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.     

Did neural pooling for night vision lead to the evolution of neural superposition eyes?

Observations of the infrared deep pseudopupil, optical determinations of the corneal nodal point, and histological methods were used to relate the visual fields of individual rhabdomeres to the array of ommatidial optical axes in four insects with open rhabdoms: the tenebrionid beetle Zophobas morio, the earwig Forficula auricularia, the crane fly Tipula pruinosa, and the backswimmer Notonecta glauca.The open rhabdoms of all four species have a central pair of rhabdomeres surrounded by six peripheral rhabdomeres. At night, a distal pigment aperture is fully open and the rhabdom receives light over an angle approximately six times the interommatidial angle. Different rhabdomeres within the same ommatidium do not share the same visual axis, and the visual fields of the peripheral rhabdomeres overlap the optical axes of several near-by ommatidia. During the day, the pigment aperture is considerably smaller, and all rhabdomeres share the same visual field of about two interommatidial angles, or less, depending on the degree of light adaptation. The pigment aperture serves two functions: (1) it allows the circadian rhythm to switch between the night and day sampling patterns, and (2) it works as a light driven pupil during the day.Theoretical considerations suggest that, in the night eye, the peripheral retinula cells are involved in neural pooling in the lamina, with asymmetric pooling fields matching the visual fields of the rhabdomeres. Such a system provides high sensitivity for nocturnal vision, and the open rhabdom has the potential of feeding information into parallel spatial channels with different tradeoffs between resolution and sensitivity. Modification of this operational principle to suit a strictly diurnal life, makes the contractile pigment aperture superfluous, and decreasing angular sensitivities together with decreasing pooling fields lead to a neural superposition eye.Abbreviations DPP deep pseudopupil - LMC large monopolar cell     

THE MICROSTRUCTURE OF THE COMPOUND EYES OF INSECTS

The apposition eyes of two diurnal insects, Sarcophaga bullata (Diptera) and Anax junius (Odonata), have been examined with the electron microscope. In the latter case only the rhabdom is described. The rhabdom of the fly consists of a central matrix and seven rhabdomeres, one for each retinula cell. The rhabdomeres show an ordered internal structure built up of transverse tubes, hexagonal in cross-section. These slender compartments running the width of the rhabdomere are 370 A in diameter. After fixation with osmium tetroxide the walls of the compartments are more electron dense than the interiors. The retinula cells contain mitochondria, and pigment granules smaller than those found in the pigment cells. These granules tend to cluster close behind the membranes which separate the retinula cells from their rhabdomeres. The rhabdom of the dragonfly is a single structure which appears to be composed of three fused "rhabdomeres," each similar to a rhabdomere of Sarcophaga. Reasons are given for believing that the rhabdom may be the site of photoreception, as well as the organ for analyzing plane-polarized light, as suggested by other workers.     

Different retina-lamina projections in mosquitoes with fused and open rhabdoms

Anopheles gambiae and Toxorhynchites brevipalpis represent the nocturnal and diurnal extremes of the mosquito light intensity range, and their eyes are structurally very different. A. gambiae has fused rhabdoms with huge acceptance angles, whereas T. brevipalpis has open rhabdoms with rhabdomere acceptance angles comparable with those of advanced (brachyceran) flies. Here, we show that the retina-lamina projections are consistent with these differences. The short receptor axons from each ommatidium in A. gambiae insert as a group between four lamina monopolar cell clusters. In T. brevipalpis axon bundles from each ommatidium undergo a twist in their passage through the nuclear layer of the lamina, and then fan out into a space the diameter of which is about twice the separation of the monopolar cell clusters. This arrangement is consistent with a neural superposition mechanism closely similar to that found in higher Diptera, but which must have evolved independently.     

