the effectiveness diminishes with distances both greater and less than this value. The ommatidial inhibitory field as measured by the spatial function of the inhibitory coefficients contains a uniform depression in the central region, a uniformly high annulus at some distance from the center, and a gradual tapering off toward the periphery. The field is large—covering over 30 % of the retina—and is somewhat elliptical in shape with its major axis in the anteroposterior direction on the lateral eye. A number of experiments reveal similar configurations in a sizable part of the eye. Control experiments show that the diminution of the inhibitory effects near the center of the field is not an artifact of the measuring technique and cannot be explained readily by local neural excitatory processes.     

Inhibition in the Limulus lateral eye in situ

Inhibition in the Limulus lateral eye in situ is qualitatively similar to that in the excised eye. In both preparations ommatidia mutually inhibit one another, and the magnitude of the inhibitory effects are linear functions of the response rate of individual ommatidia. The strength of inhibition exerted between single ommatidia is also about the same for both preparations; however, stronger effects can converge on a single ommatidium in situ. At high levels of illumination of the retina in situ the inhibitory effects are often strong enough to produce sustained oscillations in the discharge of optic nerve fibers. The weaker inhibitory influences at low levels of illumination do not produce oscillations but decrease the variance of the optic nerve discharge. Thresholds for the inhibitory effects appear to be determined by both presynaptic and postsynaptic cellular processes. Our results are consistent with the idea that a single ommatidium can be inhibited by more of its neighbors in an eye in situ than in an excised eye. Leaving intact the blood supply to the eye appears to preserve the functional integrity of the retinal pathways which mediate inhibition.     

Spatial summation of inhibitory influences in the eye of Limulus,and the mutual interaction of receptor units

The inhibitory influences exerted mutually among the receptor units (ommatidia) of the lateral eye of Limulus are additive. If two groups of receptors are illuminated together the total inhibition they exert on a "test receptor" near them (decrease in the frequency of its nerve impulse discharge in response to light) depends on the combined inhibitory influences exerted by the two groups. If the two groups are widely separated in the eye, their total inhibitory effect on the test receptor equals the sum of the inhibitory effects they each produce separately. If they are close enough together to interact, their effect when acting together is usually less than the sum of their separate effects, since each group inhibits the activity of the other and hence reduces its inhibitory influence. However, the test receptor, or a small group illuminated with it, may interact with the two groups and affect the net inhibitory action. A variety of quantitative effects have been observed for different configurations of three such groups of receptors. The activity of a population of n interacting elements is described by a set of n simultaneous equations, linear in the frequencies of the receptor elements involved. Applied to three interacting receptors or receptor groups equations are derived that account quantitatively for the variety of effects observed in the various experimental configurations of retinal illumination used.     

