The nervous system of the actinotroch larva of Phoronis muelleri (Phoronida)

Summary The nervous system of the actinotroch larva of Phoronis muelleri has been investigated with the transmission electron microscope (TEM). Attempts have been made to localize all of the major nerves and to reveal the cytoarchitecture of the apical ganglion. The nervous system is intraepithelial in position and consists of an apical ganglion, located on the epistome, with at least four different cell types, including monopolar sensory cells and mono- or multipolar neuron-like cells. From the anterior part of the apical ganglion three median nerves extend to the edge of the epistome; two of these nerves connect to nerves which follow the edge of the epistome all the way to the junction of the epistome and the mesosome. From the posterior part of the ganglion extend two lateral nerves which continue along the tentacular ring. Each tentacle has three nerves located on the frontal side which connect to the nerve ring along the tentacles. Along the posterior ciliary band is a minor nerve ring. In addition, a nerve net is found on the epistome, mesosome, and metasome, but no longitudinal nerves were observed between the posterior ciliary band and the apical ganglion. All nerve cells were found in the apical ganglion and none was observed along the nerves. Sensory cells (probably mechanoreceptors) are located in two rows on each tentacle; sensory organs such as eyes and statocysts were not observed.Abbreviations ac accessory centricle - aen anterior epistome edge nerve - af abfrontal cells - bl basal lamina - bl.c blastocoel coelomocyte - ci cilium - co collar - cp cell process - cr ciliary root - ec ₁ epistome edge cell type 1 - mne mouth nerve ring - mo mouth - mp metasomal pouch - ms mesosome - mt metasome - mu muscle - n nerve process - ne nerve - np neuropil - nu nucleus - pc ₁ posterior ciliary band cell type 1 - ec ₂ epistome edge cell type 2 - ec ₃ epistome edge cell type 3 - epi epidermis - es epistome - ese epistome edge - fc frontal cell - gc ₁ type 1 ganglion cells - gc ₂ type 2 ganglion cells - gc ₃ type 3 ganglion cells - ge gut epithelium - ij intermediate junction - laf lateroabfrontal cell - lc lateral cell - lfc laterofrontal cell - lgc lateral ganglion cell - me metacoel epithelium - lne longitudinal median epistome nerves - pc ₂ posterior ciliary band cell type 2 - pc procoel - pe procoel epithelium - pen posterior epistome edge nerve - pr posterior ciliary band - p.rec proximal recess of procoel epithelium - prne nerve ring along posterior ciliary band - sj septate junction - sne secondary nerve along the tentacular ring - t tentacle - tr tentacular ring - trne horseshoe-shaped nerve along the tentacular ring     

Modern Data on the Innervation of the Lophophore in Lingula anatina (Brachiopoda) Support the Monophyly of the Lophophorates

Evolutionary relationships among members of the Lophophorata remain unclear. Traditionally, the Lophophorata included three phyla: Brachiopoda, Bryozoa or Ectoprocta, and Phoronida. All species in these phyla have a lophophore, which is regarded as a homologous structure of the lophophorates. Because the organization of the nervous system has been traditionally used to establish relationships among groups of animals, information on the organization of the nervous system in the lophophore of phoronids, brachiopods, and bryozoans may help clarify relationships among the lophophorates. In the current study, the innervation of the lophophore of the inarticulate brachiopod Lingula anatina is investigated by modern methods. The lophophore of L. anatina contains three brachial nerves: the main, accessory, and lower brachial nerves. The main brachial nerve is located at the base of the dorsal side of the brachial fold and gives rise to the cross neurite bundles, which pass through the connective tissue and connect the main and accessory brachial nerves. Nerves emanating from the accessory brachial nerve account for most of the tentacle innervation and comprise the frontal, latero-frontal, and latero-abfrontal neurite bundles. The lower brachial nerve gives rise to the abfrontal neurite bundles of the outer tentacles. Comparative analysis revealed the presence of many similar features in the organization of the lophophore nervous system in phoronids, brachiopods, and bryozoans. The main brachial nerve of L. anatina is similar to the dorsal ganglion of phoronids and the cerebral ganglion of bryozoans. The accessory brachial nerve of L. anatina is similar to the minor nerve ring of phoronids and the circumoral nerve ring of bryozoans. All lophophorates have intertentacular neurite bundles, which innervate adjacent tentacles. The presence of similar nerve elements in the lophophore of phoronids, brachiopods, and bryozoans supports the homology of the lophophore and the monophyly of the lophophorates.     

