Daily changes in the compound eye of a beetle ( Macrogyrus )

1983 ◽  
Vol 217 (1208) ◽  
pp. 265-285 ◽  

(i) Graded index lenses in the cornea and the crystalline cone form the optical system in each ommatidium. (ii) By night the crystalline cone has a blunt ellipsoidal proximal end which contributes to the formation of a superposition image across the clear zone. By day the cone is a tapering point that is extended as a light guide through a dense layer of pigment. (iii) The action of extending the cone and moving the pigment towards the clear zone from between the cones occurs as the retinula-cell column contracts. (iv) Modelling of the ommatidial lens system shows how the superposition image is formed in the night eye, and suggests that axial rays are not well focused on the crystalline tract in the day eye. (v) All cells had peak sensitivity in the green near 552 nm. (vi) In the dark-adapted day eye, fields are ∆ ρ (acceptance angle) = 3.4–6.6°, narrowing to 2.8° minimum upon light adaptation. Sensitivity to a point source on axis is reduced during the day: the dark-adapted day eye requires 200 times more light to give the same response as the dark-adapted night eye. There is a further attenuation of about 100 upon light adaptation of the day eye. (vii) The superposition image of the night eye produces fields of width ∆ ρ = 12-15° at 50% sensitivity as recorded electrophysiologically, and therefore the image of a point source covers several rhabdoms. (viii) In recordings from single units in the night eye two bumps (effective photon captures) are counted when the intensity is such that one photon falls on the area of one facet, with parallel axial illumination at the peak of the spectral sensitivity, 552 nm. (ix) Marking of cells with Lucifer Yellow suggests that about four to six receptor units per ommatidium are involved, giving a sensitivity of eight to twelve bumps for the ommatidium at this intensity. (x) Locust apposition eyes, with facets twice the area of those in Macrogyrus eyes, give at best 0.5 bumps with the same intensity, so that the actual superposition gain is 32–48. (xi) All marked cells were of the proximal rhabdom layer; cells 1 and 8 have not been investigated.

(i) The dorsal eyes are sensitive to ultraviolet light, which is focused by the corneal lens into crystalline cones in the region where these taper progressively to columns across the clear zone. The action of these columns as light guides can be observed in fixed eyes embedded in polymerized resin. In life the light guide part of the column is surrounded by watery non-cellular haemolymph. (ii) Shadowing the eye surface with a thin wire (three facets wide) while recording from individual receptor units shows that ultraviolet light reaches each receptor by its own facet as in an apposition eye, and not, as in a superposition eye, by a group of many facets. (iii) As shown by the dye Lucifer Yellow injected from a microelectrode, the electrophysiological unit consists of all seven retinula cells in the rhabdom region. Consistent with this tight coupling of retinula cells there is no polarization sensitivity. The peak spectral sensitivity of all single units is at 345-365 nm in the ultraviolet. The acceptance angle is 2.0–2.5°. The sensitivity at the spectral peak to a point source on the optical axis of the unit is poor compared to that in other insects tested with the same equipment. (iv) The acceptance angles (∆ ρ ) in the dorsal eye are at the theoretical minimum for the facet diameter and wavelength from diffraction theory. Ultraviolet vision, therefore, has made possible a reduction in facet size but the interommatidial angle ∆ ϕ is greater than expected from the optimum sampling theory of the diffraction limited compound eye. In fact ∆ ρ ≈ ∆ ϕ ≈ 2°. (v) The dorsal eye is effectively a foveal region with greater sampling density and narrower receptive fields but less overlap of fields than the lateral eye. (vi) The square cones and yellow screening pigment strongly suggest that there is superposition by reflexion of yellow light that spreads between ommatidia across the clear zone. This yellow light might photoreisomerize the visual pigment. Attempts to prove this theory during the recording from single units have so far failed but no better function for the clear zone has been suggested.


