Comparative morphology of statocysts in the Plathelminthes and the Xenoturbellida

The general fine-structural organization of statocysts in Catenulida, Nemertodermatida, Acoela, Proseriata, Lurus (Dalyellioida), and Xenoturbella are summarized. In lithophorous (statocyst-bearing) members of the Catenulida, the statocysts exhibit a few parietal cells and one or several movable statoliths within a spacious intracapsular cavity. Statocysts in the Nemertodermatida have several parietal cells and two lithocytes, each equipped with one statolith, whereas those of the other acoelomorphan taxon, the Acoela, always have two parietal cells and one movable lithocyte. The statocysts of lithophorous members of the Proseriata represent more sophisticated systems: each has two clusters of accessory cells in addition to several parietal cells and a voluminous lithocyte in which the statolith is movable. In catenulids and proseriates, processes of outer neurons penetrate the capsule of the statocyst, whereas such innervations have not been found in the Nemertodermatida and Acoela. I conclude that the different types of statocysts have evolved independently within the Plathelminthes. Xenoturbella displays an intraepidermal statocyst with many monociliary parietal cells and several mobile cells (lithocytes) within the central cavity of the statocyst. Each of these mobile cells carries a statolith-like structure and one prominent cilium. The statocyst of Xenoturbella does not correspond to any type of plathelminth statocyst.     

An Ultrastructural Account of Otoplanid Turbellaria Neuroanatomy II. The statocyst design: evolutionary and functional implications

The statocyst architecture in the three otoplanid species Notocaryoturbella bigermaria Lanfranchi, 1969, Otoplana truncaspina Lanfranchi, 1969 and Parotoplanella heterorhabditica Lanfranchi, 1969 is compared. Common features are: (a) a fibrillar collagen-like, 0.2 μm thick, investing capsule continuous with the brain capsule; (b) an inner wall made up of six or more flattened and overlapping parietal cells; (c) a statolith forming cell hanging from the dorsal side down in the lumen, with a large statolith containing vacuole; (d) a bilateral pair of spindle shaped accessory cell groups, adjoining the statolith cell and sending projections to the wall—nerve projections run through the capsule; (e) one accessory cell enveloping the other cells of the group has a filament containing cytoplasm, the filaments coverging into a hemidesmosome making contact with a projection coming from a parietal cell; (f) muscles from the longitudinal body musculature inserting onto the capsule externally. The lack of ciliary structures differentiates the turbellarian statocyst from the majority of invertebrate statocysts. The developmental origin, the phylogenetical meaning and the functional and adaptive value of the statocyst in Turbellaria are here commented on.     

A Fine Structural Analysis of the Statocyst in Turbellaria Acoela

Ferrero, E. (Istituto di Biologia Generale, Universith di Pisa, Pisa, Italy.) A fine structural analysis of the statocyst in Turbellaria Acoela. Zool. Scr. 2 (1): 5–16, 1973.—The fine structure of the statocyst components in the acoelan Convoluta psammophila is described, namely: capsule, parietal cells, lithocyte, and the statolith. The absence of ciliary structures, the highly developed endoplasmic reticulum of the lithocyte and the layered texture of the statolith are remarkable. A functional interpretation of the muscles inserted on the statocyst and of the nerve bundle running nearby is suggested. The morphological similarities and differences between the acoelan statocyst and the statocyst of lower and higher invertebrate phyla are discussed.     