Neurobiology of the gorgonian coelenterates,Murcea californca and Lophogorgia chilensis

Summary Diffuse and synaptic nerve nets are present in the coenenchymal mesoglea and ectoderm of Muricea and Lophogorgia colonies. The nerve nets extend into the polyp column and tentacles maintaining a subectodermalmesogleal position. The density of nerve elements is low in comparison with similar nerve nets found in pennatulids.In the column of the polyp anthocodium, and throughout the oral disk region, neurons cross the mesoglea and enter the polyp endoderm. These neurons presumably connect with the endodermal nerve net which innervates the septal musculature. The trans-mesogleal neurons probably represent the connection between colonial and polyp nervous systems.In the tentacles, longitudinal ectodermal musculature is present with an overlying nerve plexus. These muscles and nerves, as well as tentacular sensory cells, are well represented in the oral side of the tentacles only.Presumed sensory cells form ciliary cone complexes in which one cell possesses an apical cilium. The other cells as well as the centrally located nematocyte contribute microvilli to the cone. The basal portion of the sensory cells is drawn into one or more neurite-like processes which enter the ectodermal nerve plexus. Similar processes form synapses with longitudinal muscle cells and nematocytes. The sensory cells of the ciliary cones presumably include chemoreceptors which can activate or modify nematocyst discharge, local muscle twitches, and tentacle bending.This work was supported by Office of Naval Research Contract N00014-75-C-0242, NSF Grant BMS 74-23242 and General Research Funds of the University of California, Santa Barbara. We wish to thank Dr. Steven K. Fisher for the use of facilities in his lab. This paper is part of a thesis to be submitted by R.A.S. to the Department of Biological Sciences, University of California, Santa Barbara in partial fulfillment of the requirements for the Ph. D.     

Myoanatomy and serotonergic nervous system of plumatellid and fredericellid phylactolaemata (lophotrochozoa,ectoprocta)

The phylogenetic position of the Ectoprocta within the Lophotrochozoa is discussed controversially. For gaining more insight into ectoproct relationships and comparing it with other potentially related phyla, we analysed the myoanatomy and serotonergic nervous system of adult representatives of the Phylactolaemata (Plumatella emarginata, Plumatellavaihiriae, Plumatella fungosa, Fredericella sultana). The bodywall contains a mesh of circular and longitudinal muscles. On its distal end, the orifice possesses a prominent sphincter and continues into the vestibular wall, which has longitudinal and circular musculature. The tentacle sheath carries mostly longitudinal muscle fibres in Plumatella sp., whereas F. sultana also possesses regular circular muscle fibres. Three groups of muscles are associated with the lophophore: 1) Lophophoral arm muscles (missing in Fredericella), 2) epistome musculature and 3) tentacle musculature. The epistome flap is encompassed by smooth muscle fibres. A few fibres extend medially over the ganglion to its proximal floor. Abfrontal tentacle muscles have diagonally arranged muscle fibres in their proximal region, whereas the distal region is formed by a stack of muscles that resemble an inverted ‘V’. Frontal tentacle muscles show more variation and either possess one or two bases. The digestive tract possesses circular musculature which is striated except at the intestine where it is composed of smooth muscle fibres. The serotonergic nervous system is concentrated in the cerebral ganglion. From the latter a serotonergic nerve extends to each tentacle base. In Plumatella the inner row of tentacles at the lophophoral concavity lacks serotonergic nerves. Bodywall musculature is a common feature in many lophotrochozoan phyla, but among other filter feeders like the Ectoprocta is only present in the ‘lophophorate’ Phoronida. The longitudinal tentacle musculature is reminiscent of the condition found in phoronids and brachiopods, but differs to entoproct tentacles. Although this study shows some support for the ‘Lophophorata’, more comparative analyses of possibly related phyla are required. J. Morphol., 2011. © 2011 Wiley Periodicals, Inc.     

