Epidermal receptor development and sensory pathways in vitally stained amphioxus (Branchiostoma floridae)

Chloromethyl (CM) DiI was applied to the exterior of living embryos, larvae, and metamorphic juveniles of amphioxus. This fluorescent dye is taken up preferentially (but not highly selectively) by epidermal receptors and often stains sensory axons to their full extent. Type I primary receptors in the epidermis first become morphologically detectable along the rostrocaudal axis of the 2.5 day larva when their epidermal perikarya extend unbranched axons to the nerve cord. These axons run posteriorly or anteriorly within the nerve cord, depending on whether their perikarya are located, respectively, rostral or caudal to the most posterior pharyngeal slit. In later larvae, axons of type I receptors are organized into a dorsal and a subdorsal sensory tract on either side of the nerve cord. In the epidermis of metamorphic juveniles, CM-DiI also stains type II receptors (which are axonless, secondary receptors) and ventral pit cells (which may not be receptors). It is probable, but not yet conclusively demonstrated, that peripheral neurites from Retzius bipolar cells (primary intramedullary sensory neurones) synapse with type II secondary epidermal receptors or ramify freely among the other epidermal cells. The discussion considers homologies among epidermal sensory receptors in chordates.     

The motor nuclei of the glossopharyngeal-vagal and the accessorius nerves in the rat

Retrograde cobalt labeling was performed by incubating the rootlets of cranial nerves IX, X and XI, or the central stumps of the same nerves, in a cobaltic lysine complex solution, and the distribution of efferent neurons sending their axons into these nerves was investigated in serial sections of the medulla and the cervical spinal cord in young rats. The following neuron groups were identified. The inferior salivatory nucleus lies in the dorsal part of the tegmentum at the rostral part of facial nucleus. It consists of a group of medium-sized and a group of small neurons. Their axons make a hair-pin loop at the midline and join the glossopharyngeal nerve. The dorsal motor nucleus of the vagus situates in the dorsomedial part of the tegmentum. Its rostral tip coincides with the first appearance of sensory fibres of the glossopharyngeal nerve, the caudal end extends into the pyramidal decussation. The constituting cells have globular or fusiform perikarya and they are the smallest known efferent neurons. The ambiguous nucleus is in the ventrolateral part of the tegmentum. The rostral tip lies dorsal to the facial nucleus, and the caudal tip extends to the level of the pyramidal decussation. The rostral one third of the ambiguous nucleus is composed of tightly-packed medium sized neurons, while larger neurons are arranged more diffusely in the caudal two thirds. The long dendrites are predominantly oriented in the dorsoventral direction. The dorsally-oriented axons take a ventral bend anywhere between the ambiguous nucleus and dorsal motor nucleus of the vagus. The motoneurons of the accessorius nerve are arranged in a medial, a lateral and a weak ventral cell column. The medial column begins at the caudal aspect of the pyramidal decussation and terminates in C2 spinal cord segment. The lateral and ventral columns begin in C2 segment and extend into C6 segment. The neurons have large polygonal perikarya and characteristic cross-shaped dendritic arborizations. The axons follow a dorsally-arched pathway between the ventral and dorsal horns. The accessorius motoneurons have no positional relation to any of the vagal efferent neurons. It is concluded that the topography and neuronal morphology of accessorius motoneurons do not warrant the designation of a bulbar accessorius nucleus and a bulbar accessorius nerve.     

Distribution and localization of immunoreactive FMRFamide-like peptides in the lancelet.

Immunofluorescence was used to study the distribution of FMRFamide (Phe-Met-Arg-Phe-NH2) in premetamorphic larvae and adults of the lancelet, Branchiostoma lanceolatum. In the larvae, FMR-Famide-containing presumably neuronal perikarya and fibers were limited to the anterior third of the dorsal nerve cord. Throughout this region, most of the immunoreactive perikarya and fibers were located ventrolaterally and ventrally within the nerve cord; in addition, in the caudal part of the cerebral vesicle, some of the immunofluorescent cells projected cytoplasmic extensions across the slot-like neural canal. In adult lancelets, immunofluorescence was detected in cells of the Hatschek's pit (a probable homologue of the anterior hypophysis of vertebrates); however, no immunofluorescence was detected in the larval preoral pit, which is the ontogenetic precursor of Hatschek's pit. Moreover, the FMR-Famide-containing elements do not show immunoreactivity to other peptides of the FaRPs family such as pancreatic polypeptide (PP). The results suggest that FMRF-amide may be involved in neuroendocrine functions of lancelets.     