Ultrastructure of the Statocysts in the Apodous Sea Cucumber Leptosynapta inhaerens (Holothuroidea, Echinodermata)

The burrowing sea cucumber Leptosynapta inhaerens possesses five pairs of statocysts, one pair on either side of each radial nerve cord where it arises from the circumoral nerve ring. The nerve cords exhibit only ectoneural components at the level of the statocysts. A sinus-like epineural canal lies superjacent to each cord. This canal is lined by a robust monociliated neuroepithelium which lacks any special support cells. Beneath the neuroepithelium, the somata of the ectoneural neurons form a perikaryal layer whereas the axons are located within the proximal parts of the cords. Glial cells have not been found. Each statocyst is a hollow sense organ. Its central cavity is lined by a monolayer of monociliated parietal cells. Axons of these parietal cells extend towards the statocyst nerve which connects each statocyst with the ectoneural pathways of the cord. A single lithocyte floats within each central statocyst cavity. This unciliated cell contains a voluminous vacuole with the statolith and several smaller vacuoles. It is concluded that statocysts do not belong to the basic organization of the Holothuroidea but have been evolved within this group. The statement, that the statocysts of apodous sea cucumbers and that of the enigmatic Xenoturbella bocki are homologous organs, is rejected.     

Fine structural study of the statocysts in the veliger larva of the nudibranch,Rostanga pulchra

Summary The two statocysts of the veliger larva of Rostanga pulchra are positioned within the base of the foot. They are spherical, fluid-filled capsule that contain a large, calcareous statolith and several smaller concretions. The epithelium of the statocyst is composed of 10 ciliated sensory cells (hair cells) and 11 accessory cells. The latter group stains darkly and includes 2 microvillous cells, 7 supporting cells, and 2 glial cells. The hair cells stain lightly and each gives rise to an axon; two types can be distinguished. The first type, in which a minimum of 3 cilia are randomly positioned on the apical cell membrane, is restricted to the upper portion of the statocyst. The second type, in which 9 to 11 cilia are arranged in a slightly curved row, is found exclusively around the base of the statocyst. Each statocyst is connected dorso-laterally to the ipsilateral cerebral ganglion by a short static nerve, formed by axons arising from the hair cells. Ganglionic neurons synapse with these axons as the static nerve enters the cerebral ganglion. The lumen of the statocyst is continuous with a blind constricted canal located beneath the static nerve.A diagram showing the structure of the statocyst and its association with the nervous system is presented. Possible functions of the statocyst in relation to larval behavior are discussed.     

Ultrastruktur der Statocyste von Ototyphlonemertes pallida (Keferstein, 1862) (Nemertini)

Zusammenfassung Ototyphlonemertes pallida (Keferstein, 1862) hat zwei Statocysten, die unmittelbar hinter den Dorsalganglien auf den verlängerten Ventralganglien liegen. Jede Statocyste besteht aus einer Statolithenkammerzelle, mehreren Nervenzellen und einer Anzahl Hüllzellen und ist von einer dicken Basalmembran umgeben. Die Statolithenkammerzelle umschließt in der Regel drei Statolithenkammern, die von einer doppelten Membran umgeben sind und untereinander in Verbindung stehen. Sie enthalten je einen frei beweglichen Statolithen. Cilien und Ciliarstrukturen fehlen. Auf der Dorsalseite der Statocyste liegen mehrere stark verästelte Nervenzellen, die einen gemeinsamen Strang bilden. In der Nähe der Statolithenkammerzelle spalten sie sich auf und bilden pro Kammer eine oder mehrere synaptische Platten mit elektrischen Synapsen. Die Statolithenkammerzelle wird von zahlreichen Hüllzellen umgeben, die durch Desmosomen fest verbunden und zusätzlich in der ventralen Hälfte der Statocyste an den Außenseiten stark miteinander verzahnt sind. Die Hüllzellen unterscheiden sich im Aufbau deutlich von den beiden anderen Zelltypen und sind nicht an der Reizperzeption oder Reizleitung beteiligt. Zu den cilienlosen Statocysten bei Coelenteraten, Turbellarien, Holothurien, Xenoturbella und Tunicaten-Larven bestehen keine engeren morphologischen Beziehungen. Die Statocyste von O. pallida stellt eine Bildung sui generis innerhalb der Nemertinen dar.