Ultrastructure and immunocytochemistry of the nervous system of the larvae ofLingula anatina andGlottidia sp. (Brachiopoda)

Summary Planktotrophic brachiopod larvae ofGlottidia sp. have been investigated for the occurrence of glyoxylic acid induced fluorescence in catecholamines (CA), and serotonin-like (5-HT) and neuropeptide FMRFamidelike (FMRFamide) immunoreactivity (ir). The location of CA, 5-HT-ir and FMRFamide-ir cells and processes were compared with the location of neurons and nerve processes found by transmission electron microscopy. The apical ganglion contains 5-HT-ir and FMRFamideir cells and processes and CA processes. From the dorsal part of the apical ganglion extend dorsal 5-HT-ir and FMRFamide-ir processes; from the nine pairs of tentacles stage (9. pt) they project to the ventral ganglion. These dorsal lophophore processes follow themusculus lophophoralis and them. brachialis. The 5-HT-ir and some of the FMRFamide-ir processes project along the muscles to the tentacles. From the ventral part of the apical ganglion extend CA, 5-HT-ir and FMRFamide-ir processes which follow the ciliary band of the lophophore and project to the tentacles. An intense band of CA processes was also observed in the lophophore, but the dorsal/ventral location could not be ascertained. The ventral ganglion contains 5-HT-ir and FMRFamide-ir cells which project either caudally on the metasome or rostrally as part of the dorsal lophophore processes. The neuropil of the ventral ganglion contains CA, 5-HT-ir and FMRFamide-ir processes. The nervous system of the planktotrophic brachiopod larvae seems to consist of a ventral lophophore system innervating the ciliary bands and a dorsal lophophore system including the ventral ganglion innervating the body musculature. The latter system develops later in ontogeny and is regarded as a specialization due to the presence of shells and associated musculature. The former system is regarded as homologous with the nervous system of actinotroch larvae (Phoronida) and planktotrophic larvae of the echinoderms.     

Immunocytochemistry of the neuromuscular systems of Loxosomella vivipara and L. parguerensis (Entoprocta: Loxosomatidae)

Little detailed information exists on the anatomy of the nervous system and the musculature of Entoprocta. Herein we describe the distribution of the neurotransmitters RFamide and serotonin as well as the myo-anatomy of adults and asexually produced budding stages of the solitary entoproct species Loxosomella vivipara and L. parguerensis using immunocytochemistry and epifluorescence as well as confocal microscopy. The development of the RFamidergic and serotonergic nervous system starts in early budding stages. In the adults, RFamide is present in the bilateral symmetric cerebral ganglion, a pair of oral nerves that innervate two pairs of nerve cell clusters in the heel of the foot, a pair of aboral nerves, the paired lateral nerves, the calyx nerves, the atrial ring nerve, the tentacle nerves, the stomach nerves, and the rectal nerves. Serotonin is only found in the cerebral ganglion, the oral nerves, and in the tentacle nerves. Some differences in the distribution of both neurotransmitters were found between L. vivipara and L. parguerensis and are most obvious in the differing number of large serotonergic perikarya associated with the oral nerves. Nerves arising from the cerebral ganglion and running in a ventral direction have not been described for Entoprocta before, and the homology of these to the ventral nerve cords of other Spiralia is considered possible. The body musculature of both Loxosomella species comprises longitudinal and diagonal muscles in the foot, the stalk, and the calyx. We found several circular muscles in the calyx. The stalk and parts of the foot and the calyx are surrounded by a fine outer layer of ring muscles. In addition to the congruent details regarding the myo-anatomy of both species, species-specific muscle structures could be revealed. The comparison of our data with recent findings of the myo-anatomy of two Loxosoma species indicates that longitudinal and diagonal body muscles, atrial ring muscles, tentacle muscles, esophageal and rectal ring muscles, as well as intestinal and anal sphincters are probably part of the ancestral entoproct muscle bauplan.     