Cell death of asynaptic neurons in regenerating spinal cord

The weakly electric fish Sternarchus albifrons possesses a unique class of asynaptic neurons, the electromotor neurons, whose axons constitute the electric organ. The cell bodies of origin of the electrocyte axons are located in the spinal cord. Both spinal cord and electromotor neurons ( electrocytes ) regenerate after amputation of the tail. Sternarchus spinal cords which have regenerated for 1 or more years show a progression in number of perikarya of electromotor neurons along the rostro-caudal axis. The most recently regenerated region of the cord is at the caudal end, which consists of a tube of ependyma. Progressing rostrally along regenerated spinal cord from the caudal end, numerous cells are generated and large numbers of electromotor neurons differentiate. The maximum number of electromotor neurons per transverse section of regenerated cord is five times higher than in normal mature cord. Rostral to this, the number of electromotor neurons decreases gradually to the normal number near the transition zone (the border with unregenerated cord). As the more rostral regenerated cord has presumably had a longer period of regeneration, we conclude that excess numbers of electromotor neurons are generated initially, and that subsequently the number of these neurons is decreased by cell death. This conclusion is supported by the fact that younger regenerates (2-4 months) have larger-than-normal numbers of perikarya of electromotor neurons extending up to the transition zone (Anderson and Waxman , 1981). No evidence of migration or depletion of electromotor neurons from unregenerated cord rostral to the amputation has been observed. Since the axons of the electromotor neurons in Sternarchus do not normally form any synapses, this study provides evidence that factors other than synaptic competition must be responsible for determining cell death during regeneration of these spinal neurons.     

Projections of neurochemically specified neurons in the porcine colon

The intramural projections of nerve cells containing serotonin (5-HT), calcitonin gene-related peptide (CGRP), vasoactive intestinal peptide (VIP) and nitric oxide synthase or reduced nicotinamide adenine dinucleotide phosphate diaphorase (NOS/NADPHd) were studied in the ascending colon of 5- to 6-week-old pigs by means of immunocytochemistry and histochemistry in combination with myectomy experiments. In control tissue of untreated animals, positive nerve cells and fibres were common in the myenteric and outer submucous plexus and, except for 5-HT-positive perikarya, immunoreactive cell bodies and fibres were also observed in the inner submucous plexus. VIP- and NOS/NADPHd-positive nerve fibres occurred in the ciruclar muscle layer while VIP was also abundant in nerve fibres of the mucosal layer. 5-HT- and CGRP-positive nerve fibres were virtually absent from the aganglionic nerve networks. In the submucosal layer, numerous paravascular CGRP-immunoreactive (IR) nerve fibres were encountered. Myectomy studies revealed that 5-HT-, CGRP-, VIP- and NOS/NADPHd-positive myenteric neurons all displayed anal projections within the myenteric plexus. In addition, some of the serotonergic myenteric neurons projected anally to the outer submucous plexus, whereas a great number of the VIP-ergic and nitrergic myenteric neurons send their axons towards the circular muscle layer. The possible function of these nerve cells in descending nerve pathways in the porcine colon is discussed in relation to the distribution pattern of their perikarya and processes and some of their morphological characteristics.     

Efferent neurons of the lateral-line system and the VIII cranial nerve in the brainstem of anurans

Summary The octavo-lateral efferent system of several anuran species was studied by means of retrograde transport of horseradish peroxidase. This system is organized similarly in all larval anurans and in all adult aglossids. All have two groups of efferent neurons in the nucleus reticularis medialis between the VIIIth and the IXth motor nucleus. The caudal group consists of efferent neurons that supply the posterior lateral-line nerve (NLLp) and a considerably smaller group of neurons supplying both the NLLp and the anterior lateral-line nerve (NLLa). The rostral group is composed of efferent neurons supplying the NLLa, neurons projecting to the inner ear and neurons supplying both the inner ear and the NLLa. Efferent neurons of the VIIIth cranial nerve exhibit a rostrocaudal cytoarchitectonic differentiation. Caudal perikarya, which are rounder in shape than those of the rostral part, have a dendritic projection to the superior olive. It is suggested that this differentiation reflects a functional differentiation of acoustic and vestibular efferent neurons.Labeled neurons were ipsilateral to the site of application of HRP. None were found in the vestibular nuclei or in the cerebellum.Efferent axons projecting to neuromasts of the NLLa leave the medulla with the VIIth nerve, axons projecting to neuromasts of the NLLp exit via the IXth nerve. Cell counts and the observation of axonal branching revealed that efferent units of both the lateral-line and the VIIIth-nerve system supply more than one receptor organ. In contrast to the lateral-line system, dendrites of efferent neurons of the VIIIth nerve project dorsally onto its nuclei, and afferents of the VIIIth nerve project onto efferent neurons. These structures most probably represent a feedback loop between the afferent and efferent systems of the VIIIth cranial nerve.     

