Evolution of invertebrate nervous systems: the Chaetognatha as a case study

Harzsch, S. and Wanninger, A. 2010. Evolution of invertebrate nervous systems: the Chaetognatha as a case study. —Acta Zoologica (Stockholm) 91 : 35–43 Although recent molecular studies indicate that Chaetognatha may be one of the earliest Bilaterian offshoots, the phylogenetic position of this taxon still is a matter of ongoing debate. In this contribution, we review recent attempts to contribute phylogenetic information on the Chaetognatha by analysing structure and development of their nervous system (neurophylogeny). Analysing this group of organisms also has a major impact on our understanding of nervous system evolution in Bilateria. We review recent evidence from this field and suggest that Urbilateria already was equipped with the genetic toolkit required to build a complex, concentrated central nervous system (CNS), although this was not expressed phenotypically so that Urbilateria was equipped with a nerve plexus and not a CNS. This implies that in the deep metazoan nodes, concentration of the ancestral plexus occurred twice independently, namely once after the protostome–deuterostome split on the branch leading to the protostomes (resulting in a ventrally positioned nerve cord) and once along the chordate line (with a dorsal nerve cord).     

Distribution and development of the serotonin-and RFamide-like immunoreactive neurons in the arrowworm,Paraspadella gotoi (Chaetognatha)

Summary The distribution and development of serotonin-and RFamide-like immunoreactivities in the nervous system of Chaetognatha, Paraspadella gotoi, were examined in whole-mount preparations. In adults, a single serotonin-like immunoreactive (5HTLI) neuron and numerous RFamide-like immunoreactive (RFaLI) neurons were found in the central nervous system. Based on the structure of the fins, hooks, and eyes, seven postembryonic developmental stages were recognized. The most obvious features of the stages are: stage 1, newly hatched young; stage 2, elongation of a continuous lateral tail fin; stage 3, separation of the lateral and tail fins; stage 4, appearance of hooks; stage 5, pigmentation of eyes, stage 6, attachment by tail adhesive fins; stage 7, prey capture. Stage 1 did not show any immunoreactivity. The 5HTLI neuron first appeared at stage 4 and its axonal pathway became similar to the adult at stage 6. On the other hand, the RFaLI neurons appeared at stage 3 in the ventral ganglion. Some of their somata disappeared at stage 5 and the neuronal architecture resembled the adult at stage 7 although the RFaLI neurons in the cerebral ganglion were complete at the juvenile stage.We are sad to announce that Dr. M. Yoshida died on 29 October 1988     

The nervous system of the tube-worm Chaetopterus variopedatus (Polychaeta)

Silver impregnation of serial histological sections of the tubeworm Chaetopterus variopedatus revealed the presence of a subepidermal nervous system. The anterior nervous system is delimited by the first 11 segments and comprises (1) two dorsolateral cerebral ganglia and lateral instead of ventral nerve cords which are widely separated and thus connected by unusually long commissures, (2) a pharyngeal ganglion in the fourth segment which is connected to the cerebral ganglia by pharyngeal nerves and constitutes along with the pharyngeal plexus a stomatogastric or enteric nervous system, and (3) small, presumably segmental ganglionic swellings along the lateral nerve cords from which emerge commissures and parapodial nerves. No subesophageal ganglion or periesophageal connective could be identified. The lateral nerve cords converge toward the midline in the 12th segment to form the posterior nervous system comprising a pair of ventromedian nerve cords with their repetitive segmental ganglia from which emerge numerous short commissures and three segmental nerves coursing toward the dorsal and ventral regions of parapods and toward the neuropod. Light and electron microscopic investigations of cerebral and segmental ganglia showed an arrangement of inner neuropile and of unipolar neuron somata at the periphery. The neuropile comprises numerous neurites ranging in diameter from 0.5 to 10 μm and making polarized or symmetrical synaptic junctions with each other. The pharyngeal ganglion consists of a similar neuropile and of a large mass of cell bodies which is traversed by an elaborate network of sinuses and harbors three types of neurosecretory cells in addition to the conventional neuron somata. These findings are interpreted in the framework of the highly specialized morphological features and habits of Chaetopterus, and the welldeveloped stomatogastric system is considered to be related to control of the feeding activities.     