Die Verarbeitung stationärer optischer Nachrichten im Komplexauge von Limulus

Summary The functional properties of the processing of visual information by the complex eye of Limulus was studied. The spatial distribution of activity that results in the optic nerve when the Limulus eye is exposed to a stationary optical pattern depends upon the transfer characteristics of two subsystems: the dioptric apparatus and the nervous interactions comprising the lateral inhibition system. — The transfer characteristic of the dioptric apparatus is determined by the sensitivity distribution function of single ommatidia. This distribution was measured and found to be approximately of Gauss-function type. The sensitivity falls off to 1/e at a distance of one ommatidium; thus the visual fields of adjacent ommatidia strongly overlap. As a consequence of the overlap, amplitudes of the spatial Fourier components, of which the brightness distribution of the optical surround is made up, are more and more reduced with increasing frequency in the intensity distribution on the receptor mosaic. The amplitude of the spatial frequency 1/=0,25 ( in units of interommatidial distance) is reduced to half of the maximum value, which is attained at zero frequency. It is shown that the amplitude frequency characteristic of the sensitivity distribution function has no zeros, which means that no loss of optical information results from overlap of visual fields. Thus the resolving power of the dioptric apparatus is limited only by the number of receptors per unit area. — The transfer characteristic of the lateral inhibition system in the Limulus eye depends on the distribution of the inhibitory coefficients around the individual receptors. This distribution function was determined from excitatory responses in the optic nerve elicited by a spatial light intensity step function on the receptor mosaic. It is found that this distribution is also Gaussian in form, but decays to 1/e at a distance of eight to nine ommatidia along the major axis of the eye. The average value of the inhibitory coefficients between adjacent ommatidia was found to be 0,025. The amplitude frequency response of the inhibitory system is constant for high spatial frequencies down to 1/=0,1 while amplitudes of lower frequency sinusoids are reduced down to nearly half of the maximum value at frequency zero. The amplitude frequency characteristic of the inhibitory system ensures a one to one correspondence between the intensity distribution on the receptor mosaic and the excitation distribution in the optic nerve. The overall transfer characteristic of the eye is derived from the transfer characteristics of the dioptric apparatus and the inhibitory system. This characteristic is of bandpass type with a maximum amplitude response at a frequency of 1/=0,07. The overall transfer characteristic was independently confirmed in a separate experiment. The nature of the overall transfer characteristic shows that the inhibitory system does not exactly correct for the overlap of the visual fields of single ommatidia, which in principal the system could do if the distributions of inhibitory coefficients and ommatidia sensitivity were equal. The overall transfer characteristic of the Limulus eye garantees a one to one correspondence between patterns in the optical surround and excitation distributions in the optic nerve. — The average values of the inhibitory coefficients derived from these experiments are at least a factor ten smaller than those determined directly by other investigators. Possible explanations of this discrepency are discussed. — In a separate chapter the overall transfer characteristic for eyes submerged in water is described. It was found that this characteristic does not differ from that determined in air for the eye region which was investigated in the experiments. This result is explained by two properties of the eye which are dependent on the refractive index of the surround medium and whose influences cancel each other: the visual fields of ommatidia are reduced under water, while the divergence angles between the optical axes of adjacent ommatidia also diminish.

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This research was supported in part by the United States Air Force under Grant No. AF-EOAR-62-41 and monitored by the European Office, Office of Aerospace Research.     

Serotonin and Inhibition in Limulus Lateral Eye

The response to light of one ommatidium is reduced or suppressed by simultaneous illumination of neighboring ommatidia. The mechanism of this lateral inhibition may be chemical synaptic transmission, based on the physiological findings of a number of investigators and on the following evidence. The fine structure of the neuropil of the lateral plexus exhibits numerous clear vesicles (ca. 400 A), dense-core vesicles (ca. 700–1400 A), Golgi regions, and other morphological features of neurochemical synapses. The indolealkylamine, serotonin (5-HT), even in nanomolar concentrations, has a potent inhibitory action. An initial, potent inhibitory dose of 5-HT produces a long lasting densensitization to subsequent doses. The desensitization affects lateral inhibition evoked by light stimulation of neighboring receptors, i.e. crossed-desensitization. Eye tissue extracts contain 5-HT and melatonin (MLT) at a level greater than 1 µg/g wet tissue and perhaps as high as 20–30 µg/g, as determined by two-dimensional thin-layer chromatography (TLC) and o-phthaldialdehyde fluorescence assay techniques. Subcellular fractionation on sucrose gradient indicates a peak in 5-HT and MLT content associated with an intermediate density fraction. 5-HT may be an inhibitory transmitter for lateral inhibition. One pathway for metabolism of 5-HT in the lateral eye may be via N-acetylserotonin and melatonin.     

The retina-lamina projection in the crab Leptograpsus variegatus

Summary In the crab, Leptograpsus variegatus, the projection of retinula cell axons to the lamina was investigated by tracing them through a series of semi-thin sections. Forty-four such axons were traced from a single group of ommatidia as far as the distal layers of the lamina. The eight receptor axons of one ommatidium project to a single lamina cartridge. Therefore, because the crab has a fused rhabdom, angular information is conserved in vision, and the outside world is projected literally onto the lamina, just as it is in the standard non-dipteran pattern of insects. The belief of previous workers that other decapod eyes show neural superposition was an inference based primarily on the patterns of penetration of the basement membrane by receptor axons, and on degeneration experiments. This evidence is reviewed, shown to be inadequate and discussed in the light of the projection now demonstrated for Leptograpsus.     

