This article presents an original hypothesis of the origin of vertebrate limbs. According to this hypothesis, the tentacles and limbs of Bilateria originate from two circles of tentacles of the coelenterate ancestor, the labial circle and marginal circle. In Bilateria, the tentacles of the coelenterate ancestor gave rise to the metameric processes of mesodermal somites. In vertebrates, fin radials are established as metameric processes of somites. Taking account that chordates (including vertebrates) are inverted relative to other bilaterians, the homologs of labial tentacles are radials of unpaired median fin fold and the homologs of marginal tentacles are radials of paired lateral fin folds. Current data on Hox gene expression and paleontological finds indicating the homology of the digits of the Tetrapoda limb to the fin radials of cartilaginous fishes and lobe-finned fishes are discussed. This makes it possible to stretch the thread of homology from the tentacles of coelenterates to the digits of vertebrates.

The brachiopods are sessile invertebrates with an unusual blood system, which consists of a long-branched dorsal vessel. It is still unknown how blood circulates in this system. In the present study, for the first time we propose the circulation of blood in brachiopod Hemithiris psittacea based on morphological and experimental data. The main heart is located on the dorsal side of the stomach and divides the dorsal vessel into anterior and posterior parts. The anterior part enters the lophophore, where it gives off blind branches to each tentacle. The posterior part passes by the funnels of the nephridia and forms a blindly closed network in the gonads. We suggest that the circulation of blood includes three successive stages. During the first phase of systole of the main heart, blood flows through the anterior dorsal vessel. During the second phase of systole, blood flows through the posterior dorsal vessel. During diastole, blood flows from the anterior and posterior vessels and fills the main heart. The origin of a peculiar blood system in brachiopods can be explained by reduction of the ventral vessel, which is probably correlates with the reduction of the ventral side of the brachiopod ancestor's body. Another peculiarity of brachiopod blood system is the presence of an ampullar heart, which functions as a blood depot and allows blood to move in the vessels in two directions in an oscillatory mode. The brachiopod blood system contains vessels lacking true endothelium and can be classified as an "incompletely closed" type.

The paper considers the origin of segmentation, tentacles, limbs, and ciliary bands of larvae of Deuterostomia. The analysis of the expression of regulatory genes (along with the data of classical comparative anatomy) allows us to prove the homology of the pharyngeal plane of Anthozoa and the sagittal plane of triploblatic Bilateria and to confirm the origin of the coelomic segments from the peripheral gastric pockets of coelenterone. At the same time, an end-to-end homology of the segments is assumed within the Bilateria. The homology of the preoral segment and the tentacular segment in all Bilateria is emphasized. It is assumed that the ancestors of Bilateria had two circles of tentacles, labial and marginal. The marginal tentacles of the coelenterates are homologous to the metameric appendages of Bilateria, and the labial tentacles are homologous to the perioral ciliated tentacles and the ventral ciliated sole. The ciliary bands of larvae are homologous to the labial ciliated tentacles transferred to the larval stage. Possible homologues of metameric appendages and ciliated tentacles in Deuterostomia are discussed. In chordates (taking into account the “upside-down theory” of their origin), the radials of the median fin are considered as homologues of the labial tentacles, while the radials of paired fins are considered as homologues of the marginal tentacles. Two-phase expression of Hox genes allows us to prove the homology of tetrapod digits and fin radials of primitive fish. The similarity of the genetic mechanisms regulating the development of limbs makes it possible to stretch the threads of homology from the coelenterate tentacles to the limbs of vertebrates.

Worm-like endosymbionts were found in the hepatic region of the digestive tract of the deep-sea acorn worm Quatuoralisia malakhovi Ezhova et Lukinykh, 2022 (family Torquaratoridae) from the Bering Sea. The symbionts were assigned to the taxon Nemertodermatida on the basis of histological examination. Torquaratoridae are similar in feeding type to holothuroids, which have also been found to have Xenacoelomorpha endosymbionts.

