Inferences concerning the evolution of invertebrate nervous systems are often hampered by the lack of a solid data base for little known but phylogenetically crucial taxa. In order to contribute to the discussion concerning the ancestral neural pattern of the Lophotrochozoa (a major clade that includes a number of phyla that exhibit a ciliated larva in their life cycle), we investigated neurogenesis in Phoronopsis harmeri, a member of the poorly studied Phoronida, by using antibody staining against serotonin and FMRFamide in combination with confocal microscopy and 3D reconstruction software.

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.

The Phoronida, which is one of the smallest phyla of invertebrates, includes only 13 valid species, although ~40 larval forms of phoronids were recently described. This report uses light microscopy and molecular methods to describe a new phoronid species, Phoronis embryolabi Temereva & Chichvarkhin, sp. nov. The morphology of P. embryolabi, which lives commensally in the burrows of Axiidea shrimp Nihonotrypaea japonica in Vostok Bay (the Sea of Japan), is extremely similar to that of Phoronis pallida Silen, 1952; the bodies of both species exhibit specific regionalisation. However, the organisation of the metanephridia differs between P. pallida and P. embryolabi. Moreover, P. embryolabi has a unique type of development, viviparity, in which mothers release fully developed larvae into the environment. In all other phoronid species, the spawning occurs as a release of fertilised eggs or early embryos. Viviparity of completely developed larvae has not been previously described for any phoronid. According to analysis of partial 28S rRNA, P. embryolabi is close to Phoronis pallida. On the other hand, analysis of partial cytochrome c oxidase subunit I indicated a unique position of P. embryolabi among phoronids. These results should be used for revision of phoronid taxonomy (i.e. the type of development should be considered as characteristic of subgenera within the genus Phoronis). This report also establishes the relationship between P. embryolabi and an Actinotrocha sp. that was described in a previous paper.

Phoronids are marine invertebrates which great significance from the perspective of comparative anatomy. Phoronids have traditionally been described as deuterostomes because they exhibit developmental and larval traits similar to those traits found in echinoderms and hemichordates. However, molecular phylogenetic evidence has consistently identified the phoronids and brachiopods as a monophyletic group within the lophotrochozoan protostomes. The nature of egg cleavage, gastrulation and the origin of coelomic mesoderm can help to resolve this contradiction. These questions were investigated with Phoronopsis harmeri. Embryos and actinotroch larvae were cultivated in the laboratory, and embryonic and larval development was followed with SEM, light and video microscopy. Egg cleavage is radial, although the furrows of the 4th and succeeding divisions are oblique, which allows for blastula formation by the 16-cell stage. The ciliated blastula hatches 10 h after the onset of cleavage and acquires the apical tuft. Gastrulation proceeds by invagination. Coelomic mesoderm derives from two, the anterior and posterior, precursors. The anterior precursor forms by cell migration from the anterior wall of the archenteron, the posterior one by enterocoelic outpouching of the midgut. The lining of preoral coelom and tentacle coelom is derived from the anterior precursor, whilst the lining of trunk coelom originates from the posterior one. The intestine forms as an ectodermal invagination of the posterior part of larval body. Protonephridial canals form as ectodermal dorso-lateral furrows on both sides of the anus. A typical actinotroch is formed 100 h after the onset of cleavage. Larvae are described at the stages of 8, 12,20 and 24 tentacles.

The Bryozoa (=Ectoprocta) is a large group of bilaterians that exhibit great variability in the innervation of tentacles and in the organization of the cerebral ganglion. Investigations of bryozoans from different groups may contribute to the reconstruction of the bryozoan nervous system bauplan. A detailed investigation of the polypide nervous system of the ctenostome bryozoan Amathia gracilis is reported here. The cerebral ganglion displays prominent zonality and has at least three zones: proximal, central, and distal. The proximal zone is the most developed and contains two large perikarya giving rise to the tentacle sheath nerves. The neuroepithelial organization of the cerebral ganglion is revealed. The tiny lumen of the cerebral ganglion is represented by narrow spaces between the apical projections of the perikarya of the central zone. The cerebral ganglion gives rise to five groups of main neurite bundles of the lophophore and the tentacle sheath: the circum-oral nerve ring, the lophophoral dorso-lateral nerves, the pharyngeal and visceral neurite bundles, the outer nerve ring, and the tentacle sheath nerves. Serotonin-like immunoreactive nerve system of polypide includes eight large perikarya located between tentacles bases. There are two analmost and six oralmost perikarya with prominent serotonergic “gap” between them. Based on the characteristics of their innervations, the tentacles can be subdivided into two groups: four that are near the anus and six that are near the mouth. Two longitudinal neurite bundles - medio-frontal and abfrontal - extend along each tentacle. The zonality of the cerebral ganglion, the presence of three commissures, and location of the main nerves emanating from each zone might have caused by directive innervation of the various parts of the body: the tentacles sheath, the lophohpore, and the digestive tract. Two alternative scenarios of bryozoan lophophore evolution are discussed. The arrangement of large serotonin-like immunoreactive perikarya differs from the pattern previously described in ctenostome bryozoans. In accordance with its position relative to the same organs (tentacles, anus, and mouth), the lophophore outer nerve ring corresponds to the brachiopod lower brachial nerve and to the phoronid tentacular nerve ring. The presence of the outer nerve ring makes the lophophore innervation within the group (clade) of lophophorates similar and provides additional morphological evidence of the lophophore homology and monophyly of the lophophorates.

The validity of the Lophophorata as a monophyletic group remains controversial. New data on the innervation of the lophophore, which is a unique feature of the lophophorates, may help clarify the status of the Lophophorata and provide new information on the early evolution of the group. In this paper, the organization of the nervous system of the lophophore is described in adults of the minute phoronid Phoronis ovalis. The lophophore nervous system includes a dorsal ganglion, a tentacular nerve ring, an inner ganglion, an inner nerve ring, and six nerves in each tentacle. The inner ganglion and inner nerve ring, which is associated with sensory cells, are described for the first time in adult phoronids. The general plan of the nervous system of the lophophore and tentacles is similar in P. ovalis and bryozoans. These new results suggest the presence of two nerve centers and two nerve rings in the last common ancestor of phoronids and bryozoans. During evolution, bryozoans may have lost the outer nerve center and outer nerve ring, whereas phoronids may have lost the inner nerve center and inner nerve ring. These morphological results evidence the lophophorates are monophyletic.