Ultrastructural study of the naupliar eye of the ostracodeVargula graminicola (Crustacea,Ostracoda)

Summary Ostracodes, like other crustaceans, have a simple naupliar eye that is built upon a theme of three eye cups surrounded by a layer of screening pigments. The single naupliar eye of the ostracodeVargula graminicola is situated medially on the dorsal-anterior side of the body and has three fused eye cups, two dorso-lateral and one ventral. Each eye cup has the following components: (1) pigment cells between the eye cups, (2) tapetal cells, (3) retinular cells with (4) microvillar rhabdomeres, and (5) axons extending into the protocerebrum. Typically two retinular cells contribute lateral microvilli to each rhabdom. The two dorso-lateral eye cups have about 40 retinular cells (20 rhabdoms) and the ventral eye cup has about 30 retinular cells (15 rhabdoms). Typical of myodocopid naupliar eyes (as reported from light microscopic studies), no lens cells or cuticular lenses were observed. The presence of tapetal cells identifies theVargula eye as a maxillopod-ostracode type crustacean naupliar eye. It is unlikely that the naupliar eye ofV. graminicola functions in image formation, rather it probably functions in the mediation of simple taxis towards and away from light.     

PHOTORECEPTOR STRUCTURES : III. DROSOPHILA MELANOGASTER

The eyes of three eye mutants of Drosophila melanogaster were fixed and thin sections studied for its structural detail in the electron microscope. Each ommatidium was found to have seven retinula cells with an equal number of rhabdomeres (visual units). The rhabdomeres average 1.2 µ in diameter and 60 µ in length. Each rhabdomere consists of osmium-fixed dense bands averaging 120 A in thickness, and with less dense interspaces 200 to 400 A. There is an average of 23 dense bands or 46 interfaces per micron within the rhabdomere. The rhabdomere as we have presented it is a single structure of packed rods or tubes. The "fine structure" within the rhabdomere is similar to that observed by electron microscopy for the retinula of the house fly, and to the retinal rods of the vertebrate eye, and to the chloroplasts of plant cells in a variety of animal and plant photoreceptor structures. In addition, the radial arrangements within the ommatidium of radially unsymmetrical units, the rhabdomeres, is probably related to the analysis of polarized light in the insect eye.     

Visual behaviour and the structure of dark and light-adapted larval and adult eyes of the New Zealand glowworm Arachnocampa luminosa (Mycetophilidae: Diptera)

Both larval and adult New Zealand cave glowworms exhibit reactions to light; their photoreceptors must, therefore, be regarded as functional. The two principal stemmata of the larva possess large biconvex lenses and voluminous rhabdoms. Approximately 12 retinula cells are present. In light-adapted larvae the diameter of the rhabdom is 8 μm and that of an individual microvillus is 49.5 nm. Dark-adapted eyes have rhabdoms that measure 14 μm in cross section and microvilli with an average diameter of 54 nm. The compound eye of the adult comprises approximately 750 ommatidia, each with a facet diameter of 27–28 μm. A facet is surrounded by 1–6 interommatidial hairs which are up to 30 μm long. The interommatidial angle is 5.5°. Cones, consisting of 4 crystalline cone cells, are of the ‘acone’ type. Pigment granules in the primary pigment cells are twice as large as those of the retinula cells which measure 0.6–0.75 μm in diameter. The rhabdom is basically of the dipteran type, i.e. six open peripheral rhabdomeres surround 2 central rhabdomers arranged in a tandem position. The microvilli of cells 1–6 and cell 8 have diameters ranging from 68 to 73 nm, but those of the distally-located central rhabdomere 7 are 20% larger. This is irrespective of whether the eye is dark or light-adapted. In the latter the cones are long and narrow, the screening pigment granules closely surround the rhabdomeres, and the rhabdom is less voluminous than that of the dark-adapted eye.     

Angular and spectral sensitivity of fly photoreceptors. II. Dependence on facet lens F-number and rhabdomere type in<Emphasis Type="Italic"> Drosophila</Emphasis>

A wave-optical model for the integrated facet lens-rhabdomere system of fly eyes is used to calculate the effective light power in the rhabdomeres when the eye is illuminated with a point light source or with an extended source. Two rhabdomere types are considered: the slender rhabdomeres of R7,8 photoreceptors and the wider, but tapering R1-6 rhabdomeres. The angular sensitivities of the two rhabdomere types have been calculated as a function of F-number and wavelength by fitting Gaussian functions to the effective light power. For a given F-number, the angular sensitivity broadens with wavelength for the slender rhabdomeres, but it stays approximately constant for the wider rhabdomeres. The integrated effective light power increases with the rhabdomere diameter, but it is for both rhabdomere types nearly independent of the light wavelength and F-number. The results are used to interpret the small F-number of Drosophila facet lenses. Presumably the small head puts a limit to the size of the facet lens and favors a short focal length.     