Comparative immunohistochemistry of the cerebral ganglion in Gastrotricha: an analysis of FMRFamide-like immunoreactivity in Neodasys cirritus (Chaetonotida), Xenodasys riedli and Turbanella cf. hyalina (Macrodasyida)

The neuropeptide FMRFamide (Phe–Met–Arg–Phe–NH₂) is part of a large and diverse family of peptidergic neurotransmitters present throughout the animal kingdom. To date, no such neuropeptides have been demonstrated in gastrotrichs despite their presence in closely related invertebrates such as nematodes. Here, the FMRFamidergic nervous system of three marine gastrotrichs is investigated with immunofluorescence, CLSM, and 3D computer imaging to gain insight into structure of the cerebral ganglion and test various phylogenetic hypotheses on its organization. Results reveal that FMRFamide-like immunoreactivity (IR) is present throughout the nervous systems of three species: Neodasys cirritus (Chaetonotida), Xenodasys riedli and Turbanella cf. hyalina (Macrodasyida). Both macrodasyidans possess FMRFamide-like IR in the central, peripheral- and stomatogastric-nervous systems, while FMRFamide-like IR is restricted to the CNS in N. cirritus. In all three species, the cerebral ganglion is dumbbell-shaped and bordered bilaterally by cerebral perikarya: numerous perikarya are present in X. riedli and N. cirritus, while few perikarya are present in T. cf. hyalina. Cerebral perikarya flank the nerve ring neuropil, which contains IR fibers in the supra- and subpharyngeal commissures of both macrodasyidans, but in N. cirritus, only contains IR fibers in the suprapharyngeal commissure. Together, these results confirm the peripharyngeal nature of the gastrotrich cerebral ganglion, but are equivocal on hypotheses of its tripartite structure. Still, the neural organization of gastrotrichs, in particular, the architecture of the cerebral ganglion, is expected to hold valuable information for future assessments of gastrotrich phylogeny, and may yet provide key insights into the evolution of this enigmatic taxon.     

On the fine structure of the cerebral ganglion of Sagitta (Chaetognatha)

Summary In this study the fine structure of the cerebral ganglion of Sagitta setosa (Chaetognatha) is investigated. The ganglion is flat and superficially positioned dorsally, below the basal lamina of the cephalic epidermis. It is surrounded by a specifically differentiated sheath. This sheath is made up of cells, which are interpreted as representing glial cells, and can be divided into an outer and an inner zone. The outer zone is composed of flat sheath cells with pale nuclei and few organelles. The inner zone consists of densely packed, extremely thin lamellar cellular processes. These attenuated lamellae, which still contain cytoplasm, resemble the myelin sheath of vertebrate axons. The intercellular space between the lamellae contains electron-dense material. In the sheath specialized intercellular contacts occur. The inner zone of this sheath extends at definite points into the centre of the ganglion and separates a zone of perikarya from the neuropil, as well as the single perikarya from each other. The perikarya are relatively uniform and do not form a cortex, but are concentrated mainly in lateral parts of the cerebral ganglion. Within the neuropil are axonal endings which have synaptic contacts with several postsynaptic elements. These anatomical findings are discussed with respect to their functional significance.     