Breakages of sequential Hox expression in Echinodermata may reflect evolutionary changes in the body plan. We quantified Hox gene expression at early larval stages, i.e., blastula at 13 hpf, gastrula at 35 hpf, prism stage at 46 hpf, and echinoplutei at 4 and 9 dpf, in the sea urchin with indirect development, Strongylocentrotus intermedius. The design of specific primers for real-time PCR (q-RT-PCR) for S. intermedius was performed. At blastula stage, only SiHox7 (medial Hox group), SiHox11/13b and SiHox11/13c (posterior Hox group) were expressed. At gastrula and prism stages, the expression of SiHox genes was lower than that in unfertilized eggs. Significant increase in the expression of all SiHox genes was observed in 4-day pluteus with further increase of SiHox1, SiHox8, SiHox9/10, and SiHox11/13b expression in 9-day pluteus. In other echinoderms, the genes of anterior Hox group are nearly not expressed in the early stages of development. At blastula, gastrula and prism stages, the genes of medial Hox group (mainly Hox7 and Hox8) and posterior Hox group (mainly Hox11/13a, 11/13b, 11/13c) are significantly expressed. This breakage of sequential Hox expression in echinoderm ancestors may be caused by loss of metameric gill slits, which are the most important synapomorphy of deuterostomes.

A box was designed to keep the acorn worm Saccoglossus mereschkowskii in laboratory conditions for 60 days and to monitor its behavior and feeding. Locomotion and construction of burrows in the sediment were found to be due to peristaltic movements of the proboscis, which periodically changes its shape from cylindrical to mushroom-like, and vice versa. Worms built U-shaped burrows connected with burrows of neighbor worms by flank anastomoses, thus producing a branched system of passages in a sediment layer up to 8 cm deep. The system is of importance for aeration of the upper sediment layer. When a worm is feeding, the proboscis sticks out from the anterior opening of the burrow and stretches along the surface of the sediment. Organic particles adhere to mucus secreted by the epidermal epithelium of the proboscis and are transported by ciliary beating to a furrow between the collar and proboscis, where the mouth is located.

Abstract Background The deep-sea acorn worm Quatuoralisia malakhovi belongs to the phylum Hemichordata, class Enteropneusta, family Torquaratoridae, which was described in 2005. Owing to their epibenthic lifestyle and deep-sea habitat features, torquaratorids differ anatomically from shallow-water acorn worms; however, their morphology and fine structure are poorly studied. We have the opportunity to make three complete detailed series of histological sections of Q. malakhovi and to study the microscopic anatomy, histology and fine structure of the reproductive system of this acorn worm using scanning and transmission electron microscopy. Results The sexes of Q. malakhovi are separate and indistinguishable externally. The lobed testes occupy the dorsal side of the genital wings and distinctly bulge into the peribranchial cavity by their mature lobes. The central part of the testis is always submerged into the genital wing and opens via a single gonad pore. The monociliary muscle cells stretch along the external wall of the testis and surround the gonad pore, probably taking part in the contraction of the testis lobes for spawning. The germinative epithelium of the testis contains spermatogenic cells at different stages of development and interstitial cells. Yolk cells are not found. Interstitial cells embrace the spermatogonia and spermatogenic columns, providing horizontal compartmentalization of the germinative epithelium, and contain numerous phagosomes with remnants of degenerating spermatogenic cells. The testis wall contains haemal lacunae, which are usually located on the side opposite the gonad pore. We describe the fine structure of spermatogonia, spermatocytes clustered in spermatogenic columns, spermatids, and spermatozoa. Spermatozoa are of the ectaquasperm type and consist of an acorn-shaped head and a flagellum 18–25 µm long. The sperm head includes a beak-shaped acrosomal part, a spherical nucleus and a midpiece containing a ring of 5 or rarely 6 mitochondria. Conclusions The male reproductive system and sperm structure of Q. malakhovi, a representative of the family Torquaratoridae, have a number of differences from shallow-water acorn worms; however, the spermatogenesis and sperm structure of Q. malakhovi generally follow the pattern of the other three enteropneust families, and the phylogenetic significance of these deviations should be the subject of further research.

Free endings of peripheral neurosecretory neurons (NNs) were found in the tegument of plerocercoids of five species of parasitic cestodes of fish in an ultrastructural study. The free terminals secreted vesicles on the tegument surface and into the host body. Secretion was experimentally shown to increase in response to the host fish blood serum. In the cestode body, NNs form paracrine-type contacts near the cell membranes of the frontal glands, the tegument, and muscles, functioning as endocrine glands. Simultaneously, NNs function as exocrine glands and secrete the so-called manipulative factors, which influence the physiology of the host.

Gutless marine worms of the family Siboglinidae have been found in the estuaries of the largest Arctic rivers Yenisei, Lena, and Mackenzie. Siboglinid metabolism is provided by symbiotic chemoautotrophic bacteria. Strong salinity stratification is characteristic of the estuaries of the largest Arctic rivers and ensures a high salinity at depths of 25–36 m, where siboglinids were found. High methane concentrations, which are necessary for siboglinid metabolism, result from dissociation of permafrost gas hydrates under the influence of river runoff in the conditions of Arctic warming.