Retinal ultrastructure of the dorsal eye region of Pararge aegeria (Linné) (Lepidoptera: Satyridae)

We examined the fine structure of dorsal rim ommatidia of the compound eye of Pararge aegeria (Lepidoptera: Satyridae) and compared them with ommatidia of the large dorsal region described by Riesenberg (1983 Diploma, University of Munich). 1. The ommatidia of the dorsal rim show morphological specializations known to be typical of the perception of polarized light: (a) the dumb-bell-shaped rhabdoms contain linearly aligned rhabdomeres with only 2 orthogonally arranged microvilli orientations. The rhabdoms are composed of the rhabdomeres of 9 receptor cells, 8 of which are radially arranged. The rhabdomeres of receptor cells VI and V5, as well as D2, D4, D6 and D8 are dorsoventrally aligned, whereas the rhabdomeres of the cells H3 and H7 are perpendicular to them. The rhabdomere of the bilobed 9th retinula cell lies basally and is dorsoventrally aligned, where retinula cell VI and V5 are already axonal. (b) There is no rhabdomeric twist, and (c) the rhabdoms are rather short. 2. However, in the ommatidia of the large dorsal region, only 2 retinula cells (H3 and H7) are suitable for perception of polarized light. 3. Lucifer yellow and horse radish peroxidase were used as tracers to visualize the projections of retinula cell axons of the dorsal rim area and the large dorsal region into the optic neuropils (lamina and medulla). Two receptors (VI and V5) from both the dorsal rim area and the large dorsal region, have long visual fibres projecting into the medulla. The 7 remaining retinula cells of both eye regions, including those that meet the structural requirements for detection of polarized light in the large dorsal region, terminate in the lamina (short visual fibres). These results provide a starting point for further studies to reveal the possible neuronal pathways by which polarized light may be processed.     

Morphological differentiation of the central visual cells R7/8 in various regions of the blowfly eye

The central rhabdomeres in the retina of the blowfly Calliphora erythrocephala and the house fly Musca domestica are not structurally uniform. In Calliphora, four classes of central rhabdomeres were found; they are formed by a total of seven types of central visual cells, clearly distinguished by the following structural features: length of the rhabdomeres R7 or R8, position of the nucleus, rhabdomere twist, fine structure in the R7/R8 transition region, and cross-sectional area of the rhabdomeres. In the lateral part of the eye only the most common central-rhabdomere class, ‘sl.’ is present, whereas in the frontal and dorsal parts classes ‘sl’ and ‘ls’ are found in a particular numerical ratio. Near the frontal eye margin the rare class ‘per’ also appears, with two separate rhabdomeres, R7per and R8s; the morphological properties of R7per are midway between those of peripheral and central visual cells. The special ommatidia at the dorsal margin of the eye are characterized by the central rhabdomeres ‘marg’. The known functional properties of the visual cells in the fly eye can be readily assigned to these classes (Table 1, Fig. 12). The non-uniform distribution of the various kinds of central rhabdomeres suggests functional differentiation of the eye region.     

Specialized ommatidia of the polarization-sensitive dorsal rim area in the eye of monarch butterflies have non-functional reflecting tapeta

