Fine structure studies of the ocelli of Polyorchis penicillatus (Hydrozoa,Anthomedusae) and their connection with the nerve ring

Summary The fine structure of the small compact ocelli (50–100 m in diameter) of Polyorchis penicillatus is described. The ocellar cup is formed of pigment cells and receptor cells. The pigment cells occur in approximately a 2:1 ratio to the receptor cells. Each pigment cell has a process that may pass through the presumed photosensory region. Pigment cells are connected to adjacent receptor cell processes by septate junctions. The sensory cells are bipolar with the apical part forming the receptor process and the basal part forming an axon 8–15 m long and 1–2 m in diameter. Each receptor cell axon forms a synapse with a single second order neuron but the sensory cells are also connected to the second order neurons postsynaptically. There are also synapses between adjacent second order neurons. The second order neurons lie outside the ocellar cup, next to the tentacular mesogloea. Each second order neuron forms an axon of about 1 m thickness. The axons on each side group together to form an optic nerve having 30–40 axons that travel around the tentacle base on either side and enter the outer nerve ring independently.     

Fine structure of the ocellus of Sarsia tubulosa (Hydrozoa,Anthomedusae)

Summary The fine structure of the ocellus of Sarsia tubulosa is described. The ocellar cup is formed of pigment cells and receptor cells. The receptor cells outnumber the pigment cells in almost a 2:1 ratio. Lateral extensions of neighbouring pigment cells enclose a distal region of 2 to 10 receptor cells. The receptor cell body is 5–7 m in diameter with an apical extension (20–60 m long) that reaches the ocellar cavity. A cilium (9+2 microtubules) arises from the distal part of the receptor cell. The ciliary membrane forms lateral microvilli. The tips of a number of cilia are swollen into large vesicles forming a cornea. The central region of the ocellar cavity contains extracellular electron dense homogeneous material surrounded by swollen ciliary tips and small vesicles. The close apposition between the plasma membrane covering the distal part of adjacent receptor cells as well as the adjacent ciliary shafts suggests the presence of gap junctions. The basal part of each receptor cell forms an axon. The axons of receptor cells form 3 to 4 nerve bundles that join to form the optic nerve. Synapses occur between receptor cell bodies, between axons and receptor cell bodies and among axons.     

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.     

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.     

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     

Fine structure of the segmental ocelli of Armandia brevis (Polychaeta: Opheliidae)

Summary Armandia brevis responds negatively to light during the benthic phase and positively to light during the epitokous phase of its life history. In addition to the prostomial photoreceptors this slender translucent marine worm possesses eleven pairs of ocelli arranged serially from the 7th to the 17th segments.Each ocellus is located at the inner edge of the epidermis slightly in front of the parapodium and contains a single photoreceptor cell which gives off approximately 15 sensory processes. These processes are composed of a central core of neurofibrils surrounded by a mitochondrial layer and a compact array of microvilli. The sensory processes project into and nearly fill the ocellar cavity which is lined by squamous glial cells.The pigment cup enclosing the photoreceptor is composed of about 30 cuboidal cells packed with brown granules. The pigment cells form a mesothelium, being in direct continuity with the coelom. The cup is separated from the glial cells by a basal lamina which separates the epidermal tissues from the mesodermal derivatives of the body wall. Slender muscle fibers traverse the coelom and pass between the cells of the pigment cup.The prostomial photoreceptors were re-examined and found in this material to be composed of microvilli rather than of folds containing labyrinthine tubular infoldings of the cell surface as previously reported.The author thanks Dr. Richard M. Eakin for support and criticism. This investigation was financed by a postdoctoral fellowship, number 1-F2-GM-20, and grant number GM 10292 from the National Institute of General Medical Science.     

