Membrane maintenance and electrical properties of photoreceptors of wild-type andrpa (receptor potential absent) mutant blowflies (Calliphora erythrocephala)

Summary The maintenance of photoreceptor cell membranes in the blowfly was investigated in relation to the diurnal cycle, age, and therpa (receptor potential absent) phototransduction mutation. The effect of disturbed membrane assembly on the electrical membrane properties was examined using single-electrode discontinuous current-clamp techniques. In wild-type flies the cross-sectional dimensions of the rhabdomeres were markedly reduced with age, and the quantity of synthetic organelles decreased concurrently, whereas no correlation was found between the diurnal cycle and membrane turnover. Therpa mutation is thought to block the visual transduction cascade in photoreceptor cells and to lead to degeneration of the photoreceptor cell bodies. The volume of rhabdomeres decreased markedly inrpa mutants and the quantity of synthetic organelles was reduced significantly, indicating an imbalance between photoreceptive membrane renewal and degradation. Also, the plasma membrane underwent degenerative changes. The passive electrical properties of photoreceptor cells — resting membrane voltages and input resistances — were only slightly changed from those of wild-type flies, although the photoreceptive membrane did not depolarize in response to light. This indicates no apparent disturbance in the function of the ionic channels in these membranes. Taken together, these results suggest that the photoreceptor cells need a functional phototransduction cascade with its feedback controls to maintain continuous renewal of rhabdomeres, but that the plasma membrane maintains its normal electrochemical properties despite extreme morphological degeneration of photoreceptor cell.     

Dynactin affects extension and assembly of adherens junctions in<Emphasis Type="Italic">Drosophila</Emphasis> photoreceptor development

Drosophila eye development is a progressive process including cell fate determination, pattern formation, and rhabdomere morphogenesis. During eye development, a dramatic change in cell shape, which involves turning and extension of the photoreceptor apical surface, occurs in the early pupal stages. It is known that assembly and extension of adherens junctions (AJs) play an important role in this process. In the present study, I show that mutation of the largest subunit of dynactin complexes encoded byGlued (GI) affects the extension and assembly of Ajs in developing photoreceptors. InGl ¹/+ mutants and transgenic flies expressing the dominant negative form of Glued, the AJs failed to properly assemble and extend. In addition, the morphogenesis of rhabdomeres was also affected in these flies. Taken together, these results suggest that the extension and assembly of AJs as well as determination of the rhabdomere domain in photoreceptor development areGl dependent.     

The distribution of actin immunoreactivity in rhabdomeres of tipulid flies in relation to extracellular membrane shedding

Summary Rhabdomeres of tipulid flies lose membrane during turnover from a shedding zone composed of microvillar tips. These distal domains lack intramicrovillar cytoskeletons and appear to be empty sacs of membrane. Recent concerns about the role of ninaC mechano-enzymes in the architecture of dipteran rhabodomeral microvilli and the dynamic role that they may play in the creation of shedding zones demand an examination of the distribution of actin in tipulid rhabdomeres. We compared rhabdomeres from tipulid retinae incubated before fixation for immunocytochemistry in a buffer without additives and a stabilising buffer that contained a cocktail of cysteine protease inhibitors; both were challenged by an anti-actin antibody for immunogold labelling after embedding in LR White Resin. Shedding zones thus processed collapse to structureless detritus. Stabilised and unstabilised shedding zones were immunonegative to anti-actin. To ensure that the negative results were not consequent upon conformational changes generated by the processing protocol, we examined microvilli of degenerating rhabdomeres of the Drosophila light-dependent retinal degeneration mutant rdgB ^KS222 (which separate and collapse without creating a shedding zone) and found the detritus they generate to be immunopositive to anti-actin. Stabilised and unstabilised regions of basal regions of tipulid rhabdomeres were equally immunopositive. We infer that (a) actin is absent from shedding zones; (b) actin is not degraded by microvillar cysteine proteases. The implications of these conclusions are discussed in relation to some functional models of arthropod photoreceptor microvilli.     