Many insects exploit sky light polarization for navigation or cruising-course control. The detection of polarized sky light is mediated by the ommatidia of a small specialized part of the compound eye: the dorsal rim area (DRA). We describe the morphology and fine structure of the DRA in monarch butterflies (Danaus plexippus). The DRA consists of approximately 100 ommatidia forming a narrow ribbon along the dorsal eye margin. Each ommatidium contains two types of photoreceptor with mutually orthogonal microvilli orientations occurring in a 2:6 ratio. Within each rhabdomere, the microvilli are well aligned. Rhabdom structure and orientation remain constant at all retinal levels, but the rhabdom profiles, as seen in tangential sections through the DRA, change their orientations in a fan-like fashion from the frontal to the caudal end of the DRA. Whereas these properties (two microvillar orientations per rhabdom, microvillar alignment along rhabdomeres, ommatidial fan array) are typical for insect DRAs in general, we also report and discuss here a novel feature. The ommatidia of monarch butterflies are equipped with reflecting tapeta, which are directly connected to the proximal ends of the rhabdoms. Although tapeta are also present in the DRA, they are separated from the rhabdoms by a space of approximately 55 μm effectively inactivating them. This reduces self-screening effects, keeping polarization sensitivity of all photoreceptors of the DRA ommatidia both high and approximately equal.     

Zonation of the optical environment and zonation in the rhabdom structure within the eye of the backswimmer,Notonecta glauca

In the compound eye of Notonecta glauca, the backswimmer, there is a small ventral region in which the rhabdoms differ in structure from those in the other parts of the eye. Here, among other unusual features, there is a special orientation of the microvilli of the central rhabdomeres, i.e., in most of the median eye region that has been examined, the microvilli of the two central rhabdomeres are aligned with one another, at an acute angle to the transverse axis of the body. In the small ventral region, the microvilli of these rhabdomeres are perpendicular to one another, those of one rhabdomere being almost exactly in parallel with the median plane of the animal, and those of the other, almost exactly at right angles to the median plane. When Notonecta is hanging under the water surface, the field of vision of the ventral part of the eye coincides with the transparent part of the water surface. Within the ventral eye region there is a bandlike zone only four ommatidia wide; the ommatidia here differ from the others in the ventral eye region by the unique orientation of their central rhabdomeres. With this zone the animal views the area ahead of it just above the water surface. When the backswimmer is flying, the ventral part of the eye views a region that begins under the animal and extends forward from the vertical over ca. 35 degrees. Possible relationships between the special orientation of the microvilli in the ventral eye region and the polarization of the light by the water surface are discussed.     

Cofilin/ADF is required for retinal elongation and morphogenesis of the Drosophila rhabdomere

Drosophila photoreceptors undergo marked changes in their morphology during pupal development. These changes include a five-fold elongation of the retinal cell body and the morphogenesis of the rhabdomere, the light sensing structure of the cell. Here we show that twinstar (tsr), which encodes Drosophila cofilin/ADF (actin-depolymerizing factor), is required for both of these processes. In tsr mutants, the retina is shorter than normal, the result of a lack of retinal elongation. In addition, in a strong tsr mutant, the rhabdomere structure is disorganized and the microvilli are short and occasionally unraveled. In an intermediate tsr mutant, the rhabdomeres are not disorganized but have a wider than normal structure. The adherens junctions connecting photoreceptor cells to each other are also found to be wider than normal. We propose, and provide data supporting, that these wide rhabdomeres and adherens junctions are secondary events caused by the inhibition of retinal elongation. These results provide insight into the functions of the actin cytoskeleton during morphogenesis of the Drosophila eye.     

Structural reactions to polarized light of microvilli in photoreceptor cells of the moth Spodoptera

Summary Intact armyworm moths (Spodoptera exempta, Farn. Noctuidae) were illuminated by polarized monochromatic light to induce structural changes in the rhabdomeres of the compound eyes. The degree of distortion of their microvilli depends on the light energy absorbed per time unit. Under polarized light, the number of quanta absorbed varies with the position of the plane of polarization relative to the axis of the microvilli (intrinsic dichroism). Therefore, in Spodoptera, different degrees of deformations could be demonstrated in differently oriented rhabdomeres of both types of ommatidia. Moreover, in rhabdoms of the lobed type with fan-like arranged microvilli, different reactions were regularly seen in differently oriented microvilli of one rhabdomere. This indicates that microvilli may react to light individually.Supported by Deutsche Forschungsgemeinschaft, Sonderforschungsbereich 114 (Bionach)     

The fine structure of some carabid beetle eyes,with particular reference to ciliary structures in the retinula cells