Ultrastructural and immunocytochemical observations of the nervous systems of three macrodasyidan gastrotrichs

The nervous systems of three macrodasyidan gastrotrichs, Dactylopodola baltica, Macrodasys caudatus and Dolichodasys elongatus, were investigated using immunocytochemistry and electron microscopy. Labelling of neural structures against serotonin revealed the presence of two pairs of cerebral cells, a dorsal cerebral connective, and paired ventral nerve cords in D. baltica. In M. caudatus and D. elongatus serotonin immunoreactivity was present in a single pair of dorsal cerebral cells and the ventral nerve cords; the dorsal connective of D. elongatus was also immunoreactive to serotonin and acetylated α‐tubulin. The presence of paired, serotonin‐like immunoreactive cells in D. baltica and other species may represent the plesiomorphic condition in Macrodasyida. The fine structure of the photoreceptors in D. baltica was also investigated to explore the potential ground pattern for eyes in the Macrodasyida. The pigmented photoreceptors of D. baltica contain a unicellular pigment cup, sheath cell and sensory receptor. The pigment cup contains numerous osmiophilic granules that presumably function to shield the eyes from downwelling light in the red part of the spectrum. Projecting into the pigment cup and sheath cell are numerous microvilli from a bipolar sensory cell. A single sensory cell may represent the plesiomorphic condition in Macrodasyida, with multiplication of sensory cells representative of more derived taxa.     

The fine structure of the ocelli of Triatoma infestans (Hemiptera: Reduviidae)

The morphology and fine structure of the ocelli of Triatoma infestans have been analyzed by means of light and electron microscopy. The two dorsal ocelli of this species are located behind the compound eyes, looking dorsally and frontally. Externally, the ocelli are marked by the corneal lenses virtually spherical in form and limited internally by a cuticular apodeme. The lens focuses the incoming rays beyond the retina. A single layer of corneagen cells lies below the cuticular lens. The corneagen cells and photoreceptors are arranged in a cup-like fashion beneath the cuticular lens. A distal retinal zone comprises the rhabdoms, which are laterally connected in an hexagonal meshwork. A middle retinal zone comprises the receptor cell segment free of rhabdom, and a proximal zone their axons. In the middle zone, the oviform nuclei and spheroids are located. Screening pigment granules are present within the retinal cell. Spherical mitochondria are homogeneously distributed in the cytoplasm of the cell body. In the axonal zone, mitochondria are found in the peripheral region. Axons from receptor cells extend into the ocellar neuropile at the base of the ocelli, to synapse with second order neurons. The large axons of second order neurons are bundled by glial cells. The ocellar plexus exhibits a high diversity of synaptic unions (i.e. axo-dendritic, axo-axonic, dendro-axonic, and dendro-dendritic).     

Statocyte and Ocellar Pigment Cell in Embryos and Larvae of the Ascidian, Styela plicata (Lesueur)

The processes of formation of two pigmented cells, the statocyte and the ocellar pigment cell, in the cerebral vesicle of larvae of the ascidian Styela plicata were investigated in whole mount specimens and serial paraffin sections by light microscopy. The pigmentations of the two cells became visible simultaneously in embryos at the stage of tail elongation, 5–6 hr after fertilization. The pigmented cells were at first located side by side in the dorsal wall of the neurocoel. Growth of the pigment mass in the ocellus ceased at about 6.5 hr, while that in the statocyte continued through the hatching period (9–10 hr) up to the swimming stage. The pigment mass in the statocyte consisted of two blocks which joined together during their growth. The statocyte migrated from the dorsal to the ventral wall of the cerebral vesicle by the swimming stage. In swimming larvae, the more ventral of the two pigment blocks of the statocyte formed an inverted pigment cup and a cluster of protuberances projected into it from the ventral wall of the cerebral vesicle. Phylogenetically, the sensory organs in the cerebral vesicle of Styela plicata seem intermediate between those in Pyuridae and Botryllinae with respect of their structure and process of differentiation.     

Ultrastructure and development of the rhabdomeric eyes in Lineus viridis (Heteronemertea, Nemertea)