Colour receptors in the eye of the digger wasp,Sphex cognatus Smith: Evaluation by selective adaptation

Summary Pigment granule migration in pigment cells and retinula cells of the digger wasp Sphex cognatus Smith was analysed morphologically after light adaptation to natural light, dark adaptation and after four selective chromatic adaptations in the range between 358 nm and 580 nm and used as the index of receptor cell sensitivity. The receptor region of each ommatidium consists of nine retinula cells which form a centrally located rhabdom. Two morphologically and physiologically different visual units can be described, defined by the arrangement of the rhabdomeric microvilli, the topographical relationship of the receptor cells with respect to the eye axes and the unique retinula cell screening pigmentation. These two different sets of ommatidia (type A and B) are randomly distributed in a ratio of 13 throughout the eye (Ribi, 1978b). Chromatic adaptation experiments with wavelengths of 358 nm, 443 nm, 523 nm and 580 nm and subsequent histological examination reveal two UV receptors, two blue receptors and four yellow-green receptors in type A ommatidia and two UV receptors and six green to yellow-green receptors in type B ommatidia. The pigments in cells surrounding each ommatidium (two primary pigment cells, 20 secondary pigment cells and four pigmented cone extensions) were not affected significantly by the adaptation experiments.     

Preferential adhesion maintains separation of ommatidia in the Drosophila eye

In the Drosophila eye, neighboring ommatidia are separated by inter-ommatidial cells (IOCs). How this ommatidial spacing emerges during eye development is not clear. Here we demonstrate that four adhesion molecules of the Irre cell recognition module (IRM) family play a redundant role in maintaining separation of ommatidia. The four IRM proteins are divided into two groups: Kirre and Rst are expressed in IOCs, and Hbs and Sns in primary pigment cells (1°s). Kirre binds Hbs and Sns in vivo and in vitro. Reducing activity of either Rst or Kirre alone had minimal effects on ommatidial spacing, but reducing both together led to direct ommatidium:ommatidium contact. A similar phenotype was also observed when reducing both Hbs and Sns. Consistent with the role of these factors in sorting ommatidia, mis-expression of Hbs plus Sns within a single IOC led to complete separation of the cell from neighboring ommatidia. Our results indicate mutual preferential adhesion between ommatidia and IOCs mediated by four IRM proteins is both necessary and sufficient to maintain separation of ommatidia.     

Localization of visual pigments within rhabdoms of the compound eye of Spodoptera exempta (Insecta,Noctuidae)

Summary In the noctuid moth Spodoptera exempta, the distribution of visual pigments within the fused rhabdoms of the compound eyes was investigated by electron microscopy. Each ommatidium regularly contains eight receptor cells belonging to three morphological types: one distal, six medial, and one basal cell (Meinecke 1981); four different visual pigments — absorption maxima at approximately 355, 465, 515, and 560 nm — are known to occur within the eye (Langer et al. 1979). The compound eyes were illuminated in situ by use of monochromatic light of different wavelengths. This illumination produced a wide scale of structural changes in the microvilli of the rhabdomeres of individual cells. Preparation of eyes by freeze-substitution revealed the structural changes in the rhabdomeres to be effects of light occurring in vivo.The degree of structural changes may be considerably different in rhabdomeres within the same ommatidium; it was found to depend on the wavelength and the duration of illumination, the intensity received by the ommatidia as well as the spectral sensitivity of the receptor cells. Therefore, it was possible to estimate the spectral sensitivities of the morphological types of receptor cells. Generally, all medial cells are green receptors and all basal cells red receptors; distal cells are blue receptors in about two-thirds of the ommatidia, while in the remaining third of them distal cells are sensitive to ultraviolet light.Supported by Deutsche Forschungsgemeinschaft, Sonderforschungsbereich 114 (Bionach)     

Electrophysiological basis for the spatial dependence of the inhibitory coupling in the Limulus retina

A technique for measuring, with total optical isolation, the inhibition between two individual receptor units in the Limulus lateral eye is described. The extracellular responses of pairs of units were recorded, using light piping microelectrodes. The inhibitory coupling between two units was found to be nonlinear and describable by a simple hyperbolic equation written in terms of saturation rate (S), half saturation (H), and threshold (ft). By plotting reciprocal frequencies, the data could be linearized and compared for different pairs of units. The magnitude of inhibition (in terms of S and H) was found to decrease monotonically as the anatomical distance between receptors increased. An electrical model of the inhibitory system was developed which accounts for many of the properties of the observed inhibitory interactions. Using the equations from the model and the experimental data, it is shown that the "electrical distances" (which are computed in terms of space constants lambda) of the inhibitory synapses from the impulse-generating region of the test unit are directly related to the anatomical distance between receptors. It is also shown that "synaptic strength" is relatively constant with separation. The electrical distances of the inhibitory synapses range from about 0.1lambda to 0.25lambda for adjacent units to greater than 0.5lambda for units seven to nine receptors away. It is concluded that the nonlinear character of the inhibitory coupling is attributable to synaptic effects, and that the decrease of inhibition with distance between receptors is caused primarily by an increase in the electrical distance of the inhibitory synapses from the test unit.     