Bridging Schwann cell transplants promote axonal regeneration from both the rostral and caudal stumps of transected adult rat spinal cord

Transplantation of cellular components of the permissive peripheral nerve environment in some types of spinal cord injury holds great promise to support regrowth of axons through the site of injury. In the present study, Schwann cell grafts were positioned between transected stumps of adult rat thoracic spinal cord to test their efficacy to serve as bridges for axonal regeneration. Schwann cells were purified in culture from adult rat sciatic nerve, suspended in Matrigel:DMEM (30:70), and drawn into polymeric guidance channels 8mm long at a density of 120×106 cells ml-1. Adult Fischer rat spinal cords were transected at the T8 cord level and the next caudal segment was removed. Each cut stump was inserted 1mm into the channel. One month later, a bridge between the severed stumps had been formed, as determined by the gross and histological appearance and the ingrowth of propriospinal axons from both stumps. Propriospinal neurons (mean, 1064±145 SEM) situated as far away as levels C3 and S4 were labelled by retrograde tracing with Fast Blue injected into the bridge. Near the bridge midpoint there was a mean of 1990±594 myelinated axons and eight times as many nonmyelinated, ensheathed axons. Essentially no myelinated or unmyelinated axons were observed in control Matrigel-only grafts. Brainstem neurons were not retrogradely labelled from the graft, consistent with growth of immunoreactive serotonergic and noradrenergic axons only a short distance into the rostral end of the graft, not far enough to reach the tracer placed at the graft midpoint. Anterograde tracing with PHA-L introduced rostral to the graft demonstrated that axons extended the length of the graft but essentially did not leave the graft. This study demonstrates that Schwann cell grafts serve as bridges that support (1) regrowth of both ascending and descending axons across a gap in the adult rat spinal cord and (2) limited regrowth of serotonergic and noradrenergic fibres from the rostral stump. Regrowth of monoaminergic fibres into grafts was not seen in an earlier study of similar grafts placed inside distally capped rather than open-ended channels. Additional intervention will be required to foster growth of the regenerated axons from the graft into the distal cord tissue.     

Segmental mode of neural patterning in sipuncula

Recent molecular phylogenetic analyses suggest a close relationship between two worm-shaped phyla, the nonsegmented Sipuncula (peanut worms) and the segmented Annelida (e.g., earthworms and polychaetes) [1-5]. The striking differences in their bodyplans are exemplified by the annelids' paired, ladder-like ventral nervous system, which contains segmentally arranged ganglia, and the sipunculans' single ventral nerve cord (VNC), which is devoid of any segmental structures [6, 7]. Investigating central nervous system (CNS) formation with serotonin and FMRFamide labeling in a representative sipunculan, Phascolosoma agassizii, we found that neurogenesis initially follows a segmental pattern similar to that of annelids. Starting out with paired FMRFamidergic and serotonergic axons, four pairs of associated serotonergic perikarya and interconnecting commissures form one after another in an anterior-posterior progression. In late-stage larvae, the two serotonergic axons of the VNCs fuse, the commissures disappear, and one additional pair of perikarya is formed. These cells (ten in total) migrate toward one another, eventually forming two clusters of five cells each. These neural-remodeling processes result in the single nonmetameric CNS of the adult sipunculan. Our data confirm the segmental ancestry of Sipuncula and render Phascolosoma a textbook example for the Haeckelian hypothesis of ontogenetic recapitulation of the evolutionary history of a species [8].     

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.     

An evaluation of myelomeres and segmentation of the chick embryo spinal cord.

