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.     

Catecholamine-containing,serotonin-like and neuropeptide FMRFamide-like immunoreactive cells and processes in the nervous system of the pilidium larva (Nemertini)

Summary Pilidium larvae at different developmental stages have been investigated for the occurrence of glyoxylic acid induced fluorescence in catecholamines (CA), and serotonin-like (5-HT) and neuropeptide FMRFamide-like (FMRFamide) immunoreactivity (ir). The distribution of CA, 5-HT-ir and FMRFamide-ir cells and processes was compared with the location of nerve processes as found by transmission electron microscopy (TEM). In the pilidium larvae the marginal and oral nerves contain CA and 5-HT-ir processes and 5-HT-ir unipolar cells. The posterior suboral nerve contain CA and 5-HT-ir processes, whereas in the anterior suboral nerve neither CA nor 5-HT-ir and FMRFamide-ir were observed. The lateral helmet nerve contains FMRFamide-ir processes and unipolar cells. In the epidermis CA and 5-HT-ir multipolar cells were found. The juvenile worm that develops inside the pilidium larva was found to contain only 5-HT-ir. A pair of lateral cords extent the whole length of the juvenile and anteriorly they form the anterior ventral cerebral commissure. Also, from the anterior part of the lateral cords projects a pair of circumrhynchodeal processes which dorsally form the dorsal cerebral commissure. A pair of proboscis processes originate from the circumrhynchodeal processes and extend the whole length of the probosics. From the dorsal cerebral commissure cephalic processes project rostrally and ventrally. Only unipolar 5-HT-ir cells were observed, and they were located along the lateral cords into which their processes extend.Abbreviations AEC 3-amino-9-ethylcarbazole - ap apical plate - arp anterior accessory ridge processes - ason anterior suboral nerve - CA catecholamines - cd cephalic discs - cp cephalic processes - crp circumrhynchodeal processes - DAB 3,3'-diaminobenzidine - dc dorsal cerebral commissure - epi epidermis - es oesophagus - fl fore lobe - FMRFamide phe—met—arg—phe—NH₂ - Go goat - GS goat serum - hl hind lobe - int intestine of the juvenile - lhn lateral helmet nerve - lhp lateral helmet processes - ll lateral lobe - lp lateral processes of the juvenile - mcb marginal ciliary band - me mesoderm - mn marginal nerve - moc monociliary cell - mp marginal processes - mu muscle - muc multiciliary cell - n 1, n 2, n 3 division of marginal nerve - on oral nerve - op oral processes - pb proboscis - pp proboscis processes - pson posterior suboral nerve - psop posterior suboral processes - Ra rabbit - sd stomodeum - st stomach - td trunk discs - tr trunk - TRITC tetramethylrhodamine isothiocyanate - vc ventral cerebral commissure - z 1, z 2 ciliary zones of marginal ciliary band - 5-HT serotonin     

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.     

Distribution of catecholamine-containing,serotonin-like and neuropeptide FMRFamide-like immunoreactive neurons and processes in the nervous system of the actinotroch larva ofPhoronis muelleri (Phoronida)

Summary Glyoxylic-acid-induced fluorescence of catecholamines and antibodies against serotonin and FMRFamide were used to study the distribution of putative neurotransmitters in the actinotroch larva ofPhoronis muelleri Selys-Longchamps, 1903. Catecholamines occur in the neuropile of the apical ganglion, in the longitudinal median epistome nerves, in the epistome marginal nerves, and in the nerve along the bases of the tentacles. The tentacles have laterofrontal and latero-abfrontal bundles of processes that form two minor nerves along the lateral ciliary band of the tentacles, and a medio-frontal bundle of processes. Monopolar cells are located on the ventro-lateral part of the mesosome. Processes are located along the posterior ciliary band and as a reticulum in the epidermis. Serotonin-like immunoreactive cells and processes are located in the apical ganglion, in the longitudinal median epistome nerves, and as a dorsal and ventral pair of bundles along the tentacle bases. Processes from the latter extend into the tentacles as the medioabfrontal processes. The latero-abfrontal processes form a minor nerve along the ciliary band. The dorsal bundles forms the major nerve ring along the tentacles and processes extend from it to the metasome. Processes are located along the posterior ciliary band. FMRFamide-like immunoreactive cells and processes are found in the apical ganglion, in the longitudinal median epistome nerves and as a pair of lateral epistome processes projecting towards the ring of tentacles. In the tentacles, a pair of latero-frontal processes are found; these form a minor nerve along the ciliary band. A band of cells can be seen along the tentacle ring.     

