Ultrastructure of the photoreceptors of Allostoma sp. (Plathelminthes,Prolecithophora)

The four eyes of the prolecithophoran Allostoma sp. are disposed in two pairs in a dorsolateral position at the periphery of the brain and beneath its capsule. They are rhabdomeric pigment-cup ocelli. Each eye in the anterior pair consists of one pigment cell and one receptor cell; each in the posterior pair is made up of a larger, single pigment cell and two photoreceptor cells. A lens in front of the pigment cell's aperture is formed by electron-dense, refractive, finger-like protrusions which arise from unpigmented cytoplasmic extensions of the pigment-cup margin. Degenerative signs are sometimes visible in the lens.     

The fine structure of the retina of the honey bee drone

Summary The eye of the honey bee drone is composed of approximately 8,000 photoreceptive units or ommatidia, each topped by a crystalline cone and a corneal facet. An ommatidium contains 9 visual or retinula cells whose processes or axons pierce a basement membrane and enter the optic lobe underlying the sensory retina. The visual cells of the ommatidium are of unequal size: six are large and three, small. In the center of the ommatidium, the visual cells bear a brush of microvilli called rhabdomere. The rhabdome is a closed-type one and formed mainly by the rhabdomeres of the six large retinula cells. The rhabdomeric microvilli probably contain the photopigment (rhodopsin), whose modification by light lead to the receptor potential in the retinula cells. The cytoplasm of the retinula cells contains various organelles including pigment granules (ommochromes), and peculiar structures called the subrhabdomeric cisternae. The cisternae, probably composed of agranular endoplasmic reticulum undergo swelling during dark adaptation and appear in frequent connection with Golgi cisternae. Three types of pigment cells are associated with each ommatidium. The crystalline cone is entirely surrounded by two corneal pigment cells. The ommatidium, including its dioptric apparatus and corneal pigment cells, is surrounded by a sleeve of about 30 elongated cells called the outer pigment cells. These extend from the base of the corneal facet to the basement membrane. Near the basement membrane the center of the ommatidium is occupied by a basal pigment cell. Open extracellular channels are present between pigment cells as well as between retinula cells. Tight junctions within the ommatidium are restricted to the contact points between the rhabdomeric microvilli. These results are discussed in view of their functional implications in the drone vision, as well as in view of the data of comparative morphology.This work was supported by a grant from the Fonds National Suisse de la Recherche Scientifique.     

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 fine structure of the lateral ocellus of the dobsonfly larva

The lateral ocelli of the dobsonfly (Protohermes grandis, Neuroptera) larva have been examined with light and electron microscopy. The larva has six ocelli on both sides of the head, each containing a single corneal lens. A conical crystalline body, of some 10–20 cells is situated immediately posterior to the lens. From 100 to 300 elongated retinular cells are arranged perpendicular to the crystalline body except at the innermost surface of the lens, where they are absent. The distal process of each retinular cell is enclosed by a tube-like rhabdom formed by the close association of microvilli from the same and adjacent distal processes. The distal process contains many mitochondria, multivesicular bodies, microtubles and pigment granules. In the dark-adapted ocellus the pigment granules are concentrated near the nucleus which lies under the rhabdomic layer. The granules diffuse toward the rhabdomic microvilli during light adaptation. Each retinular cell has a single axon, which extends from the ocellus as an ocellar nerve fiber into the optic lobe, where it frequently synapses upon second order neurons. In addition to these afferent synapses, there are two other synaptic combinations: (1) a feedback synapse from a second order neuron to a retinular axon, and (2) a synapse between second order neurons. These results suggest that photic signals reach the more proximal part of the brain via second order neurons after some degree of integration in the optic lobe.     

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.     

The distal ommatidium of the compound eye of the housefly (Musca domestica): A scanning electron microscope study

The distal aspect of the housefly ommatidium was surveyed by the scanning electron microscope. Attention was directed to the somal eminence of the superior central cell and the lens to large pigment cell junction. The underside of each lens facet exhibits six hexagonally arranged incisures. Into each of these indentations are fitted several large pigment cells. This hexagonal indentation appears to be a tenacious anchorage. Two corneal pigment cells laterally encircle the pseudocone and at their proximal extension they enclose the Semper cells and neck of the retinula. The somal eminence of the superior central cell is about 10 mum from the base of the corneal pigment cell enclosure. Micrographs were used to construct a diagram of the ommatidium above the basement membrane. Suggestions are made as to the functional correlates of the observed ommatidial structures.     

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.     

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.     

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.     