Nemerteans are undoubtedly members of the Spiralia, although their phylogenetic relationships are still a matter of debate. The apparently acoelomate organization suggests a relationship with the platyhelminths, whereas the blood-vascular system has been interpreted as an equivalent to coelomic cavities of annelids, indicating a close relation between annelids and nemerteans. Like other spiralians, most nemertean species are known to have one or several pairs of rhabdomeric and subepidermally situated eyes when adult. The development of these eyes as well as the mode in which the eyes are multiplied is as yet unknown. This is the first attempt to investigate eye formation in a nemertean. In the heteronemertean Lineus viridis (Müller, 1774) the everse rhabdomeric eyes are located deeply underneath the epidermis and consist of a few pigment cells that form a cup-like structure with interdigitating processes that contain numerous pigment granules. In hatchlings, the optical cavity contains processes of 12 sensory cells, each bearing a single cilium and various microvilli. The perikarya of these cells are located distally from the pigment cup. During further development the number of cells increases. Eye development starts with a small anlage situated underneath the epidermis, irrespective of whether this is the first eye or any additional one. The anlage consists of five unpigmented cells and three dendritic processes, each bearing apical microvilli and a single cilium. There is no evidence for an epidermal origin of the eyes. In L. viridis eye formation resembles that described in platyhelminths in which eyes also develop as cerebral derivatives. Although this result has the potential to influence the discussion on the position of Nemertea, the data have to be interpreted with care, since development of L. viridis is derived within the Nemertea.     

Scanning electron microscopy of neurons isolated from the pedal disk and body column of Hydra

Segments of pedal disk and body column were cut from specimens of Hydra littoralis and separated into epidermis and gastrodermis, then macerated to isolate neurons for scanning electron microscopy. Bipolar and multipolar ganglion cells were present in both tissue layers, whereas sensory cells were found only in the gastrodermis. A single cilium projected from the perikaryon of some bipolar and multipolar ganglion cells; the cilium was long in the pedal disk ganglion cells and short in those from the body column. Ganglion cells from the pedal disk had short, thick processes, whereas those from the body column had long, thin neurites. Gastrodermal sensory cells were characterized as unipolar by the presence of an apical cilium near the perikaryon or as asymmetrical bipolar by the presence of a narrow neck region between the perikaryon and cilium. The axon was short in pedal disk sensory cells and long in those from the body column.     

Ultrastructural investigations of presumed photoreceptors as a means of discrimination and identification of closely related species of the genus Microphthalmus (Polychaeta,Hesionidae)

Summary Differences in the ultrastructure of presumed photoreceptors of three morphologically similar Microphthalmus populations on the opposite sides of the Atlantic (German North Sea coast and coasts of North Carolina and Massachusetts) suggest the existence of three different species. Only the European M. listensis possesses three pairs of prostomial eyes, of which one pair has rhabdomeric receptors and pigment cells. The two other pairs are unpigmented and can be found in all three species. The frontal one has ciliary receptors, the posterior one rhabdomeric sensory cells. An additional unpaired potential photoreceptor organ in the segment with the first pair of tentacular cirri is present in all individuals of this species complex. It has a relatively high number of cilia with numerous microvillar projections. — For each type of ocellus there are slight but distinct and constant differences among the species such as relative position of sensory cells, presence of dilations of the ciliary shafts, number of cilia, and shape of the sensory cells. Presence of both ciliary and rhabdomeric light-sensitive cells is discussed with reference to various theories of the evolution of photoreceptors.Abbreviations ax axonema - bb basal body - cc cup cell - ci cilium - cu cuticle - epc epidermal cell - g Golgi apparatus - gp glycogen particles - mi mitochondrion - mv microvilli - mvb multivesicular body - nu nucleus - pc pigment cell - pg pigment granule - rer rough ER - smc submicrovillar cysternae - sr striated rootlet     

First evidence of mitochondrial lensing in two species of the “Typhloplanoida” (Plathelminthes,Rhabdocoela): phylogenetic implications

 Based on electron-microscopical observations the light-sensing organs of Proxenetes deltoides and Ptychopera westbladi, representatives of the ”Typhloplanoida” Trigonostominae, are described. The photoreceptors in both species belong to the type of rhabdomeric pigment cup ocelli. P. deltoides has a single pigment cell and three sensory cells. P. westbladi possesses eyes made up of a single pigmented cup cell and a single sensory cell. The dioptric apparatus in the eyes of P. deltoides is formed by three proliferations of the cup cell containing giant mitochondria. In P. westbladi, the elements focalizing incoming light also consist of modified mitochondria which are arranged in the section of the cup cell covering the eye cavity. With regard to the new findings, mitochondrial lensing is hypothesized as an autapomorphy of a monophylum encompassing distinct taxa or all members of the free-living Rhabdocoela; the Neodermata also belong to this monophylum. Accepted: 21 March 1996     