We have investigated whether the neuromeres of the developing chick spinal cord (myelomeres) are manifestations of intrinsic segmentation of the CNS by studying the patterns of cell proliferation and neuronal differentiation. Treatment of 2-day embryos with colchicine does produce exaggerated myelomeres, in confirmation of K?llén (Z. Anat. Entwickl.-Gesch. 123, 309-319, 1962). However, this does not imply that myelomeres are segmental proliferation centres: the undulations caused by colchicine are irregular alongside the unsegmented mesoderm, and another mitotic inhibitor, bromodeoxyuridine, has no such effects. In contrast to lower vertebrate embryos, there is no evidence for segmental groups of primary motor neurons in the chick: the earliest motor neurons express cholinesterase, and project their axons into the adjacent sclerotome, at random positions in relation to the somite boundaries. The population of motor neurons projecting HRP-labelled axons into a single somite lies out of phase with both myelomere and somite, and is placed symmetrically about the anterior half-sclerotome. The earliest intrinsic spinal cord neurons, as stained with zinc iodide-osmium tetroxide or anti-68 x Mr neurofilament antibody, show no segmental patterns of differentiation. We conclude that, in contrast to the rhombomeres of the developing hindbrain, myelomeres are not matched by segmental groupings of differentiating nerve cells, and result from mechanical moulding of the neuroepithelium by the neighbouring somites.     

Aminergic neuron systems of lobsters: morphology and electrophysiology of octopamine-containing neurosecretory cells

In the American lobster (Homarus americanus) the biogenic amines serotonin and octopamine appear to play important and opposite roles in the regulation of aggressive behavior, in the establishment and/or maintenance of dominant and subordinate behavioral states and in the modulation of the associated postural stances and escape responses. The octopamine-containing neurosecretory neurons in the thoracic regions of the lobster ventral nerve cord fall into two morphological subgroups, the root octopamine cells, a classical neurohemal group with release regions along second thoracic roots, and the claw octopamine cells, a group that selectively innervates the claws. Cells of both subgroups have additional sets of endings within neuropil regions of ganglia of the ventral nerve cord. Octopamine neurosecretory neurons generally are silent, but when spontaneously active or when activated, they show large overshooting action potentials with prominent after-hyperpolarizations. Autoinhibition after high-frequency firing, which is also seen in other crustacean neurosecretory cells, is readily apparent in these cells. The cells show no spontaneous synaptic activity, but appear to be excited by a unitary source. Stimulation of lateral or medial giant axons, which excite serotonergic cells yielded no response in octopaminergic neurosecretory cells and no evidence for direct interactions between pairs of octopamine neurons, or between the octopaminergic and the serotonergic sets of neurosecretory neurons was found.     

Distribution of serotonergic neurons in the eyestalk and brain of the crab,Cancer antennarius

1. Serotonin-containing neurons were localized immunocytochemically in crab cerebral ganglia and their extensions in the eyestalk.2. Approximately 155 serotonergic cells were found in identifiable regions of the brain, the largest number being localized in the anterior cell cluster (40 reactive cells) and the bilateral anterior olfactory cell clusters (40 cells each).3. Serotonin immunoreactive cells were found in all three ganglionic divisions of the eyestalk. The medulla terminalis contains up to 15 reactive cells, of which only one occurs in the X-organ (origin of neurosecretory axons in the sinus gland nerve). The m. terminalis also contains three identifiable cells in the mediolateral border adjacent to the sinus gland nerve, of which one is a giant (up to 100 μm diameter), designated MT-1. The axon of MT-1 branches profusely after entering the m. terminalis neuropil.4. No serotonin immunoreactivity was apparent within the sinus gland, the sinus gland nerve or the organ of Bellonci.5. These findings are discussed in relation to the known serotonergic control of peptide hormone secretion by the eyestalk X-organ-sinus gland complex.     

Development of Serotonergic Neurons in Embryos of the Sea Urchin, Strongylocentrotus purpuratus

The development of the serotonergic component of the nervous system of larvae of S. purpuratus is traced using indirect immunofluorescence with a polyclonal antibody against the neurotransmitter serotonin. Initially one or two neuroblasts can be detected in the thickened epithelium of the animal plate of late gastrulae (56 hr). The number of immunoreactive cells increases to about eight during formation of the pluteus (85–90 hr). Immunoreactive axons appear simultaneously from all neuroblasts present in the 79 hr prism stage larva and form the apical ganglion. It is proposed that this component of the larval nervous system is derived from a small number of ectodermal cells associated with the apical tuft.     