On the neurosecretory system of Dendrobaena atheca Cernosvitov

Four kinds of neurosecretory cells A, B, U and C are distinguished in the central nervous system of Dendrobaena atheca Cernosvitov. A cells, which show different morphological characteristics under different physiological states and during their cyclic changes, are the most active neurosecretory cells. They form the outer layer of the cortical cell zone in the cerebral ganglion. B cells are large and medium sized and are distributed in all parts of the central nervous system. U cells are found only in the sub-pharyngeal ganglion while C cells are distributed in the sub-pharyngeal as well as in the ventral nerve cord ganglion. The number and secretory activity of C cells decrease in caudal direction. Further, Gomori-positive cells are also observed in the ganglia of the vegetative nervous system. A rudimentary neurohaemal organ, the storage zone, has been observed in the cerebral ganglion and there appears to be another neurohaemal area in the ventral nerve cord ganglion. The storage zone is formed by the terminal ends of the axons of A cells. The chrome alum haematoxylin phloxin (CHP) and aldehyde fuchsin (AF) positive substances in the form of granules are found in this area. The cerebral ganglion is richly supplied by blood capillaries. The distal end of the axons of B cells are swollen like a bulb while in some cases the axons are united to form an axonal tract. Extra-cellular material is abundant in different parts of the nervous system. In all cell types, the perinuclear zone is the first to show activity in the secretory cycle. It appears that the nucleus may be involved in the elaboration of the neurosecretory material in the cells.     

On the serotonergic nervous system of two planktonic rotifers, Conochilus coenobasis and C. dossuarius (Monogononta, Flosculariacea, Conochilidae)

The serotonergic nervous systems of two non-colonial species of Conochilus were examined to obtain the first immunohistochemical insights into the neuroanatomy of species of Flosculariacea (Rotifera, Monogononta). Species of Conochilus, subgenus Conochiloides, were examined using serotonin (5-HT) immunohistochemistry, epifluorescence and confocal laser scanning microscopy, and 3D computer imaging software. In specimens of C. coenobasis and C. dossuarius, the serotonergic nervous system is defined by a dorsal cerebral ganglion, apically directed cerebral neurites, and paired nerve cords. The cerebral ganglion contains approximately four pairs of small 5-HT-immunoreactive perikarya; one pair innervates the posterior nerve cords and three pairs innervate the apical field. The most dorsal pair innervates a coronal nerve ring that encircles the apical field. Within the apical field is a second nerve ring that outlines the inner border of the coronal cilia. Together, both the inner and outer nerve rings may function to modulate ciliary activity of the corona. The other two pairs of perikarya innervate a region around the mouth. Specific differences in the distribution of serotonergic neurons between species of Conochilus and previously examined ploimate rotifers include the following: (a) a lack of immunoreactivity in the mastax; (b) a greater number of apically directed serotonergic neurites; and (c) a complete innervation of the corona in both species of Conochilus. These differences in nervous system immunohistochemistry are discussed in reference to the phylogeny of the Monogononta.     

Hydra size and budding rate: influence of feeding

Summary

In the present study of Dugesia tigrina, the development of the nervous system is followed and compared during regeneration after fission and after decapitation. Immunocytochemistry was used, with antisera raised against the biogenic mine, 5-hydroxyhyptamine (5-HT) and the two neuropeptides, neuropeptide F (NPF), and FMRFamide. The results indicate that two processes are involved in the formation of the new cerebral ganglion. First, new processes sprouting from the original main longitudinal nerve cords bend transversely, indicating the position of the developing horseshoe-shaped anterior cerebral commissure. Then new nerve cells in front of the commissure differentiate from neoblasts and their growth cones fasciculate with the fibres from the old main longitudinal nerve cords. In the cerebral ganglion, 5-HT-IR cells appear before NPF-IR cells, in contrast to the pharynx where NPF-IR cells differentiate before the 5-HT-IR cells. In the peripheral nervous system, NPF-IR fibres and cells appear at a very early stage and dominate the whole regeneration process. A role for the PNS in early pattern formation is suggested.     

Serotonin-like immunoreactivity in the central nervous system and gonad of the scallop,Patinopecten yessoensis

Summary The localization of neurons containing serotonin in the central nervous system and the gonad of the scallop, Patinopecten yessoensis, was examined immunohistochemically. In the central nervous system a large number of immunoreactive perikarya were observed in the following regions: a part of the anterior lobe of the cerebral ganglion; the posterior lobe of the cerebral ganglion; the pedal ganglion; and the accessory ganglion. No immunoreactive perikarya were found in the visceral ganglion. Numerous immunoreactive fibers were revealed in the neuropil of all central ganglia. In the gonadal region immunoreactive fibers were distributed around the gonoduct and along the germinal epithelium.This work was supported by a grant from the Ministry of Education, Science and Culture, Japan     