Structure of the brachiopod lophophore

Data on the development, structure, and functional morphology of the brachiopod lophophore are analyzed. The common origin of the tentacle apparatus in Lophophorata from the postoral ciliary band of the larva is shown. The brachiopod lophophore is based on the brachial axis consisting of the brachial fold running along the row of tentacles. The brachial axis may be attached to the brachial (dorsal) mantle lobe (trocholophe, schizolophe, and ptycholophe lophophores) or extend freely into the mantle cavity to form coiling brachia (spirolophe, zygolophe, and plectolophe lophophores). The circulation of water flows through the mantle cavity in the brachiopods with attached and free lophophores is described. A new hypothesis on the sorting of particles suspended in water during filtration is proposed.     

Neuromuscular development in Novocrania anomala: evidence for the presence of serotonin and a spiralian-like apical organ in lecithotrophic brachiopod larvae

SUMMARY The phylogenetic position of Brachiopoda remains unsettled, and only few recent data on brachiopod organogenesis are currently available. In order to contribute data to questions concerning brachiopod ontogeny and evolution we investigated nervous and muscle system development in the craniiform (inarticulate) brachiopod Novocrania anomala . Larvae of this species are lecithotrophic and have a bilobed body with three pairs of dorsal setal bundles that emerge from the posterior lobe. Fully developed larvae exhibit a network of setae pouch muscles as well as medioventral longitudinal and transversal muscles. After settlement, the anterior and posterior adductor muscles and delicate mantle retractor muscles begin to form. Comparison of the larval muscular system of Novocrania anomala with that of rhynchonelliform (articulate) brachiopod larvae shows that the former has a much simpler muscular organization. The first signal of serotonin-like immunoreactivity appears in fully developed Novocrania anomala larvae, which have an apical organ that consists of four flask-shaped cells and two ventral neurites. These ventral neurites do not stain positively for the axonal marker α-tubulin in the larval stages. In the juveniles, the nervous system stained by α-tubulin is characterized by two ventral neurite bundles with three commissures. Our data are the first direct proof for the presence of an immunoreactive neurotransmitter in lecithotrophic brachiopod larvae and demonstrate the existence of flask-shaped serotonergic cells in the brachiopod larval apical organ, thus significantly increasing the probability that this cell type was part of the bauplan of the larvae of the last common lophotrochozoan ancestor.     

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.     

Anti-tubulin labeling reveals ampullary neuron ciliary bundles in opisthobranch larvae and a new putative neural structure associated with the apical ganglion