Ultrastructure within the Lateral Plexus of the Limulus Eye

The ultrastructure of the lateral plexus in the compound eye of Limulus is investigated by serial section technique. "Cores" of tissue containing the axons, lateral plexus, and neuropile associated with one sensory ommatidium show the following features: (a) collateral branches from retinular cells do not contribute to the lateral plexus proper, but do form retinular neuropile by contacting collaterals of a self-contained cluster of retinular axons; (b) collateral branches from eccentric cell axons always branch repeatedly upon leaving the parent axon, and compose the bulk of the lateral plexus; (c) the most distal collateral branches from an eccentric cell axon appear to form neuropile and synaptic contacts with each other, whereas more proximal branches form synaptic contacts with collaterals from eccentric cell axons of neighboring ommatidia. We conclude that the ribbon synapses and associated transmitter substance in eccentric cell collaterals must be inhibitory, and that two pathways for self-inhibition may exist. We suggest, as a working hypothesis for the structure of the lateral plexus, a branching pattern with depth that mirrors the horizontal spread of lateral inhibition measured physiologically.     

Functional morphology of the ommatidia in the compound eye of the moth,Antheraea polyphemus (Insecta,Saturniidae)

Summary The fine structure of the superposition eye of the Saturniid moth Antheraea polyphemus Cramer was investigated by electron microscopy. Each of the approximately 10000 ommatidia consists of the same structural components, but regarding the arrangement of the ommatidia and the rhabdom structure therein, two regions of the eye have to be distinguished. In a small dorsal rim area, the ommatidia are characterized by rectangularly shaped rhabdoms containing parallel microvilli arranged in groups that are oriented perpendicular to each other. In all other ommatidia, the proximal parts of the rhabdoms show radially arranged microvilli, whereas the distal parts may reveal different patterns, frequently with microvilli in two directions or sometimes even in one direction. Moreover, the microvilli of all distal cells are arranged in parallel to meridians of the eyes. By virtue of these structural features the eyes should enable this moth not only discrimination of the plane of polarized light but also skylight-orientation via the polarization pattern, depending on moon position. The receptor cells exhibit only small alterations during daylight within the natural diurnal cycle. However, under illumination with different monochromatic lights of physiological intensity, receptor cells can be unbalanced: Changes in ultrastructure of the rhabdomeres and the cytoplasm of such cells are evident. The effects are different in the daytime and at night. These findings are discussed in relation to the breakdown and regeneration of microvilli and the influence of the diurnal cycle. They are compared with results on photoreceptor membrane turnover in eyes of other arthropod species.     

The organization of perpendicular fibre pathways in the insect optic lobe.

High resolution serial photomicrography has been used to plot the axonal projection patterns between retina, lamina and medulla in the optic lobes of various insects with differing ommatidial receptor arrangements. Observations are reported on the cabbage white and skipper butterflies, the bee, locust, fly, backswimmer and waterbug. The patterns of these fibre pathways have previously eluded non-rigorous analyses primarily because of their physical dimensions but are revealed in this study to have striking precision and uniformity between species when examined at the level of individually identifiable cells. Axon bundles of the tracts between retina and lamina or lamina and medulla project between a single ommatidium and its corresponding lamina cartridge or between corresponding lamina and medulla cartridges. Lateral interweaving of axons between adjacent bundles is absent. The bundles preserve the retinotopic order within their total array, so transferring the pattern of retinulae directly upon the lamina and thence after horizontal inversion in the chiasma upon the medulla. Within the lamina neuropile on the other hand the trajectories of the individual terminals from a bundle have patterns which are species-specific, sometimes involving lateral divergences. In species with open-rhabdomere ommatidia the terminals distribute to a group of lamina cartidges with a pattern which resembles the receptor pattern in the overlying ommatidium. In species with fused-rhabdome ommatidia the terminals of a single retinula behave less interestingly and all enter the same cartridge, within which, again, each occupies a position related to its cell body position within the retinula. Long visual fibres in both eye types penetrate the lamina and terminate in the particular medulla cartridge that connects with the lamina cartridge underlying their ommatidium. The perpendicular fibre pathways therefore project the visual field exactly upon the medulla in all species while the lack of interweaving between adjacent fibre bundles precludes their involvement in lateral interactions between pathways with differing visual axes. Uniformity of these projection patterns between cell layers and species differences in retinular terminal locations in the lamina can be correlated with different modes of axon growth between and within neuropile layers during optic lobe neurogenesis. Further discussion surrounds the question of which particular receptors give rise to which type of axon, for which no clear generalization has yet emerged.     