Paired centrioles and associated ciliary root material occur in all eight retinula cells in the nine species investigated. In the diurnal Notiophilus, Elaphrus and Bembidion where the distal rhabdomere of cell 7 is fused with the proximal rhabdom formed by cells 1 to 6, the roots in cells 1 to 6 extend for the entire length of the retinula. In Notiophilus their arrangement around the rhabdom suggests a complementary mechanical relationship between the six large roots and the four Semper cell processes. In five relatively nocturnal species a retinula cell column separates the distal rhabdomere from the proximal rhabdom. In cells 1 to 6 root material is associated with the distally located centrioles as follows. In Leistus roots extend into the proximal rhabdom layer. In Loricera and Agonum roots at the level of the proximal rhabdom are not continuous with the rootlets or short roots associated with the centrioles. In Pseudophonus and Feronia, and in the diurnal Cicindela, short rootlets link the centrioles. Cell movements on dark-adaptation of Notiophilus and Cicindela include shortening of the crystalline tract. In Notiophilus the entire rhabdom is apparently displaced, whereas in Cicindela the narrow distal rhabdomere becomes dissociated from the proximal rhabdom.     

Direction ofE for maximum response of a retinula cell

Summary Except for very special fused rhabdoms, e. g. those with orthogonal microvilli like the worker bee, the direction of the electric vector E of linear polarized light necessary for a maximum response from a retinula cell is not parallel (or perpendicular) to the microvilli of the recorded cell. This is because the rhabdomeres of a fused rhabdom are optically coupled, i. e. the properties of each rhabdomere influence the manner in which light is transmitted down the composite rhabdom structure. A rhabdom is analogous to a non-uniform absorbing optical crystal. Such a crystal has two coordinate (optical) axes along which E remains linear polarized as it propagates. Only when the microvilli of the recorded cell are parallel to one of these axes will the direction ofE for maximum retinula cell response be parallel to the microvilli. The locust-type of rhabdom is used as an example.     

Angular and spectral sensitivity of fly photoreceptors. I. Integrated facet lens and rhabdomere optics

Three optical components of a fly's eye determine the angular sensitivity of the photoreceptors: the light diffracting facet lens, the wave-guiding rhabdomere and the light-absorbing visual pigment in the rhabdomere. How the integrated optical system of the fly eye shapes the angular sensitivity curves is quantitatively analyzed in five steps: (1) scalar diffraction theory for low Fresnel-number lenses is applied to four different facet lenses, with diameter 10, 20, 40, and 80 micro m, respectively, assuming a constant F-number of 2.2; (2) optical waveguide theory is used to calculate waveguide modes propagating in circular cylindrical rhabdomeres with diameter 1.0, 2.0, and 4.0 micro m, respectively; (3) the excitation of waveguide modes is studied with the tip of the waveguide positioned in the focal plane as well as outside this plane; (4) the light absorption from the various propagated modes by the visual pigment in the rhabdomere is calculated as a function of the angle of the incident light wave; and (5) the angular sensitivity of the photoreceptor is obtained by normalizing the total light absorption. Four wavelengths are considered: 300, 400, 500 and 600 nm. The analysis shows that the wavelength dependency of the lens diffraction is strongly compensated by that of the waveguide modes, an effect which is further enhanced by the decrease in light absorption when the mode number increases. The angular sensitivity of fly photoreceptors is robust to defocus and largely wavelength independent for all except very slender rhabdomeres.     

The eye of Dytiscus (Coleoptera)

The eye of Dytiscus (Coleoptera) has rhabdomeres at three different levels. The crystalline threads stretch from the ends of the crystalline cones only as far as the distal layer of rhabdomeres. There is one distal rhabdo-mere per ommatidium, and in this system the ommatidia are anatomically separate. Between the distal rhabdomere and the rhabdomeres of the next six retinula cells is a wide clear zone in which light entering by one facet could possibly reach deep rhabdomeres of a different ommatidium. Of the six proximal rhabdomeres, four have rhabdomere tubules which lie horizontal with reference to the normal posture, the other two having vertically oriented tubules. The eighth cell, with nucleus near the basement membrane, has a small rhabdomere. All eight retinula cells have axons and there is no other class of axons in the eye.     