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)     

Structure of retinular cells in a Drosophila melanogaster visual mutant,rdgA, at early stages of degeneration

Summary A Drosophila visual mutant rdgA has photoreceptive cells which degenerate gradually after eclosion. Fine structure of the retinular cells of rdgA ^KS60 and rdgA ^K014 was studied during early stages of degeneration to determine the initial morphological defects. The retinular cells of these two alleles showed the following structural abnormality within 1 day after eclosion: (1) rhabdomeres were small and irregular in shape; (2) cisternae of the rough endoplasmic reticulum were more numerous than those in normal retinular cells; (3) submicrovillar cisternae were absent; and (4) lysosomes were fewer than normal. Three-dimensional reconstruction of serial sections of the ommatidia showed that the degeneration of mutant rhabdomeres proceeds more rapidly in regions remote from the nuclei. These results suggest that the process of turnover of rhabdomeric microvilli is abnormal in rdgA. We also confirmed an increase of lysosomes and destruction of cellular organelles, as reported by previous investigators at more advanced stages of degeneration.     

Use of gain-of-function study to delineate the roles of<Emphasis Type="Italic">crumbs</Emphasis> in<Emphasis Type="Italic">Drosophila</Emphasis> eye development

The cell polarity gene,crumbs (crb), has been shown to participate in the development and degeneration of theDrosophila retina. Mutations inCRB1, the human homologue ofDrosophila crb, also result in retinitis pigmentosa and Leber congential amaurosis. In this study, we used the gain-of-function approach to delineate the roles ofcrb in developingDrosophila eye. In the third-instar larval stage, eye development is initiated with photoreceptor differentiation and positioning of photoreceptor nuclei in the apical cellular compartment of retinal epithelium. In the pupal stage, differentiated photoreceptors begin to form the photosensitive structures, the rhabdomeres, at their apical surface. UsingGMR-Gal4 to drive overexpression of the Crb protein at the third-instar eye disc, we found that differentiation of photoreceptors was disrupted and the nuclei of differentiated photoreceptors failed to occupy the apical compartment. Usinghs-Gal4 to drive Crb overexpression in pupal eyes resulted in interference with extension of the adherens junctions and construction of the rhabdomeres, and these defects were stage-dependent. This gain-of-function study has enabled us to delineate the roles of Crb at selective stages of eye development inDrosophila.     

The retina of the phalangid,Opilio ravennae,with particular reference to arhabdomeric cells

Summary The retina of the phalangid, Opilio ravennae, consists of retinula cells with distal rhabdomeres, arhabdomeric cells, and sheath cells. The receptive segment of retinula cells shows a clear separation into a Proximal rhabdom, organized into distinct rhabdom units formed by three or four retinula cells, and a Distal rhabdom, consisting of an uniterrupted layer of contiguous rhabdomeres. One of the cells comprising a retinula unit, the so-called distal retinula cell (DRC), has two or three branches that pass laterally alongside the rhabdom, thereby separating the two or three principal retinula cells of a unit. The two morphologically distinct layers of the receptive segment differ with respect to the cellular origin of rhabdomeral microvilli: DRC-branches contribute very few microvilli to the proximal rhabdom and develop extremely large rhabdomeres in the distal rhabdom only, causing the rhabdom units to fuse. Principal retinula cells, on the other hand, comprise the majority of microvilli of the proximal rhabdom, but their rhabdomeres diminish in the distal rhabdom. It is argued that proximal and distal rhabdoms serve different functions in relation to the intensity of incident light.In animals fixed 4 h after sunset, pigment granules retreat from the distal two thirds of the receptive segment. A comparison of retinae of day- and night-adapted animals shows that there is a slight (approximately 15%) increase in the cross-sectional area of rhabdomeral microvilli in dark-adapted animals, which in volume corresponds to the loss of pigment granules from the receptive segment. The length of the receptive segment as well as the pattern and shape of rhabdom units, however, remain unchanged.Each retinula unit is associated with one arhabdomeric cell. Their cell bodies are located close to those of retinula cells, but are much smaller and do not contain pigment granules. The most remarkable feature is a long, slender distal dendrite that extends up to the base of the fused rhabdom where it increases in diameter and develops a number of lateral processes interdigitating with microvilli of the rhabdom. The most distal dendrite portion extends through the center of the fused rhabdom and has again a smooth outline. All dendrites end in the distal third of the proximal rhabdom and are never present in the layer of the contiguous distal rhabdom. Arhabdomeric cells are of essentially the same morphology in day- and night-adapted animals. They are interpreted as photoinsensitive secondary neurons involved in visual information-processing that channel current collected from retinula cells of the proximal rhabdom along the optic nerve. A comparison is made with morphological equivalents of these cells in other chelicerate species.     