Specialized ommatidia for polarization vision in the compound eye of cockchafers,Melolontha melolontha (Coleoptera,Scarabaeidae)

Summary The superposition eye of the cockchafer, Melolontha melolontha, exhibits the typical features of many nocturnal and crepuscular scarabaeid beetles: the dioptric apparatus of each ommatidium consists of a thick corneal lens with a strong inner convexity attached to a crystalline cone, that is surrounded by two primary and 9–11 secondary pigment cells. The clear zone contains the unpigmented extensions of the secondary pigment cells, which surround the cell bodies of seven retinula (receptor) cells per ommatidium and a retinular tract formed by them. The seven-lobed fused rhabdoms are composed by the rhabdomeres of the receptor cells 1–7. The rhabdoms are optically separated from each other by a tracheal sheath around the retinulae. The orientation of the microvilli diverges in a fan-like fashion within each rhabdomere. The proximally situated retinula cell 8 does not form a rhabdomere. This standard form of ommatidium stands in contrast to another type of ommatidium found in the dorsal rim area of the eye. The dorsal rim ommatidia are characterized by the following anatomical specializations: (1) The corneal lenses are not clear but contain light-scattering, bubble-like inclusions. (2) The rhabdom length is increased approximately by a factor of two. (3) The rhabdoms have unlobed shapes. (4) Within each rhabdomere the microvilli are parallel to each other. The microvilli of receptor 1 are oriented 90° to those of receptors 2–7. (5) The tracheal sheaths around the retinulae are missing. These findings indicate that the photoreceptors of the dorsal rim area are strongly polarization sensitive and have large visual fields. In the dorsal rim ommatidia of other insects, functionally similar anatomical specializations have been found. In these species, the dorsal rim area of the eye was demonstrated to be the eye region that is responsible for the detection of polarized light. We suggest that the dorsal rim area of the cockchafer eye subserves the same function and that the beetles use the polarization pattern of the sky for orientation during their migrations.     

Decoding vectorial information from a gradient: sequential roles of the receptors Frizzled and Notch in establishing planar polarity in the Drosophila eye

The Drosophila eye is composed of several hundred ommatidia that can exist in either of two chiral forms, depending on position: ommatidia in the dorsal half of the eye adopt one chiral form, whereas ommatidia in the ventral half adopt the other. Chirality appears to be specified by a polarizing signal with a high activity at the interface between the two halves (the 'equator'), which declines in opposite directions towards the dorsal and ventral poles. Here, using genetic mosaics, we show that this polarizing signal is decoded by the sequential use of two receptor systems. The first depends on the seven-transmembrane receptor Frizzled (Fz) and distinguishes between the two members of the R3/R4 pair of presumptive photoreceptor cells, predisposing the cell that is located closer to the equator and having higher Fz activity towards the R3 photoreceptor fate and the cell further away towards the R4 fate. This bias is then amplified by subsequent interactions between the two cells mediated by the receptor Notch (N) and its ligand Delta (Dl), ensuring that the equatorial cell becomes the R3 photoreceptor while the polar cell becomes the R4 photoreceptor. As a consequence of this reciprocal cell fate decision, the R4 cell moves asymmetrically relative to the R3 cell, initiating the appropriate chiral pattern of the remaining cells of the ommatidium.     