This investigation examines tubulin labeling associated with the apical ganglion in a variety of planktotrophic and lecithotrophic opisthobranch larvae. Emphasis is on the ampullary neurons, in which ciliary bundles within the ampulla are strongly labeled. The larvae of all but one species have five ampullary neurons and their associated ciliary bundles. The anaspid Phyllaplysia taylori, a species with direct development and an encapsulated veliger stage, has only four ampullary neurons. The cilia-containing ampulla extends to the pretrochal surface via a long, narrow canal that opens to the external environment through a very small pore (0.1 microm diameter). Cilia within the canal were never observed to project beyond the opening of the apical pore. The ampullary canals extend toward and are grouped with the ciliary tuft cells and remain in this location as planktotrophic larvae feed and grow. If, as has been reported, the ciliary tuft is motile, the pores may be continually bathed in fresh seawater. Such an arrangement would increase sensitivity to environmental chemical stimuli if the suggested chemosensory function of these neurons is correct. In general, ciliary bundles of newly hatched veligers are smaller in planktotrophic larvae than in lecithotrophic larvae. In planktotrophic larvae of Melibe leonina, the ciliary bundles increase in length and width as the veligers feed and grow. This may be related to an increase in sensitivity for whatever sensory function these neurons fulfill. An unexpected tubulin-labeled structure, tentatively called the apical nerve, was also found to be associated with the apical ganglion. This putative nerve extends from the region of the visceral organs to a position either within or adjacent to the apical ganglion. One function of the apical nerve might be to convey the stimulus resulting from metamorphic induction to the visceral organs.     

Architecture and function of the lophophore in the problematic brachiopod Heliomedusa orienta (Early Cambrian, South China)

The detailed structure of the lophophore is a key diagnostic character in the definition of higher brachiopod taxa. The problematic Heliomedusa orienta Sun and Hou, from the Lower Cambrian Chengjiang Lagerstätte of Yunnan, southwestern China, has a well-preserved lophophore, which is unlike that of any known extant or extinct brachiopods. Based on a comparative study of lophophore disposition in H. orienta and the extant discinid Pelagodiscus atlanticus, the in- and excurrent pattern and shell orientation of H. orienta are described and discussed. Reconstructions of lophophore shape and function are based on numerous specimens and comparison with P. atlanticus. The lophophore is composed of a pair of lophophoral arms that freely arch posteriorly rather than coiling anteriorly as commonly seen in fossil and recent lingulids. The lophophore is attached to the dorsal lobe of the mantle; it has neither calcareous nor chitinous supporting structures, and is disposed symmetrically on either side of the valve midline. The mouth can be inferred to be located at the base of the two brachial tubes, slightly posterior to the anterodorsal projection of the body wall. The lophophoral arms bear laterofrontal tentacles with a double row of cilia along their lateral edge, as in extant lingulid brachiopods. The main brachial axes are also ciliated, which presumably facilitated transport of mucous-bound nutrient particles to the mouth. The unique organization of the lophophore in Heliomedusa is not like any known fossil and living brachiopods. This clearly demonstrates that H. orienta is not a member of any crown group. It is here considered as a member of the brachiopod stem group, which challenges recent interpretations of a close discinid affinity.     

Mapping of serotonin-immunoreactive neurons in the larval nervous system of the flies Calliphora erythrocephala and Sarcophaga bullata

Summary Serotonin-immunoreactive (5-HTi) neurons were mapped in the larval central nervous system (CNS) of the dipterous flies Calliphora erythrocephala and Sarcophaga bullata. Immunocytochemistry was performed on cryostat sections, paraffin sections, and on the entire CNS (whole mounts).The CNS of larvae displays 96–98 5-HTi cell bodies. The location of the cell bodies within the segmental cerebral and ventral ganglia is consistent among individuals. The pattern of immunoreactive fibers in tracts and within neuropil regions of the CNS was resolved in detail. Some 5-HTi neurons in the CNS possess axons that run through peripheral nerves (antenno-labro-frontal nerves).The suboesophagealand thoracico-abdominal ganglia of the adult blowflies were studied for a comparison with the larval ventral ganglia. In the thoracico-abdominal ganglia of adults the same number of 5-HTi cell bodies was found as in the larvae except in the metathoracic ganglion, which in the adult contains two cell bodies less than in the larva. The immunoreactive processes within the neuropil of the adult thoracico-abdominal ganglia form more elaborate patterns than those of the larvae, but the basic organization of major fiber tracts was similar in larval and adult ganglia. Some aspects of postembryonic development are discussed in relation to the transformation of the distribution of 5-HTi neurons and their processes into the adult pattern.     