The Larval Nervous System of Polygordius lacteus Scheinder, 1868 (Polygordiidae,Polychaeta): Immunocytochemical Data

The nervous system of the planktotrophic trochophore larva of Polygordius lacteus has been investigated using antibodies to serotonin (5-HT) and the neuropeptide FMRFamide. The apical ganglion contains three 5-HT-ir neurons, many FMRFamide-ir neurons and a tripartate 5-HT-ir and FMRFamide-ir neuropil. A lateral nerve extends from each side of the apical ganglion across the episphere and the ventral hyposphere, where the two nerves combine to form the paired ventral nerve cord. These nerves have both 5-HT-ir and FMRFamide-ir processes. Three circumferential nerves are associated with the ciliary bands: two prototroch and one metatroch nerve. All contain 5-HT-ir and FMRFamide-ir processes. An oral nerve plexus also contain both 5-HT-ir and FMRFamide-ir processes develops from the metatroch nerve, and an esophageal ring of FMRFamide-ir processes develops in later larval stages. In young stages the ventral ganglion contains two 5-HT-ir and two FMRFamide-ir perikarya; during development the ventral ganglion grows caudally and adds additional 5-HR-ir and FMRFamide-ir perikarya. These are the only perikarya that could be found along the lateral nerve and ventral nerve cord. The telotroch nerve develops from the ventral nerve cord. The 5-HT-ir and FMRFamide-ir part of the nervous system is strictly bilateral symmetric. and much of the system (i.e. apical ganglion, lateral nerves ventral nerve cord, dorsal nerve and oral plexus) is retained in the adult.     

The nervous system in the cyclostome bryozoan <Emphasis Type="Italic">Crisia eburnea</Emphasis> as revealed by transmission electron and confocal laser scanning microscopy

Introduction

Among bryozoans, cyclostome anatomy is the least studied by modern methods. New data on the nervous system fill the gap in our knowledge and make morphological analysis much more fruitful to resolve some questions of bryozoan evolution and phylogeny.

Results

The nervous system of cyclostome Crisia eburnea was studied by transmission electron microscopy and confocal laser scanning microscopy. The cerebral ganglion has an upper concavity and a small inner cavity filled with cilia and microvilli, thus exhibiting features of neuroepithelium. The cerebral ganglion is associated with the circumoral nerve ring, the circumpharyngeal nerve ring, and the outer nerve ring. Each tentacle has six longitudinal neurite bundles. The body wall is innervated by thick paired longitudinal nerves. Circular nerves are associated with atrial sphincter. A membranous sac, cardia, and caecum all have nervous plexus.

Conclusion

The nervous system of the cyclostome C. eburnea combines phylactolaemate and gymnolaemate features. Innervation of tentacles by six neurite bundles is similar of that in Phylactolaemata. The presence of circumpharyngeal nerve ring and outer nerve ring is characteristic of both, Cyclostomata and Gymnolaemata. The structure of the cerebral ganglion may be regarded as a result of transformation of hypothetical ancestral neuroepithelium. Primitive cerebral ganglion and combination of nerve plexus and cords in the nervous system of C. eburnea allows to suggest that the nerve system topography of C. eburnea may represent an ancestral state of nervous system organization in Bryozoa. Several scenarios describing evolution of the cerebral ganglion in different bryozoan groups are proposed.

     

Neuromuscular structure of the larva to early ancestrula stages of the cyclostome bryozoan Crisia eburnea

For the first time, the development of a cyclostome bryozoan has been studied with immunochemistry and confocal laser scanning microscopy, with emphasis on nerves and muscles. The larva is covered by multiciliated cells, which are latitudinally strongly elongated and show phalloidin-stained cell junctions. We hypothesize that these cells contract at metamorphosis and squeeze the apical invagination and the adhesive sac out. Ectodermal, longitudinal muscle cells extend from the cells of the inner, conical cuticularized part of the apical invagination to the lower part of the corona, around the adhesive sac pore. These muscles are retained in the ancestrula. Scattered monociliated nerve cells are interspersed between the coronal ciliary cells. An equatorial nerve in the larva disappears at metamorphosis. The central, conical part of the cuticle becomes the terminal membrane of the ancestrula, and the underlying ectodermal and mesodermal cell layers differentiate into the polypide bud, forming a deep narrow invagination, differentiating into vestibule–atrium, mouth ring and pharynx–stomach–rectum. Tentacles develop from the ring of cells around the mouth, and a small ganglion with four nerves innervating each of the tentacles develops at the anal side of the mouth. These new findings yield further support for previous homology statements of bryozoan larvae and development.     