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Ultrastructure of the Statocyst of Ototyphlonemertes pallida (Keferstein, 1862) (Nemertini)

Summary Ototyphlonemertes pallida (Keferstein, 1862) has two statocysts, which are situated just behind the dorsal ganglions on the elongations of the ventral ganglions. Each statocyst consists of one statolith chamber cell, some nerve cells and a number of covering cells and is surrounded by a thick basement membrane. Usually the statolith chamber cell encloses three statolith chambers, which are intercommunicated and surrounded by a double membrane. Each chamber contains a single mobile statolith. Cilia and ciliary structures are lacking. Within the dorsal part of the statocyst some very ramified nerve cells are situated, which form a nerve fibre. In the vicinity of the statolith chamber cell the nerve cells split up into synaptical plates with electric synapses; there are one or several synaptical plates at the level of each chamber. The statolith chamber cell is surrounded by numerous covering cells, which are connected by desmosomes and additionally linked together at the outside in the ventral part of the statocyst. With regard to their structure the covering cells differ greatly from the other cell types, and they do not participate in impulse perception and impulse conduction. There do not exist any closer morphological relations to the statocysts lacking cilia in Coelenterates, Turbellaria, Holothuria, Xenoturbella and Tunicata. The statocyst of O. pallida represents an indigenous structure within the Nemerteans.

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Abkürzungen bm Basalmembran - d Dendrit - de Desmosom - dg Dorsalganglion - dm doppelte Membran der Statolithenkammer - ds deltaförmige Strukturen - ep Epidermis - es elektrische Synapse - hms Hautmuskelschlauch - hz Hüllzelle - k Zellkern - m Mitochondrium - ma abgewandeltes Mitochondrium - mu Muskulatur - n Nervenstrang - nv Neurosekretvesikel - nz Nervenzelle - rs Rüsselscheide - sc Statocyste - sk Statolithenkammer - skz Statolithenkammerzelle - sp synaptische Platte - st Statolith - v Vakuole - vg Ventralganglion     

Ultrastructural aspects of nervous-system and statocyst morphogenesis during embryonic development of Convoluta psammophila (Turbellaria,Acoela)

The notion that statocysts originated from an infolding of ectoderm lined by ciliated sensory cells has been challenged with evidence of capsule-limited, non-ciliary statocysts in several independent phyla. Statocysts in turbellarians primitively lack cilia and are embedded within or closely adjoined to the cerebral ganglion; they are likely to be derived from nervous tissue. We investigated the development of the simple statocyst in an acoel turbellarian, a statocyst consisting of three cells. Observations of serial TEM sections of embryos at different stages of development support the hypothesis of an inner (non-epithelial) origin of the statocyst. First, a three-cell complex is delimited by a basal lamina; it then undergoes cavitation by swelling, autophagy, and fluid secretion. The statocyst becomes discernible within the precursor ganglion cells while they still contain yolk inclusions. The two outer (parietal) cells, enclosed together by a 10-nm-thick basal lamina, arrange themselves in an ovoid of about 10 µm diameter and surround the inner statolith-forming cell. The statolith is formed later within vacuoles of the statolith-forming cell.     

Carbonic anhydrase is required for statoconia homeostasis in organ cultures of statocysts from Aplysia californica

A novel organ culture system has been developed to study the regulation of statoconia production in the gravity sensing organ in Aplysia californica. Statocysts were cultured in Leibovitz (L15) medium supplemented with salts and Aplysia haemolymph for four days at 17°C. The viability of the system was evaluated by examining four parameters: statocyst morphology, the activity of the mechanosensory cilia in the statocyst, production of new statoconia during culture and change in statoconia volume after culture. There were no morphological differences in statocysts before and after culture when ciliary beating was maintained. There was a 29% increase in the number of statoconia after four days in culture. Mean statocyst, statolith and statoconia volumes were not affected by culture conditions. The presence of carbonic anhydrase in the statocysts was shown using immunohistochemistry. When statocysts were cultured in the presence of 4.0 × 10^–4 M acetazolamide to inhibit the enzyme activity, there was a decrease in statoconia production and statoconia volume, indicating a role for this enzyme in statoconia homeostasis, potentially via pH regulation. These studies are the first to report a novel system for the culture of statocysts and show that carbonic anhydrase is involved in the regulation of statoconia volume and production.     