The phylogenetic position of entoprocta, ectoprocta, phoronida, and brachiopoda

Ectoprocts, phoronids and brachiopods are often dealt with underthe heading Tentaculata or Lophophorata, sometimes with entoproctsdiscussed in the same chapter, for example in Ruppert and Barnes(1994). The Lophophorata is purported to be held together bythe presence of a "lophophore," a mesosomal tentacle crown withan upstream-collecting ciliary band. However, the mesosomaltentacle crown of pterobranchs has upstream-collecting ciliarybands with monociliate cells, similar to those of phoronidsand brachiopods, although its ontogeny is not well documented.On the contrary, the ectoproct tentacle crown carries a ciliarysieving system with multiciliate cells and the body does notshow archimery, neither during ontogeny nor during budding,so the tentacles cannot be characterized as mesosomal. The entoproctshave tentacles without coelomic canals and with a downstream-collectingciliary system like that of trochophore larvae and adult rotifersand serpulid and sabellid annelids. Planktotrophic phoronidand brachiopod larvae develop tentacles at an early stage, buttheir ciliary system resembles those of echinoderm and enteropneustlarvae. Ectoproct larvae are generally non-feeding, but theplanktotrophic cyphonautes larvae of certain gymnolaemates havea ciliary band resembling that of the adult tentacles. The entoproctshave typical trochophore larvae and many feed with downstream-collectingciliary bands. Phoronids and brachiopods are thus morphologicallyon the deuterostome line, probably as the sister group of the"Neorenalia" or Deuterostomia sensu stricto. The entoproctsare clearly spiralians, although their more precise positionhas not been determined. The position of the ectoprocts is uncertain,but nothing in their morphology indicates deuterostome affinities."Lophophorata" is thus a polyphyletic assemblage and the wordshould disappear from the zoological vocabulary, just as "Vermes"disappeared many years ago.     

Toward locating the source of serotonergic axons in the tail nerve of <Emphasis Type="Italic">Aplysia</Emphasis>

Stimulation of the tail nerve (pedal nerve 9, p9) of the mollusk, Aplysia californica, causes release of serotonin (5-HT), which mediates sensitization of withdrawal responses. There are about 35 serotonin-immunoreactive (5-HT-ir) axons in p9, yet the cell bodies of these axons have not been located. Backfills of p9 were combined with 5-HT immunohistochemistry to locate the cell bodies of 5-HT-ir neurons with axons in p9. About 100 neurons had axons in p9. Only about ten neurons, however, were both backfilled and 5-HT-ir. These double-labeled neurons were all located in the pedal ganglion associated with p9, which had a total of approximately 42 5-HT-ir somata. The discrepancy between the number of 5-HT-ir axons and double-labeled cell bodies is not likely due to neurons having multiple axons in the nerve; intracellular fills suggest that these neurons do not branch before entering p9. Additionally, no evidence was found for peripheral 5-HT-ir cell bodies that project axons centrally through p9. Thus, approximately 70% of the neurons that give rise to the 5-HT-ir axons in tail nerve are unaccounted for, but likely to reside in the pedal ganglion.     

Anatomy and innervation of the abdominal segmental muscles in larval and adult Tenebrio molitor (Coleoptera)

The external morphology, musculature, and the innervation of the abdominal segments were examined in larvae and adult Tenebrio molitor. In the larva, there are 26 pairs of muscles arranged at four different levels in the ventral, lateral, and dorsal region of each segment. In the adult, the number of muscles has been dramatically reduced and is limited to six pairs of muscles located at the dorsal and lateral region of the segment. These muscles, in either larval or adult stages, are innervated by two main nerves, n1 and n2, which originate from the segmental ganglia. The cell bodies of the motoneurons innervating the muscles of the 3rd abdominal segment are located in the 3rd and 2nd abdominal ganglia. Some cell bodies are retained throughout metamorphosis, but others disappear during the larva-pupa transition.     