Fine structural organization of the lateral ocelli in two species of Scolopendra (Chilopoda: Pleurostigmophora): an evolutionary evaluation

The lateral ocelli of Scolopendra cingulata and Scolopendra oraniensis were examined by electron microscopy. A pigmented ocellar field with four eyes arranged in a rhomboid configuration is present frontolaterally on both sides of the head. Each lateral ocellus is cup-shaped and consists of a deeply set biconvex corneal lens, which is formed by 230–2,240 cornea-secreting epithelial cells. A crystalline cone is not developed. Two kinds of photoreceptive cells are present in the retinula. 561–1,026 cylindrical retinula cells with circumapically developed microvilli form a large distal rhabdom. Arranged in 13–18 horizontal rings, the distal retinula cells display a multilayered appearance. Each cell layer forms an axial ring of maximally 75 rhabdomeres. In addition, 71–127 club-shaped proximal retinula cells make up uni- or bidirectional rhabdomeres, whose microvilli interdigitate. 150–250 sheath cells are located at the periphery of the eye. Radial sheath cell processes encompass the soma of all retinula cells. Outside the eye cup there are several thin layers of external pigment cells, which not only ensheath the ocelli but also underlie the entire ocellar field, causing its darkly pigmented. The cornea-secreting epithelial cells, sheath cells and external pigment cells form a part of the basal matrix extending around the entire eye cup. Scolopendromorph lateral ocelli differ remarkably with respect to the eyes of other chilopods. The dual type retinula in scolopendromorph eyes supports the hypothesis of its homology with scutigeromorph ommatidia. Other features (e.g. cup-shaped profile of the eye, horizontally multilayered distal retinula cells, interdigitating proximal rhabdomeres, lack of a crystalline cone, presence of external pigment and sheath cells enveloping the entire retinula) do not have any equivalents in scutigeromorph ommatidia and would, therefore, not directly support homology. In fact, most of them (except the external pigment cells) might be interpreted as autapomorphies defining the Pleurostigmophora. Certain structures (e.g. sheath cells, interdigitating proximal rhabdomeres, discontinuous layer of cornea-secreting epithelial cells) are similar to those found in some lithobiid ocelli (e.g. Lithobius). The external pigment cells in Scolopendra species, however, must presently be regarded as an autapomorphy of the Scolopendromorpha.     

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)     

The nucleolar fibrillar centres in various cell types in vivo or in vitro

Summary The compound eye of male (haploid) Xyleborus ferrugineus beetles was examined with scanning and transmission electron microscopy. The eye externally consists of ca. 19 to 33 facets. Each ommatidium is composed of a thickly biconvex lenslet with about 50 electron dense and rare layers, but at the junction area between two lenslets there are only about 35 to 37 layers that can be distinguished. A very short (3.4–4.0 m) acone type crystalline cone is located directly beneath the lenslet. Each ommatidium is surrounded by pigment cells, and pigment granules also appear throughout the cytoplasm of the retinular cells. Some pigment granules are even present below the basement membrane. There are 8 retinular cells. The rhabdomeres of 2 centrally situated photoreceptor cells fuse into a rhabdom which is enveloped by the rhabdomeres of 6 peripheral retinular cells. The rhabdomeres of the 6 peripheral retinular cells join laterally to form a rhabdomeric ring around the central rhabdom. No tracheation was observed among the retinular cells. Virus-like particles are evident near the nucleus in each Semper cell of the crystalline cone.This research was supported by the Director of the Research Division, C.A.L.S., University of Wisconsin, Madison; and in part by research grant No. RR-00779 from the Division of Research Resources, National Institutes of Health and by funds from the Schoenleber Foundation, Milwaukee, WI to D.M.N.     