The Structure and Function of Müller Vesicles in Loxodid Ciliates1

ABSTRACT. The Müller vesicle is a characteristic organelle of loxodid ciliates. Its structure and development have been investigated using light microscopy and TEM. The organelle consists of a membrane-covered mineral body (the statolith), a vacuole, and various structures derived from the overlying kinety. There is strong evidence that the vesicle functions as a gravity sensor: a) its structure and relative dimensions fulfil the minimum requirements of a functional statocyst; b) its structure bears a close resemblance to the statocysts of some higher animals; c) re-orientation of the cell with respect to gravity produces a gravity-induced displacement of the mineral body, and d) geotaxis in Loxodes can be demonstrated experimentally. The transduction of the signal probably takes place at the level of the two kinetosomes of the organelle, one of which is in close contact with the cell membrane, while the other is connected to the statolith by a fairly rigid stalk containing a bundle of microtubules.     

Statocysts of hydromedusae

The statocysts of Leptomedusae are formed as a depression in the velum. They are lined on the inside towards the distal part of the velum by thin epithelium and towards the proximal part by ciliated sensory cells. Lithocytes are present in the centre. The concretion contains calcium sulphate and in some cases, calcium phosphate is also present in addition to some membranous material. The statocysts of Narcomedusae arise from the exumbrellar nerve ring as free sensory clubs. They have a proximal basal cushion of sensory cells from the centre of which arises a sensory club (Aegina) or a sensory papilla carrying a sensory club (Solmissus). The sensory club has an axial strand of endodermal cells covered by ciliated sensory cells. Some of the endodermal cells have a concretion. While the statocysts of Leptomedusae are totally ectodermal, those of Narcomedusae are ecto-endodermal in origin. The sensory cilia of Leptomedusae, especially those present on the sensory cells adjacent to the lithocyte, run close and parallel to the lithocyte membrane. In Narcomedusae the sensory cilia of the basal cusion and sensory papilla are tall and strong. Ciliary rootlets are missing in the sensory cilia of Leptomedusae and in the sensory club of Narcomedusae but they are strongly developed in the cilia of basal cusion and sensory papilla. The cilia have 9+2 filament content. A ring of stereocilia surrounds the kinocilium of the sensory club cells. Mechanism of statocyst function is discussed.     

Light- and electron-microscopic analysis of the statocyst of the American crayfishOrconectes limosus (Crustacea,Decapoda)

Summary The statocyst ofOrconectes limosus contains static hairs arranged in four groups. All the hairs are the same in basic structure; they differ only in length and diameter and in their positions with respect to the other hairs in the group and to the statolith. In terms of functional morphology, each static hair is part of a unit consisting of an acellular lever string, three receptor cells, a scolopale cell, sheath cells, and enveloping cells. The lever string comprises two components in a characteristic longitudinal arrangement. The structure of the receptor cells resembles that of the arthropod chemo- and mechanoreceptors studied previously. The cilium and the postciliary section lie within two receptor cavities, formed by the scolopale cell and the sheath cells; the two cavities communicate with one another. The receptor cells are fixed in position by various structures. Proximally they form desmosomes with the scolopale cell, medially they are joined by filaments to the inner wall of cavity 1, and distally they are retained by a constriction between the two cavities. Two possible stimulus-mediating mechanisms are discussed: pressure changes in the receptor cavities and shearing of the base of the cilia with respect to the preciliary region. The lever string is part of the cuticle and hence is shed during molting. Nevertheless, the statocyst remains functional during this process because new structural units are formed below the old cuticle prior to ecdysis.Abbreviations a axon - b bulb - bb basal body - c cilium - cu cuticle - d dendrite - de desmosome - dm dense material - ec enveloping cell - f fulcrum - h hair - hs hair shaft - ir inner row of hair group - l lingula - ls lever string - m mitochondrion - n nucleus - or outer row of hair group - pcd postciliary dilation - R1 receptor cavity 1 - R2 receptor cavity 2 - rc receptor cell - ro rootlet - s sheath cell - sc scolopale cell - st statoconium - t tooth     

Structure and function of the Nautilus statocyst.