Ciliary and nervous structures in juvenile females of the annelid Dinophilus gyrociliatus (O. Schmidt, 1848) (Annelida: Polychaeta)

Ciliary and nerve structures were described in juvenile female Dinophilus gyrociliatus (O. Schmidt, 1848) after immunochemical staining with tubulin, serotonin, and FMRFamide antibodies. Anti-tubulin antibodies revealed the following external structures: two head and seven body ciliary bands, a ventral ciliary band, and head ciliary fields. Gut cilia and five pairs of protonephridia were detected inside the body. The nervous system consists of an oval headed neuropile with anterior and posterior nerves extending from it, seven longitudinal nerve cords, commissures, and circular nerves. Anti-serotonin antibodies revealed the head neuropile, neurons at the base of the ventral ciliary band, an oesophageal ring, and seven longitudinal ventral cords. Anti-FMRFamide antibodies revealed approximately ten neurons in the cerebral ganglion, five longitudinal cords, and the oesophageal and caudal-nerve rings. The presented data suggest the simplification of the nervous system structure in D. gyrociliatus, which probably reflects pedomorphosis.     

Development of the tentacular apparatus in sipunculans (Sipuncula): I. Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920)

The development and arrangement of the tentacular apparatus of Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920) are described and illustrated using scanning electron microscopy. In T. nigra, the tentacular apparatus is composed of two crowns: the nuchal arc enclosing the nuchal organ and a crown of numerous oral tentacles arranged in U-shaped festoons. In early juveniles, two dorsal horn-like protrusions develop into the first, or primary, pair of tentacles of the nuchal arc. The second pair of tentacles of the nuchal arc develops dorsolaterally on the bases of the primary tentacles. Two ventrolateral lobes of the oral disk grow and become subdivided by the longitudinal ciliary groove into anlages of one set of dorsal and one set of ventral tentacles, thus forming a first oral festoon. Later, a pair of dorsolateral lobes develop between the first festoons and the nuchal arc to form a second pair of oral festoons. The third and following pairs of oral festoons develop in the dorsolateral growth zones lateral to the borders of the nuchal arc, where they meet the oral crown. The growing festoons extend down the oral disk and run alongside the head. A new oral tentacle appears directly at/on the base of the previous tentacle, thus giving rise to a typical sympodium with an alternating arrangement of tentacles. In T. pyroides, a second pair of tentacles develops from two ciliary lobes that are ventrolateral outgrowths of the circumoral ciliary field around the terminal mouth opening. The third pair of tentacles appears from the dorsolateral lobes at the base of primary tentacles, between the first two pairs of tentacles. These six tentacles determine the position of six main stems of the tentacular apparatus designated the first tentacles in the corresponding stems. The second tentacle in every stem appears as a ventrolateral outgrowth at the base of the first tentacle. The third and following tentacles in the stem are developed between the two previous tentacles according to a sympodial pattern. In both species, the distinct sympodial pattern in the arrangement of tentacles in the tentacular apparatus is well evidenced by the outlines of the ciliary oral grooves. The branched stems of T. pyroides may be homologized structurally and functionally to the oral festoons of T. nigra. J. Morphol. (c) 2006 Wiley-Liss, Inc.     

The fine structure of the buccal tentacles of Holothuria forskali (Echinodermata,Holothuroidea)

Summary Ultrastructural study of the buccal tentacles of Holothuria forskali revealed that each tentacle bears numerous apical papillae. Each papilla consists of several differentiated sensory buds.The epidermis of the buds is composed of three cell types, i.e. mucus cells, ciliated cells, and glandular vesicular cells (GV cells). The GV cells have apical microvilli; they contain bundles of cross striated fibrillae associated with microtubules. Ciliated cells have a short non-motile cilium. Bud epidermal cells intimately contact an epineural nervous plate which is located slightly above the basement membrane of the epidermis. The epineural plate of each bud connects with the hyponeural nerve plexus of the tentacle. This nerve plexus consists of an axonic meshwork surrounded in places by sheath cells. The buccal tentacles have well-developed mesothelial muscles. Direct innervation of these muscles by the hyponeural nerve plexus was not seen.It is suggested that the buccal tentacles of H. forskali are sensory organs. They would recognize the organically richest areas of the sediment surface through the chemosensitive abilities of their apical buds. Tentacles presumably trap particles by wedging them between their buds and papillae.     