Comparison of the neuromuscular systems among actinotroch larvae: systematic and evolutionary implications

A comparative analysis of the larval and presumptive juvenile neuromuscular systems among actinotroch larvae was performed using confocal laser microscopy with probes for F-actin and serotonin. Currently, there are two main categories of larval nervous systems based on the origin of the nerve fibers that innervate the larval tentacles. Characteristics of the serotonergic cells of the larval apical ganglion and juvenile nervous system have remained relatively conserved, but the structure of the secondary (hood) sense organ and the juvenile tentacles has diversified among species. Differences in larval musculature are mainly associated with differences in hood morphology. The presumptive, juvenile neuromuscular system is either integrated or separated from that of the larva based on the origin of the juvenile tentacles. Among species, the juvenile tentacles are made by remodeling the larval tentacles, developed from a basal tentacular thickening, or developed as a completely separate set in the larva. Differentiation of the neuromuscular structures of the juvenile tentacles is more diverse than their outward morphological characteristics would suggest. Importance of these larval characters is discussed in terms of current problems that exist within phoronid systematics. Evolutionary implications of these morphological characters are discussed among the phoronids, brachiopods, and related bilaterians. Overall, the integration or separation of larval and juvenile neuromuscular characters may yield insights into the evolution of lophotrochozoan body plans.     

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.     

Three-dimensional reconstruction of the musculature of Cossura pygodactylata Jones, 1956 (Annelida: Cossuridae)

The musculature of adult specimens of Cossura pygodactylata was studied by means of F-actin labelling and confocal laser scanning microscopy (CLSM). Their body wall is comprised of five longitudinal muscle bands: two dorsal, two ventral and one ventromedial. Complete circular fibres are found only in the abdominal region, and they are developed only on the border of the segments. Thoracic and posterior body regions contain only transverse fibres ending near the ventral longitudinal bands. Almost-complete rings of transverse muscles, with gaps on the dorsal and ventral sides, surround the terminal part of the pygidium. Four longitudinal bands go to the middle of the prostomium and 5–14 paired dorso-ventral muscle fibres arise in its distal part. Each buccal tentacle contains one thick and two thin longitudinal muscle filaments; thick muscle fibres from all tentacles merge, forming left and right tentacle protractors rooted in the dorsal longitudinal bands of the body wall. The circumbuccal complex includes well-developed upper and lower lips. These lips contain an outer layer of transverse fibres, and the lower lip also contains inner oblique muscles going to the dorsal longitudinal bands. The branchial filament contains two longitudinal muscle fibres that do not connect with the body musculature. The parapodial complex includes strong intersegmental and segmental oblique muscles in the thoracic region only; chaetal retractors, protractors and muscles of the body wall are present in all body regions. Muscle fibres are developed in the dorsal and ventral mesenteries. One semi-circular fibre is developed on the border of each segment and is most likely embedded in the dissepiment. The intestine has thin circular fibres along its full length. The dorsal blood vessel has strong muscle fibres that cover its anterior part, which is called the heart. It consists of short longitudinal elements forming regular rings and inner partitions. The musculature of C. pygodactylata includes some elements that are homologous with similar muscular components in other polychaetes (i.e., the body wall and most parapodial muscles) and several unique features, mostly at the anterior end.     