Immunocytochemical demonstration of neuropeptides and serotonin in the tapeworm Diphyllobothrium dendriticum

Summary The present immunocytochemical study concerns the distribution of four neuropeptides, FMRF-amide, vasotocin, leu-enkephalin and neurotensin, and of the bioamine serotonin in the plerocercoid larva of Diphyllobothrium dendriticum. Anti-FMRF-amide and vasotocin-reactivity occurs in perikarya and nerve fibres in the CNS and PNS of this worm. The peptide-containing fibres surround and seem to innervate the musculature and to terminate beneath the basal lamina of the tegument at the inner surface of the bothridia, suggesting a neurotransmitter function. Antileu-enkephalin reaction occurs in perikarya and fibres in the main nerve cords and in the PNS. Anti-neurotensin reactive fibres were observed in the neuropile of the nerve cords. Serotonin immunoreactivity was found in neurons in the ganglionic commissure of the brain and along the main nerve cords. This study is the first immunocytochemical identification of neuropeptides and serotonin in a parasitic flatworm and the information gained may be of importance for the development of new antihelminthics.     

Anatomy of the serotonergic nervous system of an entoproct creeping-type larva and its phylogenetic implications

Abstract. Although the internal phyletic relationships of Spiralia (and Lophotrochozoa) remain unresolved, recent progress has been made due to molecular phylogenetic analyses as well as developmental studies of crucial taxa such as Mollusca, Sipuncula, or Annelida. Despite this progress, the phylogenetic position of a number of phyla, such as Entoprocta, remains problematic, mainly due to their unique morphology, their aberrant mode of development, and their exclusion in most large-scale phylogenetic analyses. In order to extend the morphological dataset of this enigmatic taxon, we herein describe the anatomy of the serotonergic nervous system of the creeping-type larva of Loxosomella murmanica . The apical organ is very complex and comprises six to eight centrally positioned flask cells and eight bipolar peripheral cells. In addition, a prototroch nerve ring, an anterior nerve loop, a paired buccal nerve, and an oral nerve ring are found. Moreover, the larva of L. murmanica has one pair of pedal and one pair of lateral longitudinal nerve cords and thus expresses a tetraneurous condition. Several paired serotonergic perikarya, which form contact with the pedal nerve cords but not with the lateral ones, are found along the anterior–posterior axis. The combination of a complex larval serotonergic apical organ and (adult) tetraneury, comprising one pair of ventral and one pair of more dorsally situated lateral longitudinal nerve cords without ganglia, has so far only been reported for basal molluscs and may be diagnostic for a mollusc–entoproct clade. In addition, the larva of Loxosomella expresses a mosaic of certain neural features that are also found in other larval or adult Spiralia, e.g., a prototroch nerve ring, an anterior nerve loop, and a buccal nervous system.     

Expression patterns of HNK-1 carbohydrate and serotonin in sea urchin, amphioxus, and lamprey, with reference to the possible evolutionary origin of the neural crest

We examined deuterostome invertebrates, the sea urchin and amphioxus, and an extant primitive vertebrate, the lamprey, for the presence of structures expressing the HNK-1 carbohydrate and serotonin. In sea urchin embryos and larvae, HNK-1 positive cells were localized in the ciliary bands and in their precursor ectoderm. Serotonergic cells were exclusively observed in the apical organs. In juvenile amphioxus, primary sensory neurons in the dorsal nerve cords were HNK-1 immunoreactive. The juvenile amphioxus nerve cords contained anti-serotonin immunoreactive nerve fibers that seem to be the Rohde axons extending from amphioxus interneurons, the Rohde cells. In lamprey embryos, migrating neural crest cells and primary sensory neurons, including Rohon-Beard cells, expressed the HNK-1 carbohydrate. Lamprey larvae (ammocoetes) contained cell aggregates expressing both the HNK-1 carbohydrate and serotonin in the pronephros and in the regions adjacent to the gut epithelium. Some of these cell aggregates were present in the anti-serotonin positive visceral motor nerve net. Motor neurons and Müller fibers were serotonergic in ammocoetes. Comparison of the expression patterns of HNK-1 carbohydrate among sea urchins, amphioxus and lampreys seem to suggest the possible evolutionary origin of the neural crest, that is, ciliary bands in dipleurula-type ancestors evolved into primary sensory neurons in chordate ancestors, as inferred from Garstang's auricularia hypothesis, and the neural crest originated from the primary sensory neurons.     