Biodiversity in the Laptev Sea was assessed for gutless marine worms of the family Siboglinidae (Annelida), whose metabolism is provided by symbiotic bacteria that oxidize hydrogen sulfide and methane. Seven siboglinid species were found within the geographical boundaries of the Laptev Sea, and another species was found in an adjacent sector of the Arctic Basin. The largest number of finds and the greatest biological diversity of siboglinids were observed in the eastern part of the Laptev Sea in a field of numerous methane flares. One find was made in the estuary area of the Lena River at a depth of 25 m. A possible association of siboglinids with methane seepage areas is discussed.

Representatives of pogonophorans (Annelida, Siboglinidae), whose vital activity is provided by symbiotic chemoautotrophic bacteria that oxidize methane and hydrogen sulfide, were found in the St. Anna Trough at depths of 539 and 437 m. The finding of pogonophorans suggests high concentrations of methane, which might result from dissociation of bottom gas hydrates under the influence of the influx of warm Atlantic water into the Kara Sea along the St. Anna Trough.

The digestive system was anatomically studied in the deep-sea enteropneust Quatuoralisia mala-khovi . It was shown that lateral collar lips are twisted in such a way that they form a ciliary groove that leads to an internal channel, through which collected detritus particles are transferred to peripheral pharyngeal channels. The size of the selected particles ranges from 1–6 to 100–200 μm, which corresponds to feeding on the remains of planktonic diatoms. A fecal cord was observed to act as an anchor that holds the heavily watered jelly-like body of Torquaratoridae at the sea floor during feeding.

The discovery of new occurrences of pogonophores Siboglinum sp. and Nereilinum sp. from the St. Anna Trough (northwestern portion of the Kara Sea) is described in this paper. Previously, occurrences of pogonophores ( Crispabrachia yenisey and Galathealinum karaense ) were reported in the southern part of the Kara Sea, in the estuary of the Yenisei River. Two areas in the Kara Sea where pogonophores were found coincide with the regions of distribution of two types of gas hydrates: oceanic seafloor gas hydrates, and gas hydrates associated with permafrost. Gas hydrate deposits in the permafrost are confined to the coastal regions of the Kara Sea. A methane flux forms in areas of dissociation of gas hydrates under the influence of river runoff. This methane source is vital for the survival of pogonophores. The existence of pogonophores in the St. Anna Trough indicates the presence of a methane flux associated with the inflow of Atlantic water, which causes dissociation of seafloor gas hydrates. The possible role of Arctic warming is apparent in both processes.

The morphology of cerata and cnidosacs were studied in the nudibranch mollusk Pteraeolidia semperi (Bergh, 1870). Fine tubules arise from the gastrodermal channel of the digestive gland and contain cells with symbiotic algae (zooxanthellae). The cnidosac stores large kleptocnides. Thus, P. semperi provides a unique example of symbiotrophic feeding specialization. Morphological organization of its cerata and the digestive gland demonstrates several adaptations for housing zooxanthellae and providing them with proper conditions for active photosynthesis.

The anatomical and histological structure of the trophosome of the giant vestimentiferan Riftiapachyptila has been studied. The trophosome consists of longitudinally oriented cords. The cords of the trophosome intertwine, form diverticula and anastomose with each other. Each cord has an axial blood vessel inside, which is connected to afferent vessels on the surface of the cord by radial capillaries. Based on the data on the structure and development of the trophosome, it is suggested that the evolutionary precursor of the trophosome was a blood network connecting the ventral and dorsal vessels. The cells of the coelomic lining on the surface of the vessels grew and gave rise to the parenchymal tissue of the trophosome. At the same time, the trophosome developed from two sources, namely: due to the coelomic lining on the surface of the vessels of the intestinal plexus and due to the coelomic lining on the surface of the vessels of the circulatory plexus of the body wall.

The fine structure of the body wall and gut was for the first time studied in the competent larvae of the frenulate pogonophoran Siboglinum fiordicum. Mass apoptosis of cell nuclei was observed in the dermo-muscular body wall and coelomic epithelium. Apoptotic nuclei were found in both cell cytoplasm and outside of the larval body. In the latter case, each nucleus was surrounded by the plasmalemma, and the entire cluster was covered with the cuticle. Cells of the larval gut retained the usual structure with the cytoplasm filled with numerous yolky granules and the nucleus displaying usual morphology. Similar apoptotic processes have been described in vestimentiferans and found to be initiated by penetration of symbiotic bacteria through the integument into the dorsal mesentery. The process of apoptotic rearrangement of body wall cells and the formation of unique symbiosis with bacteria were assumed to be time-spaced in S. fiordicum, occurring sequentially rather than simultaneously, unlike in vestimentiferans.