Morphological characteristics of chemosensory,visual, and statocyst pathways in the snailHelix lucorum

The following structural characteristics of the chemosensory, visual, and vestibular pathways of the snail (Helix lucorum) were demonstrated by using a variety of histological techniques. Large and small neurons of the tentacle ganglion, the bipolar cells of the olfactory nerve, and a proportion of optic tentacle bulb chemoreceptors within the olfactory nerve all send their processes to the CNS of the mollusk. Here they are divided up into numerous bundles of fibers in the neuropil of the ipsilateral cerebral ganglion. They are joined by processes from the central nervous system put out by all neurons of the protocerebrum and the cluster of cells of the commissural section of the metacerebrum. Ocular receptors do not send processes down below the enlargement of the upper optic nerve. This enlargement is also the site where processes from cells within the CNS and the nerve itself terminate. An area of arborization of processes from the visual pathway cells is located in the neuropil of the pleural portion of the metacerebrum. Hair cells of statocysts put out processes to the cerebral ganglion, whence axons of small metacerebral neurons extend towards the organ of balance. Some processes from vestibular pathway cells form an arborization zone at the ipsilateral cerebral ganglion, while others pass through the cerebral commissure to form their area of arborization in the contralateral ganglion. Processes from vestibular and visual pathway cells arborize in exactly the same area.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 18, No. 1, pp. 7–16, January–February, 1986.     

Nervous system development of the sea cucumber Stichopus japonicus

The nervous system development of the sea cucumber Stichopus japonicus was investigated to explore the development of the bilateral larval nervous system into the pentaradial adult form typical of echinoderms. The first nerve cells were detected in the apical region of epidermis in the late gastrula. In the auricularia larvae, nerve tracts were seen along the ciliary band. There was a pair of bilateral apical ganglia consisted of serotonergic nerve cells lined along the ciliary bands. During the transition to the doliolaria larvae, the nerve tracts rearranged together with the ciliary bands, but they were not segmented and remained continuous. The doliolaria larvae possessed nerves along the ciliary rings but strongly retained the features of auricularia larvae nerve pattern. The adult nervous system began to develop inside the doliolaria larvae before the larval nervous system disappears. None of the larval nervous system was observed to be incorporated into the adult nervous system with immunohistochemistry. Since S. japonicus are known to possess an ancestral mode of development for echinoderms, these results suggest that the larval nervous system and the adult nervous system were probably formed independently in the last common ancestor of echinoderms.     

Ultrastructure of tentacles in the sipunculid worm <Emphasis Type="Italic">Thysanocardia nigra</Emphasis> Ikeda, 1904 (Sipuncula)

The ultrastructure of the tentacles was studied in the sipunculid worm Thysanocardia nigra. Flexible digitate tentacles are arranged into the dorsal and ventral tentacular crowns at the anterior end of the introvert of Th. nigra. The tentacle bears oral, lateral, and aboral rows of cilia; on the oral side, there is a longitudinal groove. Each tentacle contains two oral tentacular canals and an aboral tentacular canal. The oral side of the tentacle is covered by a simple columnar epithelium, which contains large glandular cells that secrete their products onto the apical surface of the epithelium. The lateral and aboral epithelia are composed of cuboidal and flattened cells. The tentacular canals are lined with a flattened coelomic epithelium that consists of podocytes with their processes and multiciliated cells. The tentacular canals are continuous with the radial coelomic canals of the head and constitute the terminal parts of the tentacular coelom, which shows a highly complex morphology. Five tentacular nerves and circular and longitudinal muscle bands lie in the connective tissue of the tentacle wall. Similarities and differences in the tentacle morphology between Th. nigra and other sipunculan species are discussed.Original Russian Text Copyright © 2005 by Biologiya Morya, Maiorova, Adrianov.     