The night flying scarabaeid beetle Anoplognathus provides an example of a dark-adapted clear-zone compound eye in which rays from a distant point source, entering by a large patch of facets, are imperfectly focused upon the receptor layer. The optical system of the eye was investigated by six methods, all of which give similar results: (1) ray tracing through structures of known refractive index, (2) measurement of visual fields of single receptors, (3) measurement of the divergence of eyeshine, and (4) of the optomotor response to stripes of decreasing width, and (5) by direct observation of distribution of light within the eye. Finally (6) anatomically there is no single plane upon which an image could be focused. In each ommatidium, beneath the thick cornea, with its short corneal cone, lies a non-homogeneous crystalline cone (range of r. i. 1.442-1.365) that is significant in partially focusing rays across the wide clear zone (340 μm) in the dark-adapted eye. On the proximal side of the clear zone the rhabdoms form 7-lobed columns, isolated from each other over half their length by a tracheal tapetum. In the light-adapted eye the cone cells extend to form a crystalline tract (70-90 μm long) which is sur­rounded by dense pigment, and the optical path across the clear zone is completed by retinula cell columns that are of higher density than the surrounding cells. Pigment movement upon adaptation takes about 10 min to complete. Dark adaptation can be induced only at night on account of a strong diurnal rhythm. Eyeshine can be seen in the dark-adapted eye so long as the distal pig­ment leaves free the tips of the crystalline cones. Eyeshine falls to 50% at an angle of 12° from the direction of a parallel beam shining on the eye, as is consistent with a partial focus in which the distribution of light on the receptor layer is 18°-24° wide at the 50% contour. This distribution was confirmed by direct examination of the inside of the eye and by measure­ment of receptor fields as follows. The mean acceptance angle for 13 light-adapted units was 12.57° ± 1.97° s. d. and that of 10 dark-adapted ones 20.3° ± 3.36° s. d. The sensi­tivity to a point source on axis is increased at least 1000 fold by dark adaptation. Rays traced through a scale drawing of the eye, with refractive index measured for each component, show how the eye as a whole comes to be partially focused, and predicts an acceptance angle of 12° in the light-adapted and 20°-24° in the dark-adapted eye. In optomotor experiments dark-adapted Anoplognathus does not respond to stripes narrower than 18° repeat period, but light-adapted beetles respond down to 10°. The optomotor experiments also show a 1000 fold increase in sensitivity when dark-adapted at night. The eye has poor acuity that goes with wide visual fields of its recep­tors, and this is surprising when other excellently focused clear zone eyes are known. A possible compensation for the poor acuity is that the aperture of the eye can be larger, so that sensitivity although only to large objects, is that much increased.


1973 ◽  
Vol 183 (1071) ◽  
pp. 159-178 ◽  

1. The compound eye of Cybister is anatomically similar to that of Dytiscus and Hydrophilus . 2. The cornea and crystalline cone in the compound eye of Cybister (Dytiscidae) are composed of layers of unequal refractive index. With the exception of the outer 10 µ m of the cornea (where they are horizontal) the layers are arranged concentrically around a region of highest refractive index on the axis. 3. The refractive index of the cornea decreases from the central layer (1.724) to the periphery (1.561). The corresponding values for the crystalline cone are 1.435 and 1.366. The refractive index of the area between cornea and cone is 1.343; that of the clear zone is 1.341, and that of the proximal rhabdom is 1.361. 4. Parallel rays entering a facet converge to a focal region which extends from the proximal part of the cornea to the distal part of the cone. Rays cross the clear zone in a direction which depends on the angle to the axis and position on the facet. Up to an angle of 32° to the axis the rays are mainly bent back into the quadrant of origin so that rays entering many facets converge to a second focal region beyond the clear zone. These findings are consistent with the report of a first inverted although fuzzy image in the cone and a second image (Exner 1891). 5. Ray diagrams were constructed for three different positions of the distal pigment. If the cone tip is completely exposed, the receptor acceptance angle is 46°. With the pigment in the typical dark-adapted position the field of view is 38° wide. A light-adapted ommatidium would have an admission function about 18° wide according to ray tracing, but this could be reduced by the properties of the crystalline tract down which the light must pass. 6. It is concluded from the ray tracing that acuity is poor, but summation across the clear zone could confer a high sensitivity for the dark-adapted eye.