On the organization of the lophophore in phoronids (Lophophorata: Phoronida)

The organization of the lophophore is the main feature used for the identification of phoronid species. The structure of the lophophore and tentacles in seven phoronid species (Phoronis ovalis, P. ijimai, P. hippocrepia, P. svetlanae, P. australis, Phoronopsis harmeri, and Ph. malakhovi) collected in different areas of the World Ocean was studied. Two new patterns of the phoronid lophophore structure were found: “transition to horseshoe-shaped” (as in P. ovalis from Aniva Bay and in P. ijimai from the coast of Iturup Island, Sea of Okhotsk) and “transition to spiral” (in burrowing specimens P. hippocrepia from Aniva Bay, P. svetlanae and Ph. harmeri from Vostok Bay, Sea of Japan). For the first time it was shown that phoronid species with different types of the lophophore structure possess different kinds of tentacles. Thus, five types of phoronid tentacles were identified that vary in the shape of their cross section: rounded, oval, ellipsoid, rectangular, and skittle-shaped. A correlation was found between lophophore organization and the type of tentacles in phoronids. A table of the correlation between body size, lophophore organization, tentacle structure, and mode of life in different phoronid species is proposed.     

Rejection Mechanism of Plectolophous Lophophore of Brachiopod <Emphasis Type="Italic">Coptothyris grayi</Emphasis> (Terebratulida,Rhynchonelliformea)

Brachiopoda is a relict group of invertebrate filter feeders that used a tentacle organ, lophophore, for capturing food particles from the water column. Brachiopod extinction apparently occurred due to low productivity of their filtering organ in comparison with more advanced filter-feeders. Investigation of the filtering mechanism of modern brachiopods is essential to understanding their evolutionary fate. This study is devoted to the rejection mechanism of large waste particles from the plectolophous lophophore of brachiopod Coptothyris grayi. The waste particles gather inside of the lophophore on the outer side of the brachial fold. The particles form rows along frontal grooves of outer tentacles and are carried successively to the tentacle tips and move along them, slimed by mucus. One portion of the particles comes off the lophophore and falls down the mantle, while another part is carried to the abfrontal surface of the tentacles. Due to repeated reversals of abfrontal cilia, the particles wavily move along the abfrontal surface of tentacles. Such movement contributes to the secretion of mucus and the formation of particle clots. The clots come off the lophophore and fall down the mantle. The particles are transported along the mantle by cilia to the anterior part of the mantle margin. Here the ciliary reversals that facilitate secretion of mucus and formation of pseudofeces also take place. The latter takes away from the mantle cavity. Thus, only outer tentacles participate in the rejection of large waste particles from the lophophore. Ciliary reversals of the abfrontal surface of tentacles and the mantle are discovered in brachiopods for the first time. This facilitates the additional secretion of mucus and formation of pseudofeces, easing their exit from the mantle cavity. The results contribute to the knowledge of lophophore function and evolution of tentacle organs in Bilateria.     

Structure and metamorphic remodeling of the larval nervous system and musculature of Phoronis pallida (Phoronida)

The structure of the larval nervous system and the musculature of Phoronis pallida were studied, as well as the remodeling of these systems at metamorphosis. The serotonergic portion of the apical ganglion is a U-shaped field of cell bodies that send projections into a central neuropil. The majority of the serotonergic cells are (at least) bipolar sensory cells, and a few are nonsensory cells. Catecholaminergic cell bodies border the apical ganglion. The second (hood) sense organ develops at competence and is composed of bipolar sensory cells that send projections into a secondary neuropil. Musculature of the competent larva includes circular and longitudinal muscle fibers of the body wall, as well as elevators and depressors of the tentacles and hood. The juvenile nervous system and musculature are developed prior to metamorphosis and are integrated with those of the larva. Components of the juvenile nervous system include a diffuse neural net of serotonergic cell bodies and fibers and longitudinal catecholaminergic fibers. The juvenile body wall musculature consists of longitudinal fibers that overlie circular muscle fibers, except in the cincture regions, where this pattern is reversed. Metamorphosis is initiated by the larval neuromuscular system but is completed by the juvenile neuromuscular system. During metamorphosis, the larval nervous system and the musculature undergo cell death, and the larval tentacles and gut are remodeled into the juvenile arrangement. Although the phoronid nervous system has often been described as deuterostome-like, these data show that several cytological aspects of the larval and juvenile neuromuscular systems also have protostome (lophotrochozoan) characteristics.