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.     

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.     

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.     

Ultrastructure of the larval eye of the scorpionfly Panorpa dubia (mecoptera: Panorpidae) with implications for the evolutionary origin of holometabolous larvae

The evolutionary origin of holometabolous larvae is a long‐standing and controversial issue. The Mecoptera are unique in Holometabola for their larvae possessing a pair of compound eyes instead of stemmata. The ultrastructure of the larval eyes of the scorpionfly Panorpa dubia Chou and Wang, 1981 was investigated using transmission electron microscopy. Each ommatidium possesses a cornea, a tetrapartite eucone crystalline cone, eight retinula cells, two primary pigment cells, and an undetermined number of secondary pigment cells. The rhabdomeres of the eight retinula cells form a centrally‐fused, tiered rhabdom of four distal and four proximal retinula cells. The rhabdomeres of the four distal retinula cells extend distally into a funnel shape around the basal surface of the crystalline cone. Based on the similarity of the larval eyes of Panorpidae to the eyes of the hemimetabolous insects and the difference from the stemmata of the holometabolous larvae, the evolutionary origin of the holometabolous larvae is briefly discussed. Morphol., 2012. © 2012 Wiley Periodicals, Inc.     

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.     

Structure of the visual system of the two-spotted spider-mite, Tetranychus urticae

The pair of eyes on each side of Tetranychus urticae consists of fifteen retinular cells, six pigment cells, six corneal cells, and one vitreous cell. Five rhabdomeres lie beneath the anterior lens, twelve beneath the posterior lens. The pigment and corneal cells appear to determine the positions at which rhabdomeres occur. The volumes of all rhabdomeres have been measured. A pair of fibrous masses are associated with tendons at the dorsal apodemes; the nerves of these ‘apodemal organs’ join the optic nerves en route to the brain. There are a pair of optic neuropiles, each surmounted by a large secretory cell.     

Immunohistochemical evidence for multiple photosystems in box jellyfish

Cubomedusae (box jellyfish) possess a remarkable visual system with 24 eyes distributed in four sensory structures termed rhopalia. Each rhopalium is equipped with six eyes: two pairs of pigment cup eyes and two unpaired lens eyes. Each eye type probably captures specific features of the visual environment. To investigate whether multiple types of photoreceptor cells are present in the rhopalium, and whether the different eye types possess different types of photoreceptors, we have used immunohistochemistry with a range of vertebrate opsin antibodies to label the photoreceptors, and electroretinograms (ERG) to determine their spectral sensitivity. All photoreceptor cells of the two lens eyes of the box jellyfish Tripedalia cystophora and Carybdea marsupialis displayed immunoreactivity for an antibody directed against the zebrafish ultraviolet (UV) opsin, but not against any of eight other rhodopsin or cone opsin antibodies tested. In neither of the two species were the pigment cup eyes immunoreactive for any of the opsin antibodies. ERG analysis of the Carybdea lower lens eyes demonstrated a single spectral sensitivity maximum at 485 nm suggesting the presence of a single opsin type. Our data demonstrate that the lens eyes of box jellyfish utilize a single opsin and are thus color-blind, and that there is probably a different photopigment in the pigment cup eyes. The results support our hypothesis that the lens eyes and the pigment cup eyes of box jellyfish are involved in different and specific visual tasks.     

Photoreceptors with mitochondrial lenses inPogaina suecica (Plathelminthes,Rhabdocoela)

Summary The photoreceptors ofPogaina suecica correspond to the type of pigment cup ocelli. Each eye consists of one cup cell and three sensory cells. The most conspicuous differentiations of these eyes are lens elements formed by giant mitochondria densely filled with homogeneous electron-dense material. From electron microscopical findings available to date it is hypothesized that mitochondrial lensing might be an autapomorphy of a taxon comprising the Provorticidae Kirgisellinae, Dalyelliidae and Graffillidae groups which are ascribed to the paraphyletic Dalyellioida.Abbreviations l _1–3 lenses - npc nucleus of the pigment cell - pc pigment cell - pg pigment granule - rh rhabdomeres - sc _1–3 sensory cells     

The larval ocelli of Golfingia misakiana (Sipuncula, Golfingiidae) and of a pelagosphera of another unidentified species