Ultrastructure of the single-chamber stemmata of Arge pagana (Panzer, 1798) (Hymenoptera: Argidae)

Stemmata are peculiar visual organs of most larvae in holometabolous insects. In Hymenoptera, Symphyta larvae exclusively possess a pair of stemmata, whose cellular organizations have not been thoroughly elucidated to date. In this paper, the morphology and fine structure of stemmata were investigated in the large rose sawfly Arge pagana (Panzer, 1798) using light and electron microscopy. The larvae possess a pair of stemmata, which belong to the “unicorneal composite eye” or single-chamber stemmata. Each stemma is composed of a biconvex cornea lens, a layer of corneagenous cells, numerous pigment cells, and hundreds of retinula cells. According to the number of retinula cells forming a rhabdom, the stemma can be divided into two regions, the larger Region I and the smaller Region II. The former occupies the largest area of the stemma and contains the majority of rhabdoms, each of which is formed by the rhabdomeres of eight retinula cells. The latter occupies a narrow posterior margin, where each rhabdom consists of nine retinula cells. Based on the different cellular organizations of rhabdoms, the stemma of Argidae is likely developed by the fusion of two types of ommatidial units.     

The compound eye of<Emphasis Type="Italic"> Scutigera coleoptrata</Emphasis> (Linnaeus, 1758) (Chilopoda: Notostigmophora): an ultrastructural reinvestigation that adds support to the Mandibulata concept

The lateral compound eye of Scutigera coleoptrata was examined by electron microscopy. Each ommatidium consists of a dioptric apparatus, formed by a cornea and a multipartite eucone crystalline cone, a bilayered retinula and a surrounding sheath of primary pigment and interommatidial pigment cells. With reference to the median eye region, each cone is made up of eight cone segments belonging to four cone cells. The nuclei of the cone cells are located proximally outside the cone near the transition area between distal and proximal retinula cells. The connection between nuclear region and cone segment is via a narrow cytoplasmic strand, which splits into two distal cytoplasmic processes. Additionally, from the nuclear region of each cone cell a single cytoplasmic process runs in a proximal direction to the basement membrane. The bilayered rhabdom is usually made up of the rhabdomeres of 9–12 distal retinula cells and four proximal retinula cell. The pigment shield is composed of primary pigment cells (which most likely secrete the corneal lens) and interommatidial pigment cells. The primary pigment cells underlie the cornea and surround, more or less, the upper third of the crystalline cone. By giving rise to the cornea and by functioning as part of the pigment shield these pigment cells serve a double function. Interommatidial pigment cells extend from the cornea to the basement membrane and stabilise the ommatidium. In particular, the presence of cone cells, primary pigment cells as well as interommatidial pigment cells in the compound eye of S. coleoptrata is seen as an important morphological support for the Mandibulata concept. Furthermore, the phylogenetic significance of these cell types is discussed with respect to the Tetraconata.     

Fine structure of the larval eyes of Mantispa sp. (Neuroptera : Planipennia,Mantispidae)

The stemmata of the first-instar larvae of Mantispa sp. (Neuroptera) were studied by scanning (SEM) and transmission (TEM) electron microscopy. These preparasitic larvae have a pair of anterior eyes and a single posterior eye on each side of the head. Each eye possesses an outer lens; beneath it, there is a well-developed crystalline body and a 3-tiered retina made up of a maximum of 12 sensory cells. The central fused rhabdom appears always to be composed of 4 sensory cells, each filled with pigment granules. The nuclear region shows Golgi bodies and abundant rough endoplasmic reticulum; the rhabdomeric regions contain vesicles, prominent multi-vesicular bodies and lysosomes. The eyes, whether double or single, are surrounded by a perineurium, to which muscle cells are attached.     

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.     

Elimination of egg pigment from developing ocular tissues in the frog Rana pipiens

The method by which egg pigment is eliminated from the developing retina, corneal epithelium and lens in Rana pipiens was studied with light and electron microscopy. The retina expells egg pigment into the space between the retina and pigment epithelium. This pigment is then engulfed by the pigment epithelial cells. The corneal epithelium eliminates egg pigment directly to the outside via the free surface of the epithelial cells. Egg pigment accumulates in a few cells in the lens. These cells probably degenerate and are extruded. These ectodermal derivatives in the eye are free of egg pigment long before ectodermal derivatives in other parts of the embryo lose their pigment. The early elimination of egg pigment from ocular tissues may related to the fact that these tissues must be transparent in order that light may pass freely to the photoreceptors.     