In the Russian Arctic seas and adjacent areas of the Arctic basin, 120 sites of siboglinid records are currently known. Individuals belonging to 15 species have been collected. The largest number (49.2%) of records were made in the Barents Sea, followed by the Laptev Sea (37.5%) and the Arctic basin (10 records; 8.3%). No siboglinids have been reported from the Chukchi Sea. The largest number of species has been identified in both the Laptev Sea and Arctic basin (seven species each). Seventy-eight percent of the records were discovered at water depths down to 400 m. Many of the siboglinid records in the Arctic seas of Russia are associated with areas of high hydrocarbon concentrations. In the Barents Sea, Nereilinum murmanicum has been collected near the largest gas fields. The records of Oligobrachia haakonmosbiensis, N. murmanicum, Siboglinum ekmani, Siboglinum hyperboreum, Siboglinum norvegicum, as well as two undetermined species of siboglinids are associated with the marginal areas of bottom gas hydrates where methane emissions can occur. The Arctic seas of Russia feature vast areas of permafrost rocks containing gas hydrates flooded by the sea. Under the influence of river runoff, gas hydrates dissociate, and methane emissions occur. Crispabrachia yenisey and Galathealinum karaense were found in the Yenisei estuary, and O. haakonmosbiensis was found in the Lena estuary.

The article describes the parameters of maintaining the gutless symbiotrophic annelid Siboglinum fiordicum in laboratory conditions outside the marine environment for 64 days.

The common ancestor of Ambulacraria was a mobile bilaterally symmetrical organism. Its body was subdivided into a preoral (proboscis) region, a perioral (collar) region with ciliated tentacles, and a trunk region with a metameric coelom and metameric gill slits. Like in other Bilateria, the preoral and tentacular regions of Ambulacraria are free of Hox gene expression, which begins in the area of the first pair of gill slits in compliance with the colinearity. The axial complex is a synapomorphy of the Ambulacraria clade. A bilaterally symmetrical ancestor of Echinodermata lay on its dorsal side, so its anus was shifted to the ventral side as in present-day echinoderm larvae. This stage of evolution corresponds to the Early Paleozoic bilaterally symmetrical forms, such as Protocinctus, Ctenocystis , and Ctenoimbricata . The common ancestor of echinoderms had a symmetrical tentacular apparatus consisting of five ciliated tentacles on each side of a collar. At the next stage, the ancestors of echinoderms lay on the right side that resulted in the reduction of the tentacles on the right side and the right hydrocoel. This evolution stage includes various Early Paleozoic forms ( Cothurnocystis , Dendrocystoides , Syringocrinus , Castericystis , Coleicarpus , Rhenocystis , etc.). The next stage is related to the sedentary lifestyle. During this stage, the mouth and tentacles occupied an apical position, the anus has shifted up, and thus an intestinal loop was formed. The five primary tentacles of the left side of the ancestor of Ambulacraria predestined the formation of pentaradial symmetry of echinoderms. The primary tentacles remained only in Holothuroidea. The secondary ciliary grooves were formed between the primary tentacles. These grooves were accompanied by hydrocoelic canals, which gave rise to the water-vascular (ambulacral) system. The present-day echinoderms are characterized by several metameric rings formed by the derivatives of the left somatocoel. They derive from the coelomic segments of the left side of the trunk of the ambulacrarian ancestor and, thus, the echinoderms retain coelomic metamerism. The attachment of the crinoid larvae by the preoral lobe reflects the ancient method of locomotion of deuterostomes using the proboscis, but, in reality, the Pelmatozoa stalk is homologous to the Pterobranchia stalk, i.e., the posterior end of the body, because the right somatocoel grows into it. During metamorphosis, the internal coelomic complex of the larva is inverted by 180°. The significance of this inversion for the translocation of the anterior genes of the Hox cluster is discussed. The ancestors of Eleutherozoa began to crawl on the oral surface, which led to a shift of the anus to the aboral side. Thus, the ontogeny and phylogeny of echinoderms exhibits a change from bilateral symmetry to dissymmetry, and then the development of pentameric symmetry.

The article presents a report on the findings of representatives of frenulate pogonophorans Nereilinum murmanicum in the northern and central parts of the Barents Sea, which significantly expands the range of this species and provides guidance on its distribution in this basin. Here we present the coordinates of new finds with an indication of the depth. Find points were associated with data on known and potential hydrocarbon deposits.