The two equilibrium receptor organs (statocysts) of Nautilus are avoid sacks, half-filled with numerous small, free-moving statoconia and half with endolymph. The inner surface of each statocyst is lined with 130,000-150,000 primary sensory hair cells. The hair cells are of two morphological types. Type A hair cells carry 10-15 kinocilia arranged in a single ciliary row; they are present in the ventral half of the statocyst. Type B hair cells carry 8-10 irregularly arranged kinocilia; they are present in the dorsal half of the statocyst. Both type of hair cells are morphologically polarized. To test whether these features allow the Nautilus statocyst to sense angular accelerations, behavioural experiments were performed to measure statocyst-dependent funnel movements during sinusoidal oscillations of restrained Nautilus around a vertical body axis. Such dynamic rotatory stimulation caused horizontal phase-locked movements of the funnel. The funnel movements were either in the same direction (compensatory funnel response), or in the opposite direction (funnel follow response) to that of the applied rotation. Compensatory funnel movements were also seen during optokinetic stimulation (with a black and white stripe pattern) and during stimulations in which optokinetic and statocyst stimulations were combined. These morphological and behavioural findings show that the statocysts of Nautilus, in addition to their function as gravity receptor organs, are able to detect rotatory movements (angular accelerations) without the specialized receptor systems (crista/cupula systems) that are found in the statocysts of coleoid cephalopods. The findings further indicate that both statocyst and visual inputs control compensatory funnel movements.     

Ultrastructure of the jugular body of Rana pipiens

Summary The jugular bodies in adult Rana pipiens, are surrounded by a capsule of mesothelium and connective tissue, and their parenchyma consists of cell cords arranged in a sinusoidal network. The cell cords are formed by irregular reticular cells, showing numerous filaments and joined together by zonulae adhaerents. The intercellular spaces are filled by reticular fibres and free cells. These latter are small and medium lymphocytes, lymphoblasts, and developing and mature plasma cells. Additionally, free macrophages, neutrophils and acidophils also occur. Sinusoidal blood vessels show thin walls with numerous filaments and pinocytotic vesicles. They exhibit a discontinuous basement membrane, and tight junctions frequently occur between endothelial cells. Occasionally, lymphatic vessels are found and the innervation is principally vasomotor, although nerve endings appear remarkably near reticular cells and lymphocytes. The jugular bodies of adult R. pipiens are plasma cell and antibody-forming organs, whose functional significance is discussed in relation to their ultrastructural organization.     

The structure of some cephalopod statoliths

Summary The statoliths of Sepia officinalis, Octopus vulgaris, Alloteuthis subulata and Taonius megalops have a smooth outline, but an irregular shape. They have projections and indentations. The statoliths from a pair of statocysts are usually quite similar in size and shape, and the general pattern is probably maintained throughout the size range of the species. Statoliths from large animals are marginally larger than those from smaller ones. The statolith usually occupies only a small part of the cavity of the statocyst, and it is situated in the anterior part of the statocyst. They are joined to the macula by hairs extending from it. These hairs are very delicate and easily broken during preparation of the specimens. The hairs are much longer and narrower than the receptor cilia of the macula. The receptor cilia are enclosed within holes in the tangled hairlike anchoring fibrils.The statolith is made up of crystalline subunits, the statoconia. The crystals vary in size, they are usually elongated, hexagonal with pointed ends. The statolith consists of a closely packed mass of these crystals, sometimes they are irregularly arranged, where in others they are stacked with their long axes parallel. In Sepia officinalis and Taonius megalops, the crystals are arranged in regular shaped packets and these packets of crystals are stacked together. These larger subunits are not always arranged in a regular way, and their major axes can be organised in several different ways. The size and outline of these large subunits do vary in different parts of the statolith.The external surface of the statolith is macroscopically smooth. Over some parts there is a surface layer covering the rod-like crystals that make up the major bulk of the stone. In other regions, the surface is rough at a microscopic level, the roughness is produced by the exposed ends of the filamentous crystals. The crystals are composed of calcium carbonate in the form of aragonite.I wish to thank Professor J.Z. Young, FRS, for considerable help, advice and encouragement throughout this study. Dr. A. Boyde generously allowed me to use his scanning electron microscope and gave freely of his expertise and time. Dr. J. Fitch kindly gave me some fossil statoliths and Dr. J. Elliott examined them with his x-ray diffraction apparatus. Dr. Marion Nixon helped me to collect and prepare the specimens. Mrs. E. Bailey, Miss P. Stephens and Mr. R. Moss provided the expert technical assistance     