Putative sensory structures in marine bryozoans

Abstract. SEM studies of 21 species of marine bryozoans demonstrated that the abfrontal side of the tentacles bears a row of mono- or multiciliated cells, which are presumably sensory. In stenolaemates, the abfrontal cells, as well as the cells at the tentacle tips and the laterofrontal cells, are monociliated. In the 17 gymnolaemate species studied, each tentacle tip bears at least 3 multiciliated cells, each with a tuft of 5–7 stiff cilia of various lengths. On the abfrontal tentacle surface, mono- and multiciliated cells alternate, but all species studied have multiciliated cells at the base and the tip of each tentacle. In live animals, single cilia perform occasional flicks, whereas the tufts of 7–15 cilia on the multiciliated cells are immotile. Length and number of abfrontal cilia vary between species. Two types of multiciliated, putative sensory organs were found on the introvert of some gymnolaemates. One has an apical knob surrounded by a ring of cilia; the other has an apical tuft of cilia. The ultrastructure of the sensory cells of tentacles and introvert was studied in Rhamphostomella ovata . Our observations on both fixed and living material all suggest that these cells are primitive mechanoreceptors. The few species lacking ciliary structures on the introvert have long proximal ciliary tufts on the abfrontal tentacle surface.     

Neuronal background of positioning of the posterior tentacles in the snail Helix pomatia

The location of cerebral neurons innervating the three recently described flexor muscles involved in the orientation of the posterior tentacles was investigated by applying parallel retrograde Co- and Ni-lysine tracing via the olfactory and the peritentacular nerves. Their innervation patterns in the flexor muscles were studied by applying anterograde neurobiotin tracings via these nerves. The labeled neurons are clustered in eight groups in the cerebral ganglion. They send both common and distinct innervation pathways to the flexor and the tegumental muscles and to the tentacular retractor muscle. The common pathway reaches the muscles via the olfactory nerve, whereas the distinct pathways innervate via the internal and external peritentacular nerves. The three anchoring points of the three flexor muscles at the base of the tentacle outline the directions of three force vectors generated by the contraction of the muscles and enable the protracted tentacle to bend around a basal pivot. In the light of earlier physiological and the present anatomical findings, we suggest that the common innervation pathway to the muscles is required for tentacle withdrawal and the retractor mechanism, whereas the distinct pathways primarily serve the bending of the protracted posterior tentacles during foraging.     

The fine structure of the newly discovered propodial ganglia of the veliger larva of the nudibranch Onchidoris bilamellata

Summary Two pairs of ganglia are found in the propodial region of the veliger of Onchidoris bilamellata: the anterolateral pair is located at the foremost corners of the propodium, and the frontal pair is located beside the propodial midline. Both sets of ganglia are positioned below the epidermis, and they are joined to the cerebral ganglia by large, common connectives. Each ganglion possesses sensory cells, nerve cells and sheath cells, and the frontal pair contains a complement of secretory cells. Externally, the propodial ganglia are manifested as sensory fields. The fields of the anterolateral pair are elliptical in shape, and each appears as a band of cilia bordering an unciliated zone. The region devoid of cilia is composed of ordinary epidermal cells, whereas the ciliated portion is comprised of dendritic endings originating from cells in the ganglion. Dendrites arise from one type of sensory cell and pass through the epidermis in bundles. Each dendrite terminates as a single cilium at the epidermal surface. Sensory fields of the frontal ganglia are key-shaped and oppose one another on the anterior end of the foot. Each field appears as a flat, circular, unciliated region which extends into a ciliated groove that runs dorsally toward the mouth. The groove contains the terminals of secretory cells, ciliated sensory cells, and the cell bodies of nonciliated sensory cells. The nonciliated sensory cells, characterized by a microvillous apex, are the dominant cells in the flattened circular zone. The space between the frontal ganglia and the epidermis is bridged by bundles of processes which are similar to those of the anterolateral ganglia. However, these tracts contain collections of the apical processes of secretory cells, the dendrites of ciliated sensory cells, and the axons of nonciliated sensory cells. Morphological and behavioral evidence indicates that the propodial ganglia serve a chemosensory function during settlement and metamorphosis.