 The inverse cerebral ocelli of the pelagosphera larva of Golfingia misakiana and of another unidentified larva are composed of two or three sensory cells and one supportive pigmented cell. The sensory cells bear an array of microvilli as well as a single cilium with poor undulation of its membrane; the photoreceptive organelles are regarded as the rhabdomeric type. A striking feature of these cells is the cores, which extend within the microvilli from the tip into the midregion of the cell. It is suggested that these structures are identical with the submicrovillar cisternae found in the cerebral inverse eyes of larvae of Polychaeta. The findings allow the conclusion that in the pelagosphera of the Sipuncula, contrary to the teleplanic veliger larvae of Gastropoda, a lengthy pelagic cycle is not correlated with the development of a ciliary photoreceptor. Additionally, it is assumed that the pigment cup ocelli in larvae of Sipuncula are homologous with the cerebral inverted pigment cup ocelli of larvae of Polychaeta. Accepted: 19 March 1997     

The ultrastructure of the nauplius eye ofSapphirina (Crustacea: Copepoda)

Summary The ultrastructure of the specialized nauplius eye of three species of the copepod genusSapphirina was investigated. The gross morphology described earlier (Elofsson, 1966a) was confirmed. The ventral cup is covered by a red pigment and the lateral cups by a red and a black pigment. The ultrastructural configuration of the pigment granules was found to differ in the two kinds of pigment cells. The black pigment cell, moreover, contains a large number of transversely banded fibrils and is able to produce reflecting crystals. The pigment granules of the black pigment cell show a variation in electron density. An intimate connexion exists between the black pigment cell and large retinula cells in the lateral cups, indicating an exchange of material. The tapetal cells present in all three cups form crystal platelets contained in two sets of membranes. It is suggested that the ventral cup and part of the lateral cups function as thePecten-eye (Land, 1965). The rhabdomeres of the retinula cells are composed of microvilli measuring 400 Å. The orientation of these seems to exclude polarotactic behaviour. The ventral cup and the four small cells of the lateral cups contain some retinula cells with microvilli arranged parallel to the incoming light. The retinula cells further develop an intricate system of membrane-invaginations penetrating deep into the cell and associated with numerous mitochondria. Retinula cells of the ventral cup and part of the lateral cups contain clear portions filled with granular material only. Retinula and other cells contain attenuated mitochondria with parallel tubuli. The proximal lens in front of each lateral cup consists of one cell. A development from the conjunctival cells is suggested. The results are evaluated in terms of function and evolution.This work has been supported by a grant from the Swedish Natural Science Research Council (2760-2).     

The fine structure of the pallial eyes of Laternula truncata (Bivalvia: Anomalodesmata: Pandoracea)

The structure of the pallial eyes of Laternula truncata (Lamarck 1818) has been studied using the light and electron microscopes. The eye is complex and can be- considered to be- the most advanced yet described for a bivalve mollusc. The cornea consists of modified flattened epithelial cells with an external border of microvilli. The cornea covers a large, circular, multinucleate lens. The lens comprises (1) centrally located translucent lens cells, (2) laterally located supporting cells from which cell processes interdigitate with processes from the lens cells. The retina is two layered and inverted. The proximal and distal retinae are made up of concentrically arranged laminae derived from the membranes of ciliary basal bodies. The cilia comprise a base and feet, but no root system and have a 9+0 arrangement of filaments.
The pigment cup or tapetum is bounded by a sclerotic coat and is three layered, each layer possessing characteristic pigment granules. From the base of the eye arises a large optic nerve.
The eye possesses an eye appendage, the epithelium of which is invaginated on its internal border to form a groove within which are found some 28 cilia. The cilia, it is thought, make contact with the microvilli of the epithelium when the appendage is touched. Such an action serves to protect the delicate eye from damage.
The structure of the eye is compared with that of other molluscs, particularly members of the Bivalvia.     

Alterations of the eyes of Carinaria lamarcki (Gastropoda, Heteropoda) during the long pelagic cycle

 The eyes of different larval stages of Carinaria lamarcki were examined ultrastructurally. In all larval stages the eyes consist of a cornea, a lens and an everse retina. The photoreceptors in young larvae are exclusively of the ciliary type. In old larvae, however, two types of photoreceptors are present and the retina is composed of two segments: a posterior segment with altered ciliary photoreceptors (=type I sensory cells) and an anterior segment with what are presumably rhabdomeric photoreceptors (=type II sensory cells). The anterior retina is interpreted here as an accelerted character. Furthermore, the arrangement of the pigment granules changes during the long larval development being cup shaped in young larvae versus ribbon shaped in old larvae. The findings allow for the conclusions that: (a) the ciliary photoreceptors are correlated with the long larval period of Heteropoda and that (b) the eyes are altered continuously during the larval cycle. Accepted: 6 July 1998     

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.     