A transmission electron microscopic investigation of the sensory vesicle in the brain of Oikopleura dioica (Appendicularia)

Summary The structure of three cell types in the sensory vesicle is described: (1) The statocyte, with its intracellular statolith, is attached to the medial wall of the vesicle via delicate shaft cells. (2) Cells along the dorsal, ventral and lateral walls which contact the surface of the statocyte with long, slender cilia. These cells are presumed to be primary sensory cells. (3) Presumed secretory cells, along the rostral and dorsal walls, may have a dual function: (a) secretion of the vesicle fluid, and (b) stabilization of the wall by turgor created in characteristic intercellular cavities. The sensory vesicle in Oikopleura contains undoubtedly typical statocyst components adequate for a free-swimming animal, whereas the ascidian system is suggested to be a device that responds to gravitational stimuli and, together with temporary photoreceptors, aids the larva in finding optimal settling conditions.     

Hair Cell Interactions in the Statocyst of Hermissenda

Hair cells in the statocyst of Hermissenda crassicornis respond to mechanical stimulation with a short latency (<2 ms) depolarizing generator potential that is followed by hyperpolarization and inhibition of spike activity. Mechanically evoked hyperpolarization and spike inhibition were abolished by cutting the static nerve, repetitive mechanical stimulation, tetrodotoxin (TTX), and Co⁺⁺. Since none of these procedures markedly altered the generator potential it was concluded that the hyperpolarization is an inhibitory synaptic potential and not a component of the mechanotransduction process. Intracellular recordings from pairs of hair cells in the same statocyst and in statocysts on opposite sides of the brain revealed that hair cells are connected by chemical and/or electrical synapses. All chemical interactions were inhibitory. Hyperpolarization and spike inhibition result from inhibitory interactions between hair cells in the same and in opposite statocysts.     

Ultrastructure of the ovary of <Emphasis Type="Italic">Amphilina japonica</Emphasis> Goto &; Ishii, 1936 (Cestoda) and its implications for phylogenetic studies

The ultrastructure of the ovary of the amphilinidean cestode Amphilina japonica Goto & Ishii, 1936 from the body-cavity of the American sturgeon Acipenser transmontanus Richardson is described using transmission electron microscopy. The characters of the ovary of Amphilina japonica are different from those of all other cestodes. The most important difference is in the nature of the relationship between the germ and accessory cells within the ovary. In A. japonica the oocytes and accessory cells form numerous different intercellular contacts (desmosome-like junctions and zonulae adherentes). Gap junctions are present between the narrow cytoplasmic processes of the accessory cells. Numerous micropinocytotic vesicles and vacuoles from the accessory cells discharge their content into spaces between the oocytes and the accessory cells. The accessory cells are closely associated with the oocytes during the early and middle stages of oogenesis. As the volume of oocytes increases, the accessory cells gradually lose their association with the oocyte surfaces. Peripherally located individual accessory cells of A. japonica give rise to a cellular epithelial layer of irregular shape and thickness which breaks down via numerous invaginations of the basal membrane and underlying basal matrix. The different arrangements of the interconnection of cell components in the Amphilinidea compared with the Gyrocotylidea and Eucestoda (the absence of specialised cell contacts and the syncytial nature of the accessory ‘interstitial’ cells) are evidence suggesting the presence of unrelated groups within the Cestoda. The nature of the association of the accessory and germ cells in ovary of A. japonica more closely resembles the ovary of non-platyhelminth invertebrates rather than that of other neodermatans.     