Sipunculid‐like ocellar tubes in a polychaete,Fauveliopsis cf. adriatica (Annelida,Fauveliopsidae): implications for eye evolution

Abstract. A retractable head region somewhat resembling the introvert of sipunculans is a characteristic feature of members of the annelid taxon Fauveliopsidae. The morphology of fauvelopsids is not well known, and additional data might help to resolve their relationships with other annelids and sipunculans. Ultrastructural investigations of the anterior end of adults of Fauveliopsis cf. adriatica revealed peculiar brain and sensory structures. From the neuropil of the brain, two pairs of lobes mainly composed of neuronal somata extend posteriorly into the peristomium and the following segment. The nuchal organs are embedded in the median pair of lobes, as are additional photoreceptor‐like sensory structures, the ocellar tubes, which are found at the bases of epidermal follicles that extend deeply into the brain. The retractor muscles of the prostomium are attached to the apices of these follicles, which are lined by tendon and supportive cells. The lumen of each follicle is completely filled with cuticular material that forms a rod. Monociliary sensory cells are present all along the length of each follicle; their cilia extend into the cuticle, and are oriented parallel to the longitudinal axis of the tube. Basally, each follicle forms an ovoid extension that is devoid of cuticular material and densely filled with numerous sensory processes—microvilli and cilia—of bipolar sensory cells. The terminal end of the 40‐μm‐deep follicle is formed by two conspicuous cells that contain numerous densely packed vesicles that resemble pigment granules. The ocellar tubes of fauveliopsids are strikingly similar to the ocular tubes of sipunculids. These similarities may reflect common ancestry or may represent convergent evolution; both alternatives are partially supported by previous morphological and molecular studies.     

Evolution of cerebral vesicles and their sensory organs in an ascidian larva

Sorrentino M., Manni L., Lane N. J. and Burighel P. 2000. Evolution of cerebral vesicles and their sensory organs in an ascidian larva. —Acta Zoologica (Stockholm) 81 : 243–258 The ascidian larval nervous system consists of the brain (comprising the visceral ganglion and the sensory vesicle), and, continuous with it, a caudal nerve cord. In most species two organs, a statocyst and an ocellus with ciliary photoreceptors, are contained in the sensory vesicle. A third presumptive sensory organ was sometimes found in an ‘auxiliary’ ganglionic vesicle. The development and morphology of the sensory and auxiliary ganglionic vesicles in Botryllus schlosseri and their associated organs was studied. The sensory vesicle contains a unique organ, the photolith, responding to both gravity and light. It consists of a unicellular statocyst, in the form of an expanded pigment cup receiving six photoreceptor cell extensions. Presumptive mechano‐receptor cells (S1 cells), send ciliary and microvillar protrusions to contact the pigment cup. A second group of distinctive cells (S2), slightly dorsal to the S1 cells, have characteristic microvillar extensions, resembling photoreceptor. We concur with the idea that the photolith is new and derived from a primitive statocyst and the S2 cells are the remnant of a primitive ocellus. In the ganglionic vesicle some cells contain modified cilia and microvillar extensions, which resemble the photoreceptor endings of the photolith. Our results are discussed in the light of two possible scenarios regarding the evolution of the nervous system of protochordates.     

The ultrastructure of the eyes in the veliger-larvae of Aporrhais sp. and Bittium reticulatum (Mollusca,Caenogastropoda)

Summary The cerebrally innervated larval eyes of Aporrhais sp. and Bittium reticulatum are investigated by means of transmission electron microscopy. Each organ consists of a pigmented cup containing an acellular lens. The cornea overlaps the anterior portion of the eye. The retina is composed of sensory cells and supportive cells. The sensory cells of Aporrhais sp. bear one cilium and in Bittium reticulatum two cilia, the ciliary membrane being folded into numerous finger-shaped evaginations. The supportive cells contain the pigment granules and most of them bear one or two cilia, the plasmalemma of which is likewise folded. It is supposed that: (a) these cilia have a transportive function for lens material and (b) that the ciliary photoreceptor of Aporrhais sp. and Bittium reticulatum is a functional adaptation to a relatively long larval period.Abbreviations bb basal body - bp basal plate - c cilium - cc corneal cell - cm ciliary membranes - cw ciliary whorl - gd Golgi dictyosomes - gm granular material - l lens - m mitochondrion - mt microtubules - mv microvilli - mvb multivesicular body - n nucleus - pb pigment border - pg pigment granule - rer rough endoplasmic reticulum - sc sensory cell - sj septate junctions - spc supportive cell     

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.     