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.     

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.     

Ultrastructure of the eyes of the larva of Neoheterocotyle rhinobatidis (Platyhelminthes, Monopisthocotylea), and phylogenetic implications.

The oncomiracidium of Neoheterocotyle rhinobatidis (Monogenea, Monopisthocotylea, Monocotylidae) has two pairs of eyes, each eye with a lens and pigment cup. The anterior eyes have a single rhadomere; the posterior ones, two rhadomeres. Lenses are part of the pigment cup cells, as indicated by cytoplasmic connections between them and the pigment cups, and they are of mitochondrial origin because mitochondrial cristae are present in the periphery of the lenses. This is the first time that mitochondrial lenses have been shown to exist in a neodermatan. Such lenses may be a synapomorphy of a large taxon comprising the Neodermata and its turbellarian sister groups, or they may have evolved convergently in several not closely related groups as a result of strong selection pressure to find a suitable habitat or host.     

Retinal ultrastructure may mediate polarization sensitivity in larvae of the Sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae)

A number of invertebrates are known to be sensitive to the polarization of light and use this trait in orientation, communication, or prey detection. In these animals polarization sensitivity tends to originate in rhabdomeric photoreceptors that are more or less uniformly straight and parallel. Typically, polarization sensitivity is based on paired sets of photoreceptors with orthogonal orientation of their rhabdomeres. Sunburst diving beetle larvae are active swimmers and highly visual hunters which could potentially profit from polarization sensitivity. These larvae, like those of most Dytiscids, have a cluster of six lens eyes or stemmata (designated E1 through E6) on each side of the head capsule. We examined the ultrastructure of the photoreceptor cells of the principal eyes (E1 and E2) of first instar larvae to determine whether their rhabdomeric organization could support polarization sensitivity. A detailed electron microscopical study shows that the proximal retinas of E1 and E2 are in fact composed of photoreceptors with predominantly parallel microvilli and that neighboring rhabdomeres are oriented approximately perpendicularly to one another. A similar organization is observed in the medial retina of E1, but not in the distal retinas of E1&2. Our findings suggest that T. marmoratus larvae might be able to analyze polarized light. If so, this could be used by freshly hatched larvae to find water or within the water to break the camouflage of common prey items such as mosquito larvae. Physiological and behavioral tests are planned to determine whether larvae of T. marmoratus can actually detect and exploit polarization signals.     

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.     

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.     

Development and differentiation of the eye in Platynereis dumerilii (Annelida,Polychaeta)

The nereid polychaete, Platynereis dumerilii, possess two pairs of post-trochophoral eyes with one vitreous body each. The development of these eyes has first been observed in 2-day-old larvae. Whether the eye anlagen arise from stem cells or from undifferentiated ectodermal tissue was not determined. At first, the anlagen of the anterior and the posterior eyes adjoin each other. They separate in late 3-day-old larvae. The first separated eye complexes consist each of two supporting and two sensory cells. The supporting cells synthesize two different kinds of granules, the pigment granules of the pigment cup and the prospective tubules of the vitreous body. These tubules accumulate in the distal process of the supporting cell. The vitreous body is formed by compartments of the supporting cells filled with the osmiophilic vitreous body tubules. The short, bulbar photosensory processes bear microvilli that emerge into the ocular cavity. At the apex of each sensory cell process, a single cilium (or occasionally two) arises. The sensory cells contain a different kind of pigment granule within their necks at the level of the pigment cup. The rate of eye development and differentiation varies. New supporting cells are added to the rim of the eye cup. They contribute to the periphery of the vitreous body like onion skins, and sensory cells move between supporting cells. The older the individual compartments of the vitreous body are, the more densely packed is their content of vitreous body tubules. Elongation of the sensory and supporting cell processes of the older cells increases the volume of the eye. The eyespots of the trochophore are briefly described as of the two-celled rhabdomeric type with a single basal body with ciliary rootlet.