Diffusible signals and fasciculated growth in reticulospinal axon pathfinding in the hindbrain

We have addressed the control of longitudinal axon pathfinding in the developing hindbrain, including the caudal projections of reticular and raphe neurons. To test potential sources of guidance signals, we assessed axon outgrowth from embryonic rat hindbrain explants cultured in collagen gels at a distance from explants of midbrain-hindbrain boundary (isthmus), caudal hindbrain, or cervical spinal cord. Our results showed that the isthmus inhibited caudally directed axon outgrowth by 80% relative to controls, whereas rostrally directed axon outgrowth was unaffected. Moreover, caudal hindbrain or cervical spinal cord explants did not inhibit caudal axons. Immunohistochemistry for reticular and raphe neuronal markers indicated that the caudal, but not the rostral projections of these neuronal subpopulations were inhibited by isthmic explants. Companion studies in chick embryos showed that, when the hindbrain was surgically separated from the isthmus, caudal reticulospinal axon projections failed to form and that descending pioneer axons of the medial longitudinal fasciculus (MLF) play an important role in the caudal reticulospinal projection. Taken together, these results suggest that diffusible chemorepellent or nonpermissive signals from the isthmus and substrate-anchored signals on the pioneer MLF axons are involved in the caudal direction of reticulospinal projections and might influence other longitudinal axon projections in the brainstem.     

The ultrastructure and innervation of the ear of the gar, Lepisosteus osseus

The endorgans of the inner ear of the gar were examined using transmission and scanning electron microscopy as well as nerve staining. The ultrastructure of the sensory hair cells and supporting cells of the gar ear are similar to cells in other bony fishes, whereas there are significant differences between the gar and other bony fishes in the orientations patterns of the sensory hair cells on the saccular and lagenar sensory epithelia. The saccular sensory epithelium has two regions, a main region and a secondary region ventral to the main region. The ciliary bundles on the main region are divided into two groups, one oriented dorsally and the other ventrally. Furthermore, as a result of curvature of the saccular sensory epithelium, the dorsal and ventral ciliary bundles on the rostral portion of the epithelium are rotated ninety degrees and are thus oriented on the animal's rostro-caudal axis. Hair cells on the secondary region are generally oriented ventrally. The lagenar epithelium has three groups of sensory hair cells. The groups on the rostral and caudal ends of the macula are oriented dorsally, whereas the middle group is oriented ventrally. Hair cell orientations on the utricular epithelium and macula neglecta are similar to those in other bony fishes. Nerve fiber diameters can be divided into three size classes, 1-8 microns, 9-13 microns, and 14 microns or more, with the smallest size class containing the majority of fibers. The distribution of the various classes of fiber diameters is not the same in nerve branches to each of the end organs. Similarly, the ratio of hair cells to axons differs in each end organ. The highest hair cell to axon ratio is in the utricle (23:1) and the smallest is in the macula neglecta (7:1). The number of sensory hair cells far exceed the number of eighth nerve axons in all sensory epithelia.     

Acetylcholinesterase-containing nerve cells and their distribution in the pineal organ of the goldfish,Carassius auratus

Summary Histochemically, an intense acetylcholinesterase (AChE) reaction has been observed in the perikarya of the nerve cells and in the neuropil formations of the pineal organ in the goldfish, Carassius auratus. A group of AChE-rich nerve cells has also been observed between the caudal end of the pineal stalk and the habenular ganglion. No component of the complex revealed butyrylcholinesterase (BuChE) activity.Two different types of nerve cells were recognized on the basis of their size, AChE activity and distribution. Type I cells are characterized by large perikarya possessing a moderate AChE activity and by the presence of an extensive AChE-rich neuropil formation in their vicinity; they are restricted to the rostro-lateral regions of the pineal vesicle. Type II cells are situated in the medio-rostral area of the pineal vesicle and along the entire length of the stalk, and are smaller than Type I cells; they show an intense AChE activity in their perikarya.The neuropil formations in the medio-rostral area of the pineal vesicle are almost as large as those in the vicinity of the Type I cells; they exhibit a strong AChE activity. In the rostral half of the vesicle several sensory cells are associated with each nerve cell, while in the caudal portion only a few cells are apposed to each nerve cell. Thus, the ratio of the number of sensory cells to that of AChE-containing nerve cells in the anterior half of the pineal vesicle is high when compared with the remaining area. In the anterior half of the vesicle the outer segments of the sensory cells are more distinct and their inner segments possess a higher AChE activity than those in the posterior region and the stalk. A gradation in the degree of development of neuropil formations along the pineal axis is remarkable; their size and AChE activity gradually diminish in a caudal direction. In view of the structural specialization of the rostral region of the pineal organ, it has been argued that its terminal portion is more photosensitive.This work was supported by a fellowship from the Alexander von Humboldt Foundation, Federal Republic of Germany.