Deoxyglucose mapping of nervous activity induced inDrosophila brain by visual movement

Summary Autoradiographs of the brains of the visual mutantsouter rhabdomeres absent ^JK84 (ora),small optic lobes ^KS58 (KS58) andno object fixation E ^B12 (B12) have been obtained by the deoxyglucose method. The patterns of metabolic activity in the optic lobes of the visually stimulated mutants is compared with that of similarly stimulated wildtype (WT) flies which was described in Part I of this work (Buchner et al. 1984b).In the mutantKS58 the optomotor following response to movement is nearly normal despite a 40–45% reduction of volume in the visual neuropils, medulla and lobula complex. InB12 flies the volume of these neuropils and the optomotor response are reduced. In autoradiographs of both mutants the pattern of neuronal activity induced by stimulation with moving gratings does not differ substantially from that in the WT. It suggests that only neurons irrelevant to movement detection are affected by the mutation. However, in the lobula plate of someKS58 flies and in the second chiasma of allB12 flies, the pattern of metabolic activity differs from that observed in WT flies. Up to now no causal relation has been found between the modifications described in behaviour or anatomy and those observed in the labelling of these mutants.In the ommatidia ofora flies the outer rhabdomeres are lacking while the central photoreceptors appear to be normal. Stimulus-specific labelling is absent in the visual neuropil of these mutants stimulated with movement or flicker. This result underlines the importance of the outer rhabdomeres for visual tasks, especially for movement detection.Abbreviations DG deoxyglucose - KS58 small optic lobes^KS58 - B12 no object fixation E^B12 - JK84 ora outer rhabdomeres absent ^JK84 - WT wildtype     

Ultrastructure of the eye of Urastoma cyprinae (Turbellaria,Alloeocoela)

Summary Urastoma cyprinae (Graff) is a microturbellarian which has been recorded both as a free-living organism by Westblad (1955) and Marcus (1951) and as a commensal in various lamellibranch molluscs (see Burt & Drinnan 1968). The material used in this study came from oysters, Crassostroea virginica, collected off the coast of Prince Edward Island, in which hosts it occurs in large numbers especially during the summer months when the oysters are spawning (Fleming et al. 1981). When U. cyprinae is exposed to light as happens, for example, when an oyster is opened, it shows a marked negative phototactic response.Preliminary work on the fine structure of the photoreceptors in U. cyprinae shows that the two eyes each consists of: (1) a single cup cell full of relatively large, electron-dense pigment granules; (2) a tripartite conical lens system; and (3) what appear to be two photosensitive rhabdomes. The pigment cup cell has a single, well defined nucleus situated basally and close to the membrane of the pigment cell furthest away from the rhabdomeres. The lens system consists of a cone made up of three, separate but equal, parts. Each part has two, flat inner surfaces which join at an angle of 120°, an outer rounded surface, and a rounded upper surface. When these three parts fit together, the cone-shaped lens is formed with the apex of the lens within the cup of the pigment cell and the rounded, convex, broad end of the cone lying more or less at the same level as the top of the pigment cup and below the epidermis layer. The rhabdomeres lie between the electron dense lenses and the inside of the pigment cup. They show connections to the visual cells which are bipolar: one extension joining the rhabdomeres; the other constituting the axon which extends into the centrally situated brain or into the longitudinal, lateral nerves. The axons that enter the brain, form connections with other axons from the other eye. The axons that extend posteriorly in a lateral position, presumably play a role in facilitating the avoidance reaction.The chemical nature of the unusual lens has not yet been determined. This is presently under investigation and will be reported later at which time our work will be discussed in relation to other types of rhabdomeric eyes in the Turbellaria.     