Distribution and ultrastructure of neurosecretory cells in the cerebral ganglion of the earthworm

Neurosecretory (Nsy) cells within the cerebral ganglion of Lumbricus terrestris were classified ultrastructurally. The Nsy cells within the subesophageal ganglion, nerve cord ganglion, and the peripheral nervous system were also examined. A comparative survey of Nsy cells of four other species of oligochaetes, Eisenia feotida, octolasion cyaneum, Dendrobeona subrubicunda, and Allolophora longa, was also carried out. Seven cell types (A1, A2, A3, A4, A5, C, and SEF), distinguished by special cytological and ultrastructural features, were found within the cerebral ganglion. Distribution of these cells inside and outside the cerebral ganglion was studied in detail by light and electron microscopy. The nerve terminals of each cell type were followed into the neuropile region. Exocytosis from cell bodies appears to be the main release mechanism for the Nsy granules, whereas small Nsy vesicles are released through synapses in the neuropile. Peripheral fibers of some cell types (A1, A2, and A3) extend through the capsule to the pericapsular epithelium. It is possible that Nsy cells secrete hormones from their cell bodies and peripheral processes and that their centrally directed axons release modulators/transmitters within the neuropile.     

Distribution of GABA-like immunoreactive neurons in the slug Limax maximus

Summary Immunohistochemical techniques were used to study the distribution of gamma-amino butyric acid (GABA)-like immunoreactive neurons in the nervous system of the slug Limax maximus. Approximately 170 GABA-like immunoreactive cell bodies were found in the central nervous system. These were located in the cerebral, buccal and pedal ganglia. Most GABA-like immunoreactive neurons had small cell bodies, which were aggregated into discrete clusters within the cerebral and pedal ganglia. Three pairs of longer, uniquely identifiable, GABA-like immunoreactive cells were found in the cerebral ganglion. GABA-like immunoreactive nerve fibres were also found in all of the central ganglia but were absent from peripheral nerves. These results suggest that GABA acts as a central neurotransmitter in the slug. The possible roles of GABA-ergic neurotransmission in the slug are discussed.     

Distribution of serotonin-like immunoreactive neurons in the slug<Emphasis Type="Italic"> Limax valentianus</Emphasis>

Immunohistochemical techniques were used to study the distribution of serotonin-containing neurons in the nervous system of the slug Limax valentianus. Approximately 350 serotonin-like immunoreactive cell bodies were found in the central nervous system. These were located in the cerebral, pedal, visceral and right parietal ganglia. Most serotonin-like immunoreactive neurons had small cell bodies, which were aggregated into discrete clusters. A pair of previously identified metacerebral giant cells were found on the anterior side of the cerebral ganglion, and two additional pairs of uniquely identifiable, serotonin-like immunoreactive cells were found on the posterior side of the cerebral ganglion. The whole-mount maps of these stained neurons will be useful in further physiological and biochemical studies of olfactory learning at the cellular level in Limax valentianus.This study was supported by Grants-in-Aid for Scientific Research from the Ministry of Education, Culture, Science, Sports and Technology, Japan (nos. 12307053 and 13771353)     

Serotonin immunoreactivity in the nervous system of the Pandora larva, the Prometheus larva, and the dwarf male of Symbion americanus (Cycliophora)