Effects of pH on asexual reproduction and statolith formation of the scyphozoan, <Emphasis Type="Italic">Aurelia labiata</Emphasis>

Although anthropogenic influences such as global warming, overfishing, and eutrophication may contribute to jellyfish blooms, little is known about the effects of ocean acidification on jellyfish. Most medusae form statoliths of calcium sulfate hemihydrate that are components of their balance organs (statocysts). This study was designed to test the effects of pH (7.9, within the average current range, 7.5, expected by 2100, and 7.2, expected by 2300) combined with two temperatures (9 and 15°C) on asexual reproduction and statolith formation of the moon jellyfish, Aurelia labiata. Polyp survival was 100% after 122 d in seawater in all six temperature and pH combinations. Because few polyps at 9°C strobilated, and temperature effects on budding were consistent with published results, we did not analyze data from those three treatments further. At 15°C, there were no significant effects of pH on the numbers of ephyrae or buds produced per polyp or on the numbers of statoliths per statocyst; however, statolith size was significantly smaller in ephyrae released from polyps reared at low pH. Our results indicate that A. labiata polyps are quite tolerant of low pH, surviving and reproducing asexually even at the lowest tested pH; however, the effects of small statoliths on ephyra fitness are unknown. Future research on the behavior of ephyrae with small statoliths would further our understanding of how ocean acidification may affect jellyfish survival in nature.     

Fine structure of the statocyst sensilla of the mysid shrimp Neomysis integer (Leach, 1814) (Crustacea,Mysidacea)

The fine structure of the statocyst sensilla of Neomysis integer was investigated. The statocyst contains about 35 sensilla, which are composed of two bipolar sensory cells, nine enveloping cells, and a seta. The sensory cells consist of an axon, a perikaryon, and a dendrite. The dendrite contains a proximal segment with a ciliary rootlet and at least one basal body, and a distal segment with a ciliary axoneme (9 × 2 + 0) at its base. The distal segment extends along the peripheral wall of the seta and is in close contact with the wall of the hair shaft. The enveloping cells surround the proximal and distal segments of the dendrite. The innermost enveloping cell contains a scolopale rod. It surrounds the receptor lymph cavity and secretes flocculent material into this cavity. From the tip of the cell a dendritic sheath, which encloses the distal segment of the dendrite, emerges. A peculiar feature of the second enveloping cell is the presence of a scolopale-like rod, which is more slender and less pronounced than in the first enveloping cell. The seta consists of three parts: a socket, a tubular midpart, and a gutter-like apical part, the tip of which penetrates into the statolith. The seta shows over its full length a bilaterally symmetrical axis that is coplanar with the plane in which the seta is bent toward the statolith. The structure of the seta and the position of the distal segments provide morphological evidence for directional sensitivity of the sensilla and for the magnitude of shear on the setal wall being an adequate stimulus.     

ULTRASTRUCTURE DES STATOCYSTES CHEZ SCROBICULARIA PLANA ET TELLINA TENUIS (MOLLUSQUES LAMELLIBRANCHES TELLINACEA)

The statocysts of Scrobicularia plana and Tellina tenuis arecomposed of 8 sensory hair-cells separated by giant cells characterizedby a fibrous cytoskeleton. The sensory cells bear few ciliaespecially in S. plana. These cilia are responsible both forrotation of the statolith and for the transduction of stimuli.Their basal bodies bear a side rootlet facing a basal foot anda crown composed of 9 spokes. Such statocysts seem to receiveonly multidirectional stimuli and to allow a less diversifiedbehaviour than the statocysts of Pecten. (Received 16 September 1980;