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     

Assembly of rhabdomeric membrane from smooth endoplasmic reticulum can be activated by light in chromophore-deprived photoreceptors of Manduca sexta

Summary Deficiency of the photopigment chromophore, resulting from carotenoid/retinoid (vitamin A) deprivation, that severely impairs the visual function of Manduca sexta also leads to the hypertrophy of smooth endoplasmic reticulum in the photoreceptors. The excess endomembrane accumulates in the stacked cisternae of myeloid bodies. Although 11-cis retinal promotes substantial recovery of function in the retinas of deprived moths maintained in darkness, the myeloid bodies remain. When such recovering photoreceptors were exposed to light of moderate intensities, the amount of endomembrane diminished to normal levels over a period of several hours, while rhabdomeres grew larger. Since there was no endocytolysis, the myeloid bodies must have provided the membrane for rhabdomere enlargement. Bright light similarly mobilized the myeloid bodies in deprived receptors. Thus the persistence of myeloid bodies in moderately illuminated chromophoredeficient receptors is a consequence of their insensitivity. However, the initial hypertrophy of endomembrane does not appear to result from the lack of adequate stimulation: normal, chromophore-replete photoreceptors maintained in darkness from before the period of retinal development had large rhabdomeres and no myeloid bodies. The development of myeloid bodies during the differentiation of vitamin A-deprived photoreceptors appears to entail an influence of the chromophore at another level of receptor cell function.     

Freeze-fracture study of the Drosophila photoreceptor membrane: mutations affecting membrane particle density

The photoreceptor membrane of Drosophila melanogaster (wild type, vitamin A-deprived wild type, and the mutants ninaAP228, ninaBP315, and oraJK84) was studied by freeze-fracture electron microscopy. The three mutations caused a decrease in the number of particles on the protoplasmic face of the rhabdomeric membrane. The ninaAP228 mutation affected only the peripheral photoreceptors (R1-6), while the ninaBP315 mutation affected both the peripheral (R1-6) and the central photoreceptors (R7). The oraJK84 mutation, which essentially eliminates R1-6 rhabdomeres, was found to drastically deplete the membrane particles in the vestigial R1-6 rhabdomeres but not in the normal rhabdomeres of R7 photoreceptors, suggesting that the failure of the oraJK84 mutant to form normal R1-6 rhabdomeres may be due to a defect in a major R1-6 photoreceptor-specific protein in the mutant. In all cases in which both the rhabdomeric particle density and rhodopsin content were studied, the mutations or vitamin A deprivation was found to reduce both these quantities, supporting the idea that at least the majority of the rhabdomeric membrane particles are closely associated with rhodopsin. Vitamin A deprivation and the mutations also reduced the number of particles in the plasma membrane as in the rhabdomeric membrane, suggesting that both classes of membrane contain rhodopsin.     

Twisted rhabdomeres in the dipteran eye

Summary The rhabdomeres of the visual cells in the blowflyCalliphora erythrocephala and the fruit flyDrosophila melanogaster are twisted along their long axes.In rhabdomeres of the visual cells R1–6 it is possible to distinguish 3 regions differing in twist rate. In the proximal and distal regions the twist is slight (e.g., 0.52°/m) or absent, whereas in the middle the twist rate is high (e.g., 2.40°/m). The twisting of the rhabdomeres of R1–3 is congruent and codirectional, and that of R4–6 is its mirror image. The significance of twisting with regard to the dichroic absorption of the microvilli and to the polarization sensitivity and the self-screening of R1–6 is discussed. In particular, it is shown that the dichroic absorption of a single microvillus of R1–6 must be greater than 2; it follows that the absorbing dipoles of the visual pigment molecules must be more or less parallel to the axes of the microvilli. Finally, it can be shown that the twisting of the rhabdomeres R1–6 prevents self-screening — despite high microvillar absorption. Because the microvilli are not uniformly oriented, the twisted rhabdomeres R1–6 are especially effective in absorbing unpolarized light.This work was supported by a grant from the Deutsche Forschungsgemeinschaft (Sm 16/3)     

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.     