Cycliophora is a recently described phylum of enigmatic metazoans with a very complex life cycle that includes several sexual and asexual stages. Symbion pandora and Symbion americanus are the only two cycliophoran species hitherto described, of which morphological and genetic knowledge is still deficient to clarify the phylogenetic position of the phylum. Aiming to increase the database on the cycliophoran neural architecture, we investigated serotonin immunoreactivity in the free swimming Pandora larva, the Prometheus larva, and the adult dwarf male of S. americanus. In the larval forms, serotonin is mainly expressed in a ring-shaped pattern at the periphery of the antero-dorsal cerebral ganglion. Additionally, several serotonergic perikarya emerge from both sides of the cerebral ganglion. Thin neurites project anteriorly from the cerebral ganglion, while a pair of ventral longitudinal neurites emerges laterally and runs along the anterior-posterior body axis. Posteriorly, the ventral neurites fuse and extend as a posterior projection. In the dwarf male, serotonin is found mainly in the commissural neuropil of the large anterior cerebral ganglion. In addition, serotonin immunoreactivity is present in the most anterior region of the ventral neurites. Comparative analysis of spiralian nervous systems demonstrates that the neuroanatomy of the cycliophoran larval stages resembles much more the situation of adult rather than larval spiralians, which may be explained by secondary loss of larval structures and heterochronic shift of adult components into the nervous system of the Pandora and the Prometheus larva, respectively.     

Comparative studies on cholinergic activities in the central nervous system ofLocusta migratoria

Summary Comparative studies on biochemical and pharmacological aspects of cholinergic transmission in the cerebral ganglion of locust and the mouse cerebral cortex were performed. The results point to common features and significant differences in the cholinergic pharmacology of insect and vertebrate nervous system: ACh content as well as AChE and ChAT activities reflect a highly cholinergic innervation of the locust cerebral ganglion. While the nicotinic ACh-receptor type predominates in locust nervous system, the muscarinic type is the main receptor type in mouse brain.The kinetic data for the high affinity choline uptake of locust synaptosome preparations corroborate the cholinergic properties of nerve endings from locust head ganglion.Abbreviations ACh acetylcholine - ChAT cholineacetyltransferase - AChE acetylcholinesterase - QNB quinuclidihylbenzilate - -BTX -bungarotoxin The author is indebted to Prof. Lueken for support and encouragement during the course of this work. I also gratefully acknowledge the excellent technical assistance of Mrs. M. Düwel. This work was financially supported by the Deutsche Forschungsgemeinschaft (Br 712/2-2).     

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.

     

A SURVEY OF NEUROENDOCRINE PHENOMENA IN NON-ARTHROPOD INVERTEBRATES

• 1 Neurosecretory phenomena are apparently ubiquitous among Metazoa.
• 2 In Hydra, neurosecretory products are probably involved in the control of growth and reproduction.
• 3 Secretory elements in the central nervous system of turbellarians probably promote fission, regeneration and reproduction.
• 4 The cerebral ganglia of nemertines are the source of a hormone which exercises an inhibitory influence on maturation of the gonads and the development of somatic sexual characteristics. A principle secreted by the ganglia and/or the associated cerebral organs controls weight regulation. A sex hormone controls sexual differentiation.
• 5 Secretory neurons may influence the production of exsheathing fluid and ecdysis in nematodes.
• 6 In nereid polychaetes, a single hormone which originates from the cerebral neuroendocrine system apparently promotes segment proliferation and inhibits maturation. However, in certain other polychaetes, contrasting endocrine mechanisms seem to operate.
• 7 In lumbricid oligochaetes, a hormone secreted by the cerebral ganglion (and possibly by other nervous centres also) promotes gonadal activity and the differentiation of somatic sexual characters. In some species the ganglion exerts an inhibitory influence on the associated processes of ‘diapause’ and posterior regeneration. The cerebral and suboesophageal ganglia are implicated in the control of osmoregulation. Cerebral neurosecretory cells in limicolous oligochaetes may influence regenerative growth and osmoregulation.
• 8 The cerebral ganglia of leeches secrete a gonadotrophic hormone.
• 9 In gastropod molluscs, hormones secreted by the cerebral ganglia, glandular ‘dorsal bodies’ and/or optic tentacles are responsible for the control of the development of the reproductive tract, and the activity of the gonads. Contrasting mechanisms are thought to operate in the different gastropod groups and there is conflicting evidence particularly with respect to the endocrine functions of the optic tentacles and the gonads. The ‘bag cells’ of the abdominal ganglion of ApZysiu secrete a hormone which induces egg-laying. Cells in the pleural and parietal ganglia probably control osmoregulation in Lymnaea.
• 10 The optic glands of cephalopods secrete a gonadotrophic hormone, but sex hormones are apparently absent. The functional significance of the neurovenous tissues (presumptive neurosecretory complexes of unusual character) is obscure in most cases.
• 11 The radial nerves of starfish are the source of a hormone which induces the production of I-methyl adenine by the follicle cells of the gonad. This second principle stimulates oocyte maturation and the shedding of male and female gametes.
• 12 The relevance of such information to the development of certain biological concepts and to various aspects of comparative physiology is briefly discussed.