Structure and development of the larval visual system in embryos of Lytta viridana leconte (coleoptera,meloidae)

At hatching (252–264 hr. at 25 ± 0.5°C), the visual system in larvae of Lytta viridana consists of paired stemmata, stemmatal nerves, optic neuropiles, and inner and outer imaginal optic lobe anlagen. It originates between 64 and 72 hr. with invagination of an optic lobe primordium in the side of each protocephalic lobe. These primordia later differentiate into protocerebral ganglion cells and the imaginal optic lobe anlagen. Each stemma arises at 72 hr. from epidermis below and behind the optic lobe invagination and subsequently becomes cupshaped, closes over, and differentiates. At hatching, it consists of a planoconvex corneal lens, a corneagenous layer, and an everse retina of numerous, pigmented retinular cells, each with a terminal rhabdomere. Between 96 and 104 hr, proximal ends of the retinular cells grow posteromedially into a transverse, horizontal fold in the posterior wall of each optic lobe invagination and along its length to the protocerebral neuropile, which they contact by 112 hr. As the brain withdraws posteriorly within the head, these axons elongate correspondingly. Sheath cells of stemmata and stemmatal nerves descend either from protocerebral perineurium or the optic lobe primordia. Structure and development of the larval visual system in L. viridana are compared with those of other insects and its various components are shown to be homologous throughout the Insecta. However, the stemmata of this insect more closely resemble the atypical imaginal eyes of male scale insects than the photoreceptors of other holometabolous larvae–a similarity arising through convergence.     

The role of cell death during morphogenesis of the mammalian eye

Serial sections of embryonic rat eyes were stained with hematoxylin and eosin, quantified (by counting pycnotic and viable nuclei), reproduced by camera lucida on wax plates, and moulded into reconstructions in order to study the normal progression of cellular death during morphogenesis. At least nine distinct necrotic loci (A through I) can be distinguished. Immediately following contact between the retina and surface ectoderm (day 11) degenerating cells were observed in (A) the ventral extent of the optic vesicle, beginning in the mid-retinal primordium and continuing ventrally in the optic stalk, (B) in the rostral optic stalk base, and (C) in the surface ectoderm encircling the early lens placode. No degeneration was observed in the dorsal half of the presumptive retina, in the entire pigment epithelium, or in the lens placode proper. During day 11.5 the lens placode thickens and forms a degenerating locus (D) in its ventral portion opposite the underlying pycnotic zone in the retina (A). During day 12 the ventral pycnotic zone (A) divides into two subunits (A₁ and A₂). Invagination of the lens displaces its marginal and ventral components (C and D) so that they come to occupy the lens pore area and presumptive corneal epithelium. Simultaneous invagination of the retinal rudiment juxtaposes the pigment epithelium which concurrently forms a necrotic area (E) adjacent ventrally to that in the retina (A₁). Degeneration appears in the caudal optic stalk (I). The density of viable cells decreases adjacent to pycnotic areas in the retina and pigment epithelium and increases within these death centers. During day 13 the optic fissure forms within the subunits of the ventral pycnotic zone (A₁ and A₂). Degenerations are seen in the dorsal optic stalk (F) and in the walls of the optic fissure (G and H). Throughout these stages necrosis appears only in those portions of the eye rudiment where invagination is either retarded or completely absent. In part, these observations suggest that cell death serves (1) to retard or inhibit invagination within death centers, (2) to integrate the series of invaginations which mould the dorsal optic cup and optic fissure, (3) to assist formation of the pigment epithelium monolayer, and (4) to orient the lens vesicle within the eye cup. The spatio-temporal relationship between necrotic loci suggests that pycnotic cells in the retina may influence their production in the lens and pigment epithelium. Preliminary observations on the mouse, pig, and human substantiate those on the rat.     

Fine structural description of the lateral ocellus of Craterostigmus tasmanianus Pocock, 1902 (Chilopoda: Craterostigmomorpha) and phylogenetic considerations