Microspectrophotometric characterization and localization of three visual pigments in the compound eye ofNotonecta glauca L. (Heteroptera)

Summary The absorption maxima ( _max) of the visual pigments in the ommatidia ofNotonecta glauca were found by measuring the difference spectra of single rhabdomeres after alternating illumination with two different adaptation wavelengths. All the peripheral rhabdomeres contain a pigment with an extinction maximum at 560 nm. This pigment is sensitive to red light up to wavelengths > 700 nm. In a given ommatidium in the dorsal region of the eye, the two central rhabdomeres both contain one of two pigments, either a pigment with an absorption maximum in the UV, at 345 nm, or — in neighboring rhabdoms — a pigment with an absorption maximum at 445 nm. In the ventral part of the eye only the pigment absorbing maximally in the UV was found in the central rhabdomeres. The spectral absorption properties of various types of screening-pigment granules were measured.     

Fine structure of photoreceptors in larval trematodes

Summary The fine structure of photoreceptors is described in miracidia of Fasciola hepatica, Heronimus chelydrae, Allocreadium lobatum, and Spirorchis sp., and in a spirorchiid cercaria. All have in common eyespots consisting of pigment cells with chambers occupied by rhabdomeres consisting of retinular cell dendrites with numerous microvilli. Photoreceptors of the miracidia show a bilateral asymmetry which is most pronounced in H. chelydrae with a pair of well separated eyespots unequal in size. The smaller right one consists of a pigment cell and two rhabdomeres; the larger left eyespot has an anterior pigment cell with two rhabdomeres and a posterior cell containing one rhabdomere. Photoreceptors in the other species of miracidia also have five rhabdomeres but contain only two pigment cells which are closely apposed. Each contains a pair of lateral rhabdomeres and a fifth one occupies a posteromedian extension of the left pigment cell. In the number of rhabdomeres, their relationship to pigment cells and the resulting asymmetry, photoreceptors are more alike in the distantly related species of miracidia studied than they are in ocellate cercariae or even in the miracidium and cercaria of the same species or two closely related ones. From the asymmetry of photoreceptors in larvae of certain flatworms other than digenetic trematodes, it seems that eyespots of miracidia have retained an ancestral pattern whereas the diversity of photoreceptors in cercariae reflects the varied phototactic behavior of those larvae which complete their life cycles by all the means known for cercariae with a free-swimming period. In both miracidia and cercariae, photoreceptors show an anterior-posterior organization that would seem to be concerned with orientation of the larvae with respect to light.Supported in part by a David Ross Fellowship of the Purdue Research Foundation and in part by U.S.P.H.S. Grants 1T1 GM 1392 01 and 2T1 Al 106 07. We express thanks to Dr. Keith Dixon for aid in obtaining and processing miracidia of Fasciola hepatica; to Prof. Clark P. Read for his valuable comments and suggestions; and to Profs. Charles W. Philpott and Richard H. White for advice concerning electron microscopy.     

The compound eye of Parnara guttata (Insecta,Lepidoptera, Hesperiidae): Fine structure of the ommatidium

Summary The fine structure of an ommatidium of a skipper butterfly, Parnara guttata, has been studied using the electron microscope. Each ommatidium has nine retinula cells, which were classified into three groups: two distal, six medial and one basal retinula cells. The rhabdomeres of the distal retinula cells are localized in the distal part of the rhabdom, while those of the six medial retinula cells appear throughout most of the rhabdom. The rhabdomere of the basal retinula cell occupies only the basal part of the rhabdom. The rhabdomeres of four medial cells are constructed of parallel microvilli, while fan-like microvilli form the rhabdomeres of other two medial retinula cells. The distal and basal retinula cells have rhabdomeres consisting of both parallel and fan-like microvilli. This is the first time the construction of the rhabdomeres of the distal and basal retinula cells has been described in such fine detail for a skipper butterfly. Nine retinula cell axons of each ommatidium extend to the first neuropile of the optic lobe, the lamina ganglionaris. No difference was found in the number of retinula cells of an ommatidium or the shape of the rhabdom between the dorsal and ventral regions of the compound eye.