1990 ◽  
Vol 45 (1-2) ◽  
pp. 137-142 ◽  
Author(s):  
Eric J. Warrant ◽  
Robert B. Pinter

Abstract Intracellular recordings of angular sensitivity from the photoreceptors of Aeschnid dragonflies (Hemianax papuensis and Aeschna brevistyla) are used to determine the magnitude and time course of acuity changes following alterations of the state of light or dark adaptation. Acuity is defined on the basis of the acceptance angle, Δρ (the half-width of the angular-sensitivity function). The maximally light-adapted value of Δρ is half the dark-adapted value, indicating greater acuity during light adaptation. Following a change from light to dark adaptation, Δρ increases slowly, requiring at least 3 min to reach its dark-adapted value. In contrast, the reverse change (dark to light) induces a rapid reduction of Δρ , and at maximal adapting luminances, this reduction takes place in less than 10 sec.


1959 ◽  
Vol 14 (4) ◽  
pp. 273-278 ◽  
Author(s):  
Jost Bernhard Walther ◽  
Eberhard Dodt

Behaviour experiments have shown that insects react to ultraviolet light. Almost no data are available within this spectral range, however, on the sensitivity of their light sense organs.In this investigation the relative spectral sensitivity (1/Q) of the compound eye of the fly, Calliphora erythrocephala, and various areas of the compound eye of the cockroach, Periplaneta americana, was measured including the ultraviolet range down to 290 mμ. Equal amplitudes of the electroretinogram indicated equal efficiencies of the stimuli.The sensitivity curve in both species shows, besides the known maximum in the blue green, a second maximum in the ultraviolet. This second maximum was found between 341-369 mμ depending on the species and the particular area of the eye. At still shorter wave lengths sensitivity decreases. In the fly eye and the upper part of the cockroach eye the sensitivity maximum in the ultraviolet is higher than in the bluegreen, whereas in the ventral part of the cockroch eye it is lower. Monochromatic light adaptation selectively influences the relative sensitivity of the upper part of the cockroach eye.The sensitivity curves are discussed with regard to visual pigments and types of receptors. Fluorescence of the eye media is considered to have only negligible if any influence on the high sensitivity for ultraviolet light.


1985 ◽  
Vol 116 (1) ◽  
pp. 385-393 ◽  
Author(s):  
F. E. DOUJAK

1. The absolute-intensity threshold of the optokinetic response in the crab Leptograpsus variegatus was measured using a moving continuous monochromatic point source. The results are compared to the photoreceptor responses of the same animal (Doujak, 1984), to determine the photoreceptor signal for the same behavioural threshold stimuli. 2. Optokinetic eye movements demonstrate that at the peak of spectral sensitivity (499 nm), the minimum intensity of light the animal can detect is 4.0 ± 1.5 × 105 photons cm−2 s−1 (mean ± S.D., N = 18) incident on the eye, which is equivalent to a photon flux of about 6 photons facet−1 s−1. 3. Comparison of behavioural and electrophysiological studies shows that at the above behavioural thresholds, the retinula cells respond with a train of discrete membrane depolarizations (bumps). The mean bump rate recorded in retinula cells at the absolute-intensity threshold of the optokinetic response to a moving point source is 22 ± 5 bumps min−1 (mean ± S.D., N = 6). 4. Optokinetic experiments reveal an absolute sensitivity of the crab's apposition eye to a point source that is only about 900 times less sensitive than the human eye: theoretical estimates based on quantum capture efficiency and lens size predict a much larger difference. The experimental findings provide the first definitive proof that an animal possessing a compound eye can see a star, albeit only stars of 0.5 magnitude and brighter.


Author(s):  
Steven C. Chamberlain

The lateral eye of the horseshoe crab, Limulus polyphemus, is an important model system for studies of visual processes such as phototransduction, lateral inhibition, and light adaptation. It has also been the system of choice for pioneering studies of the role of circadian efferent input from the brain to the eye. For example, light and efferent input interact in controlling the daily shedding of photosensitive membrane and photomechanical movements. Most recently, modeling efforts have begun to relate anatomy, physiology and visually guided behavior using parallel computing. My laboratory has pursued collaborative morphological studies of the compound eye for the past 15 years. Some of this research has been correlated structure/function studies; the rest has been studies of basic morphology and morphological process.