The lateral lens eye of adult Craterostigmus tasmanianus Pocock, 1902 (a centipede from Australia and New Zealand) was examined by light and electron microscopy. An elliptical, bipartite eye is located frontolaterally on either side of the head. The nearly circular posterior part of the eye is characterized by a plano-convex cornea, whereas no corneal elevation is visible in the crescentic anterior part. The so-called lateral ocellus appears cup-shaped in longitudinal section and includes a flattened corneal lens comprising a homogeneous and pigmentless epithelium of cornea-secreting cells. The retinula consists of two kinds of photoreceptive cells. The distribution of the distal retinula cells is highly irregular. Variable numbers of cells are grouped together in multilayered, thread-like unions extending from the ventral and dorsal margins into the center of the eye. Around their knob-like or bilobed apices the distal retinula cells give rise to fused polymorphic rhabdomeres. Both everse and inverse cells occur in the distal retinula. Smaller, club-shaped proximal retinula cells are present in the second (limited to the peripheral region) and proximal third of the eye, where they are arranged in dual cell units. In its apical region each unit produces a small, unidirectional rhabdom of interdigitating microvilli. All retinula cells are surrounded by numerous sheath cells. A thin basal lamina covers the whole eye cup, which, together with the distal part of the optic nerve, is wrapped by external pigment cells filled with granules of varying osmiophily. The eye of C. tasmanianus seemingly displays very high complexity compared to many other hitherto studied euarthropod eyes. Besides the complex arrangement of the entire retinula, the presence of a bipartite eye cup, intraocellar exocrine glands, inverse retinula cells, distal retinula cells with bilobed apices, separated pairs of proximal retinula cells, medio-retinal axon bundles, and the formation of a vertically partitioned, antler-like distal rhabdom represent apomorphies of the craterostigmomorph eye. These characters therefore collectively underline the separate position of the Craterostigmomorpha among pleurostigmophoran centipedes. The remaining retinal features of C. tasmanianus agree with those known from other chilopod eyes and, thus, may be considered plesiomorphies. Characters like the unicorneal eye cup, sheath cells, and proximal rhabdomeres with interdigitating microvilli were already present in the ground pattern of the Pleurostigmophora. Other retinal features were developed in the ancestral lineage of the Phylactometria (e.g., large elliptical eyes, external pigment cells, polygonal sculpturations on the corneal surface). The homology of all chilopod eyes (including Notostigmophora) is based principally on the possession of a dual type retinula.     

Fundamental differences in the optical structure of the eyes of nocturnal and diurnal mosquitoes

We have studied the anatomy and optics of the eyes of a range of mosquito species from the wholly dark-active blood-feeding Anopheles gambiae to the diurnal plant-feeder Toxorhynchites brevipalpis. Consistent with studies by Satô in the 1950s, we find that dark-active and crepuscular species have short fused rhabdoms with a conical construction. This maximises the amount of light the rhabdoms receive from the almost hemispherical wide-aperture lenses. Toxorhynchites, on the other hand, has long narrow rhabdomeres that are separated from each other over their entire length, and so resemble the open rhabdoms of advanced flies (Brachycera and Cyclorrhapha). These findings are confirmed by studies of the pseudopupil, whose form indicates the layout of the rhabdomere tips in the focal plane of each ommatidial lens. In anopheline species the pseudopupil is a single undivided ellipse, indicating a fused rhabdom structure, whereas in Toxorhynchites there is a ring of six outer elements surrounding a central one. This means that each rhabdomere views a separate direction in space, and our measurements indicate that, as in higher Diptera, adjacent rhabdomeres share their fields of view with one of the rhabdomeres in the immediately adjacent ommatidia. This in turn means that in the diurnal type of mosquito eye there is a basis for neural superposition, but the fused construction of anopheline rhabdoms precludes this. The Aedes species studied were similar to Anopheles but with lenses of less extreme aperture, and Sabethes cyaneus, a diurnal blood-feeder, was intermediate in structure, with fused conical rhabdoms in the centre of the eye and unfused rhabdomeres around the periphery.     

Fine structure of the anterior median eyes of the funnel-web spider Agelena labyrinthica (Araneae: Agelenidae)

Only few electron microscopic studies exist on the structure of the main eyes (anterior median eyes, AME) of web spiders. The present paper provides details on the anatomy of the AME in the funnel-web spider Agelena labyrinthica. The retina consists of two separate regions with differently arranged photoreceptor cells. Its central part has sensory cells with rhabdomeres on 2, 3, or 4 sides, whereas those of the ventral retina have only two rhabdomeres on opposite sides. In addition, the rhabdomeres of the ventral retina are arranged in a specific way: Whereas in the most ventral part they form long tangential rows, those towards the center are detached and are arranged radially. All sensory cells are wrapped by unpigmented pigment cell processes. In agelenid spiders the axons of the sensory cells exit from the middle of the cell body; their fine structure and course through the eye cup is described in detail. In the central part of the retina efferent nerve fibres were found forming synapses along the distal region of the receptor cells. A muscle is attached laterally to each eye cup that allows mainly rotational movements of the eyes. The optical performance (image resolution) of these main eyes with relatively few visual cells is discussed.