Ein Mechanismus zur Steuerung des Lichtflusses in den Rhabdomeren des Komplexauges von Musca

Summary In the ommatidia of Musca, the light flux transmitted by each one of the rhabdomeres of sense cells no. 1 to 6 decreases as a function of time if light falls onto these rhabdomeres. With a similar time course the light flux reflected from these rhabdomeres increases. These changes take place within a few seconds following illumination. The results have been established in the intact animal using changes in the appearance of the pseudopupil as indicator and also in surviving preparations of the eye with direct inspection of the rhabdomeres.The changes are interpreted as a consequence of interactions between pigment granules in the sense cells and electromagnetic fields induced outside the rhabdomeres by light travelling on the inside: In the dark adapted situation the granules are quite distant from the rhabdomeres, the interaction is negligible. During light adaptation the granules move close to the rhabdomeres, and as a consequence, total reflection of the light in the rhabdomere is frustrated. The relatively rapid changes in the optical characteristics of the rhabdomeres are explained by the fact that the distance, the granules have to move in order to switch from one condition to the other is in principle on the order of the wavelength of light.The results indicate, that the changes in the position of the granules are induced by the excitation of the respective sense cells themselves, for instance by the degree of their depolarisation. No interaction between the sense cells of one ommatidium nor between those of different ommatidia could be found.The function of the movement of the pigment granules is interpreted as a means to protect the sense cells no. 1 to 6 against strong illumination. — Movement of pigment granules is not induced in sense cells no. 7 and 8 with light intensities which give maximal response in sense cells no. 1 to 6.

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Wertvolle Diskussionen verdanken wir Herrn Dr. K. G. Götz sowie Herrn Prof. W. Reichardt. Wir danken Fräulein T. Wiegand für Mithilfe bei den Experimenten sowie Herrn E. Freiberg für das Fertigstellen der Abbildungen.     

Retina and dioptric apparatus of the dung beetle Euoniticellus africanus

Retinal fine structure and optics of the eye of the dung beetle Euoniticellus africanus have been studied and compared with those of three other scarabaeid beetles: Repsimus manicatus, Anoplognathus pallidicollis and Sericesthis geminata. The eye of Euoniticellus, in common with that of the other three species, possesses a dioptric system in which light first passes through a thick optically homogeneous cornea, and then enters a non-homogeneous crystalline cone. The lens cylinder properties of the latter cause the light rays to become partially focused across the clear-zone upon the rhabdom layer. Rays traced through a large scale drawing of the eye, with refractive indices measured for each component, predict an acceptance angle of approximately 26°. Since no significant aperture changes, lengthening of crystalline thread, cell or pigment migrations appear to be associated with dark/light adaptation, the eye may be assumed to be permanently poorly focused. In optomotor experiments the beetles did not show their characteristic antennal following response to black and white stripes when the latter had repeat periods of <30°. Structurally the eye of Euoniticellus differs markedly from that of other scarabaeids. It is totally divided into dorsal and ventral eye which are of a different size (the dorsal eye is smaller), but whose structural organization is basically the same. Principal pigment cells (they do not fully surround the cone) as well as accessory pigment cells (they accompany the retinula cells in an extraordinarily regular fashion as far as to the basement membrane) exhibit some unusual features. On the proximal side of the clear-zone, at a level where all retinula cell membranes form complex meanders and convolutions, cell 1 is the first to possess a rhabdomere. In it, all microvilli run parallel. This rhabdomere becomes part of the rectangular proximal rhabdom over the upper 20% of its length. Below this level the rhabdom consists of 6 rhabdomeres, but throughout its length microvilli are oriented in 2 orthogonal directions. It is thought that polarization sensitivity in dung beetles generally is related to the rhabdom organization described for Euoniticellus. An eighth (basal) cell is present in each ommatidium, but it lacks a rhabdomere. A tracheal tapetum is not developed. Finally, the point is made not to regard all different eye structures in insects as perfect adaptations to a particular environment or way of living, for specializations of photoreceptors may either follow, parallel or precede any ecological adaptation.