1961 ◽  
Vol 44 (6) ◽  
pp. 1089-1102 ◽  
Author(s):  
Donald Kennedy ◽  
Merle S. Bruno

(1) The spectral sensitivity function for the compound eye of the crayfish has been determined by recording the retinal action potentials elicited by monochromatic stimuli. Its peak lies at approximately 570 mµ. (2) Similar measurements made on lobster eyes yield functions with maxima in the region of 520 to 525 mµ, which agree well with the absorption spectrum of lobster rhodopsin if minor allowances are made for distortion by known screening pigments. (3) The crayfish sensitivity function, since it is unaffected by selective monochromatic light adaptation, must be determined by a single photosensitive pigment. The absorption maximum of this pigment may be inferred with reasonable accuracy from the sensitivity data. (4) The visual pigment of the crayfish thus has its maximum absorption displaced by 50 to 60 mµ towards the red end of the spectrum from that of the lobster and other marine crustacea. This shift parallels that found in both rod and cone pigments between fresh water and marine vertebrates. In the crayfish, however, an altered protein is responsible for the shift and not a new carotenoid chromophore as in the vertebrates. (5) The existence of this situation in a new group of animals (with photoreceptors which have been evolved independently from those of vertebrates) strengthens the view that there may be strong selection for long wavelength visual sensitivity in fresh water.


1972 ◽  
Vol 56 (1) ◽  
pp. 119-128
Author(s):  
U. YINON

The electroretinogram pattern in the compound eye of T. molitor and the appearance of irregular small potentials and spikes superimposed on the ERG are influenced during dark and light adaptation procedures. The amplitude of the principal negative potential reflects bleaching and recovery of the photochemical process. This is not true for the latency values. The delay of the electrical response increases in the dark and decreases in the light adapted eye. These changes were influenced by the intensity of the adapting light. Mutant eyes only lack screening pigment and have normal visual neural pathways. The absence of this pigment lowered the threshold sensitivity of the unscreened eye in dark adaptation. The difference between the adaptation processes in mutants and normal animals has been suggested as a criterion for measuring the net effect of the screening pigment in the compound eye.


1967 ◽  
Vol 50 (9) ◽  
pp. 2267-2287 ◽  
Author(s):  
Robert M. Chapman ◽  
Abner B. Lall

Electrical responses (ERG) to light flashes of various wavelengths and energies were obtained from the dorsal median ocellus and lateral compound eye of Limulus under dark and chromatic light adaptation. Spectral mechanisms were studied by analyzing (a) response waveforms, e.g. response area, rise, and fall times as functions of amplitude, (b) slopes of amplitude-energy functions, and (c) spectral sensitivity functions obtained by the criterion amplitude method. The data for a single spectral mechanism in the lateral eye are (a) response waveforms independent of wavelength, (b) same slope for response-energy functions at all wavelengths, (c) a spectral sensitivity function with a single maximum near 520 mµ, and (d) spectral sensitivity invariance in chromatic adaptation experiments. The data for two spectral mechanisms in the median ocellus are (a) two waveform characteristics depending on wavelength, (b) slopes of response-energy functions steeper for short than for long wavelengths, (c) two spectral sensitivity peaks (360 and 530–535 mµ) when dark-adapted, and (d) selective depression of either spectral sensitivity peak by appropriate chromatic adaptation. The ocellus is 200–320 times more sensitive to UV than to visible light. Both UV and green spectral sensitivity curves agree with Dartnall's nomogram. The hypothesis is favored that the ocellus contains two visual pigments each in a different type of receptor, rather than (a) various absorption bands of a single visual pigment, (b) single visual pigment and a chromatic mask, or (c) fluorescence. With long duration light stimuli a steady-state level followed the transient peak in the ERG from both types of eyes.


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