Ultrastructural investigations of presumed photoreceptive organs in two Saccocirrus species (polychaeta,saccocirridae)

In addition to the pigmented ocelli, four different types of photoreceptor-like organs without shading pigment have been found in Saccocirrus papillocercus and S. krusadensis. The sensory cells of these presumed ocelli are either ciliary or rhabdomeric with ciliary rudiments. With the exception of the multicellular type-2 ocelli they are bicellular consisting of a sensory cell and a supportive cell. In each ocellus the supportive cell forms a thin cup-shaped envelope around the sensory elements. In the type-2 ocellus, 7 supportive cells form an ovoid cavity leaving openings through which dendritic processes of an equal number of sensory cells enter the cavity. The pigmented ocelli possess an ocellar cavity communicating with the exterior through a pore in the eyecup, ciliary rudiments in both sensory and supportive cell, and additional non-photoreceptive sensory cells in the opening of the eyecup. The sensory organs show characteristic differences between the two species, such as presence or absence of a particular type of ocellus (type 2 is absent in S. krusadensis, type 3 in S. papillocercus), number of cilia in type-4 ocelli, density of microvilli, number of non-photoreceptive sensory cells in the pore of the pigmented ocellus, etc. These differences provide important characters which can be used for discrimination either of species or of subgeneric taxa in Saccocirrus. The phylogenetic significance of the different photoreceptive organs is discussed.     

First evidence of unpigmented rhabdomeric photoreceptors in a Microstomum species (Plathelminthes, Macrostomorpha, Microstomidae) and ultrastructure of photoreceptor-like ciliary aggregations

Microstomum spiculifer possesses a pair of intracerebral photoreceptors each consisting of a single rhabdomeric sensory cell and two cup or mantle cells. The mantle cells are devoid of pigment. In addition, four so-called ciliary aggregations, presumed to have a light-sensing function, are present. Each ciliary aggregation represents a specialized cell with an internal cavity filled with axonemes of modified cilia. Rhabdomeric photoreceptors consisting of one to three sensory cells and a single pigmented or unpigmented mantle cell are widespread within taxa of the Plathelminthes Rhabditophora. On the contrary, the existence of two mantle cells forming the eye cup is only known for M. spiculifer and a few other species of the Macrostomida. Therefore, at least two hypotheses are possible: (1) two cup cells are a basic characteristic of the Rhabditophora and a reduction from two to one cup cell has occurred secondarily or (2) the stem species of the Rhabditophora possessed rhabdomeric eyes with one cup cell, and two mantle cells have evolved within the Macrostomorpha. The existence of ciliary aggregates has been documented for several taxa of the Plathelminthes Rhabditophora. From their distribution it can not be concluded whether these differentiations are either a basic feature of the Rhabditophora or have evolved several times convergently. Accepted: 26 September 1999     

Electrophysiology of the Insect Dorsal Ocellus : III. Responses to flickering light of the dragonfly ocellus

The ERG of the dragonfly ocellus has been analyzed into four components, two of which originate in the photoreceptor cells, two in the ocellar nerve fibers (Ruck, 1961 a). Component 1 is a sensory generator potential, component 2 a response of the receptor axons. Component 3 is an inhibitory postsynaptic potential, component 4, a discharge of afferent nerve impulses in ocellar nerve fibers. Responses to flickering light are examined in terms of this analytic scheme. It has been found that the generator potential can respond to higher rates of flicker—up to 220/sec.—than can the receptor axon responses, the postsynaptic potential, or the ocellar nerve impulses. The maximum flicker fusion frequency as measured by fusion of the ERG is that of the sensory generator potential itself.