     

Musculature and nervous system of<Emphasis Type="Italic"> Gnathostomula peregrina</Emphasis> (Gnathostomulida) shown by phalloidin labeling,immunohistochemistry, and cLSM,and their phylogenetic significance

Musculature and nervous system of Gnathostomula peregrina (Gnathostomulida, Scleroperalia) were reconstructed from whole animals by immunohistochemistry and confocal laser scanning microscopy. The F-actin muscular subset, stained with FITC-labeled phalloidin, consists of: (1) eleven pairs (four ventral, one ventrolateral, one dorsolateral, five dorsal) of longitudinal muscles; (2) two types of diagonal muscles (thin fibers throughout the body, and slightly thicker fibers of which seven pairs occur ventrally and two pairs dorsally); (3) evenly spaced thin circular fibers that gird the posterior half of the body, continuing less prominently into the anterior half; and (4) a complex pharyngeal and genital musculature. Dorsoventral muscles are absent. The organization of the FMRFamidergic nervous system shows: (1) a central nervous system with a frontal ganglion and one pair of longitudinal nerves ending in a terminal commissure, and one median ventral nerve; (2) eight to ten unipolar perikarya above, and up to ten bipolar perikarya in front of the brain; (3) a total of five (one unpaired, two paired) longitudinal nerves of the peripheral nervous system with two to four accompanying perikarya; and (4) a buccal ganglion of the stomatogastric nervous system with six to eight perikarya above the pharyngeal bulbus. Our results reveal the musculature and nervous system of Gnathostomula to be more complex than hitherto reported.     

Patterns of serotonin-immunoreactive neurons in the central nervous system of the earthworm Lumbricus terrestris L.

Summary The distribution patterns of serotonin-immunoreactive somata in the cerebral and subpharyngeal ganglion, and in the head and tail ganglia of the nerve cord of Lumbricus terrestris are described from whole-mount preparations. A small number of serotonin-immunoreactive neurons occurs in the cerebral ganglion, in contrast to the large population of serotonin-immunoreactive neurons that exists in all parts of the ventral nerve cord. From the arrangement of serotonin-immunoreactive somata in the subpharyngeal ganglion, we suggest that this ganglion arises from the fusion of two primordial ganglia. In head and tail ganglia, the distribution of serotonin-immunoreactive somata resembles that in midbody segments. Segmental variations in the pattern and number of serotonin-immunoreactive somata in the different body regions are discussed on the background of known developmental mechanisms that result in metameric neuronal populations in annelids and arthropods.Abbreviations CG1, CG2 cerebral soma group 1, 2 - CNS central nervous system - GINs giant interneurons - 5-HT 5-hydroxytryptamine, serotonin - 5-HTi 5-HT-immunoreactive - N side nerve - SG19 subpharyngeal soma group 1–9 - SN segmental nerve     

A morphological study of the anterior nervous system of the praesoma of Neoechinorhynchus cylindratus (Acanthocephala: Neoechinorhynchidae)

The nerve pathways in the praesoma, based on light microscopy of serial transverse, sagittal, and longitudinal sections stained with Ehrlich's acid hematoxylin are described for the first time for a memeber of Neoechinorhynchus. The route from the cerebral ganglion to the musculature and sense organs of the proboscis and body wall for 11 nerves, five pair and one single, the presence and structure of the Stutzzelle (support cell) and its association with the neck sense organs are described. A comparison with the nervous system in the praesoma of Paulisentis fractus is discussed.