Thermal Sensitivity of Lateral Inhibition in Limulus Eye

The effectiveness of lateral inhibition, measured as spike response decrement in a test ommatidium, produced by activity in a group of neighboring ommatidia, decreases as temperature decreases (Q₁₀ of 2.6). The corresponding sensory transducer-spike encoding processes have a weaker temperature dependence (Q₁₀ of 1.6). Relative synaptic delay, the time difference between the latency of inhibition onset and the latency of test receptor excitation, has a strong temperature dependence (Q₁₀ of 5), while receptor potential onset latency (Q₁₀ of 1.4) and optic nerve spike conduction velocity (Q₁₀ of 1.7), two factors inherent in relative synaptic delay, are less temperature sensitive. Oscillations of optic nerve spike response ("bursting") may be produced by thermal adjustment of temperature-sensitive parameters of the lateral inhibitory network in the retina. Burst interval has a strong temperature dependence (Q₁₀ of 2.4) and broad interspike interval distribution compared to the thermal sensitivity (Q₁₀ of 1.4) and narrow spike interval spectrum of the response of a single unit within the bursting group.     

THE VISUAL ACUITY OF THE HONEY BEE

1. Bees respond by a characteristic reflex to a movement in their visual field. By confining the field to a series of parallel dark and luminous bars it is possible to determine the size of bar to which the bees respond under different conditions and in this way to measure the resolving power or visual acuity of the eye. The maximum visual acuity of the bee is lower than the lowest human visual acuity. Under similar, maximal conditions the fineness of resolution of the human eye is about 100 times that of the bee. 2. The eye of the bee is a mosaic composed of hexagonal pyramids of variable apical angle. The size of this angle determines the angular separation between adjacent ommatidia and therefore sets the structural limits to the resolving power of the eye. It is found that the visual angle corresponding to the maximum visual acuity as found experimentally is identical with the structural angular separation of adjacent ommatidia in the region of maximum density of ommatidia population. When this region of maximum ommatidia population is rendered non-functional by being covered with an opaque paint, the maximum visual acuity then corresponds to the angular separation of those remaining ommatidia which now constitute the maximum density of population. 3. The angular separation of adjacent ommatidia is much smaller in the vertical (dorso-ventral) axis than in the horizontal (anterio-posterior) axis. The experimentally found visual acuity varies correspondingly. From this and other experiments as well as from the shape of the eye itself, it is shown that the bee''s eye is essentially an instrument for uni-directional visual resolution, functional along the dorso-ventral axis. The resolution of the visual pattern is therefore determined by the vertical angular separation of those ocular elements situated in the region of maximum density of ommatidia population. 4. The visual acuity of the bee varies with the illumination in much the same way that it does for the human eye. It is low at low illuminations; as the intensity of illumination increases it increases at first slowly and then rapidly; and finally at high intensities it becomes constant. The resolving power of a structure like the bee''s eye depends on the distance which separates the discrete receiving elements. The data then mean that at low illuminations the distance between receiving elements is large and that this distance decreases as the illumination increases. Since such a moving system cannot be true anatomically it must be interpreted functionally. It is therefore proposed that the threshold of the various ommatidia are not the same but that they vary as any other characteristic of a population. The visual acuity will then depend on the distance apart of those elements whose thresholds are such that they are functional at the particular illumination under investigation. Taking due consideration of the angular separation of ommatidia it is possible to derive a distribution curve for the thresholds of the ommatidia which resembles the usual probability curves, and which describes the data with complete fidelity.     

Localization of the pupil trigger in insect superposition eyes

Summary In the superposition eyes of the sphingid moth Deilephila and the neuropteran Ascalaphus, adjustment to different intensities is subserved by longitudinal migrations of screening pigment in specialized pigment cells. Using ophthalmoscopic techniques we have localized the light-sensitive trigger that controls pigment position.In both species, local illumination of a small spot anywhere within the eye glow of a dark-adapted eye evokes local light adaptation in the ommatidia whose facets receive the light. Details of the response pattern demonstrate that a distal light-sensitive trigger is located axially in the ommatidium, just beneath the crystalline cone, and extends with less sensitivity deep into the clear zone. The distal trigger in Deilephila was shown to be predominantly UV sensitive, and a UV-absorbing structure, presumably the distal trigger, was observed near the proximal tip of the crystalline cone.In Ascalaphus we also found another trigger located more proximally, which causes local pigment reaction in the ommatidia whose rhabdoms are illuminated (the centre of the eye glow). The light-sensitive trigger for this response appears to be the rhabdom itself.