The 'goatee' of goatfish: innervation of taste buds in the barbels and their representation in the brain

Goatfish use a pair of large chin barbels to probe the sea bottom to detect buried prey. The barbels are studded with taste buds but little else is known about the neural organization of this system. We found that the taste buds of the barbel are innervated in a strict orthogonal fashion. The barbel is innervated by a main nerve trunk running in the core of the barbel. A longitudinal nerve bundle originates from the main trunk and, after running a short distance distally, divides into two circumferential nerve bundles (CNB) extending respectively, medially and laterally around the barbel. Approximately 15 CNBs innervate each 1 mm length of barbel. At each transverse level, the CNB innervates two clusters of taste buds, each containing 14 end-organs. The primary taste centre in the brain is similarly extraordinary. The sensory inputs from the barbel terminate in a derived dorsal facial lobe, which has a highly convoluted surface forming a multitude of tubercles. Electrophysiological mapping experiments show that the entire barbel is somatotopically represented in a recurved elongate tubular fashion within the dorsal facial lobe.     

Morphology of the brain of Sarotherodon mossambicus (Peters) and correlation with its habits

The correlation between the brain of an exotic fish, Sarotherodon mossambicus, and its habits was studied. The poorly developed olfactory lobes are indicative of poor olfactory sense, while the large optic lobes and stout, thick optic nerves correlate with this fish's excellent sense of vision. Similarly, its tactile and gustatory sense are determined by the extent of development of the facial and vagal lobes.     

Mechanical sensitivity of the facial nerve fibers innervating the anterior palate of the puffer,Fugu pardalis,and their central projection to the primary taste center

Mechanical and chemical sensitivity of the palatine nerve, ramus palatinus facialis, innervating the anterior palate of the puffer, Fugu pardalis, and their central projection to the primary taste center were investigated. Application of horseradish peroxidase (HRP) to the central cut end of the palatine nerve resulted in retrogradely labeled neurons in the geniculate ganglion but no such neurons in the trigeminal ganglion, suggesting that the palatine nerve is represented only by the facial component. Tracing of the facial sensory root in serial histological sections of the brain stem suggested that the facial sensory nerve fibers project only to the visceral sensory column of the medulla. Peripheral recordings from the palatine nerve bundle showed that both mechanical and chemical stimuli caused marked responses. Mechanosensitive fibers were rather uniformly distributed in the nerve bundle. Intra-cranial recordings from the trigeminal and facial nerves at their respective roots revealed that tactile information produced in the anterior palate was carried by the facial nerve fibers. Elimination of the sea water current over the receptive field also caused a marked response in the palatine nerve bundle or facial nerve root while this did not cause any detectable responses in the trigeminal nerve root. Single fiber analyses of the mechanical responsiveness of the palatine nerve were performed by recording unit responses of 106 single fibers to mechanical stimuli (water flow), HCl (0.005 M), uridine-5'-monophosphate (UMP, 0.001 M), proline (0.01 M), CaCl2 (0.5 M), and NaSCN (0.5 M). All these fibers responded well to one of the above stimuli; however, most taste fibers did not respond well to the inorganic salts. The palatine fibers (n = 36), identified as mechanosensitive, never responded to any of the chemical stimuli, whereas chemosensitive fibers (n = 70) did not respond to mechanical stimuli at all. The chemosensitive units showed a high specificity to the above stimuli: they tended to respond selectively to hydrochloric acid, UMP, or proline. The responses of the mechanosensitive units consisted of phasic and tonic impulse trains and the sensitivity of the units varied considerably. The results reveal that the facial nerve fibers innervating the anterior palate of the puffer contain two kinds of afferent fibers, chemosensory and mechanosensory respectively, and suggest that the convergence of the tactile and gustatory information first occurs in the neurons of the primary gustatory center in the medulla.     

Morphologic characteristics and responses of the neurons of the right parietal ganglion in Lymnaea stagnalis to stimulation of the sensory structures

Intracellular recordings have been made of the responses of 22 neurons of the central part of the dorsal surface of the right parietal ganglion of the snail Lymnaea stagnalis to adequate stimulation of chemo-, photo- and mechanoreceptor cells of the mantle and head skin including tentacles and lips. It was shown that the main bulk of the neurons investigated has broad receptive fields in the body wall and mantle, being able to respond to all types of the applied stimuli. Alongside, single neurons were revealed which receive single-mode input, either a mechanosensory or chemosensory one. Morphological studies indicate that the neurons are unipolar and have usually one, sometimes several projections. They differ in the pattern of branching as well as in the projections to peripheral nerves. However, almost all of them have vast dendritic regions in the central nervous system including central sensory nucleus of the right parietal ganglion.     

Cephalic sensory pathways in the central nervous system of larval Manduca sexta (Lepidoptera : Sphingidae)

Central projections of neurons innervating sensory structures on the head of larval Manduca sexta were traced by using methods of anterograde cobalt-diffusion. Regions of the deutocerebrum and tritocerebrum in the brain receive input from the antenna, labrum, maxilla, labial palps, hypopharynx and other unidentified regions of the buccal cavity. Antennal, maxillary and labial inputs project to the larval antennal centre (LAC) of the deutocerebrum. Stemmatal neurons and a few antennal neurons project into the protocerebrum. The suboesophageal ganglion (SEG) receives input from mechanosensory neurons in all parts of the head and its sensory appendages. Some mechanosensory neurons project further to the first thoracic ganglion. In addition to receiving input from chemosensory neurons of the maxilla, the SEG may also receive chemosensory input from epipharyngeal sensilla of the labrum.     

Unit responses in the vagal lobe of the carp brain to stimulation of chemoreceptors of the skin,mouth, and gills

Experiments on curarized carp showed that neurons of the vagal lobe of the medulla respond to stimulation not only of the mouth and gills, but also of chemoreceptors of the skin of the head. The vagal lobe was shown to contain considerably fewer neurons responding selectively to stimulation of the skin than neurons responding to stimulation of the mouth or gills. Differences were found between responses of neurons to a natural food stimulus (extract from a fish food product) and to solutions of hydrochloric acid and common salt. The different roles of the vagal and facial lobes in the processing of impulsation from skin chemoreceptors is demonstrated.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 12, No. 4, pp. 397–404, July–August, 1980.     

Overlaping taste and tactile maps of the oropharynx in the vagal lobe of the channel catfish,Ictalurus punctatus

Microelectrode mapping experiments indicate an ipsilateral representation of the oropharynx and a well-defined, bilateral input from the proximal portion of the maxillary barbels and snout region within the vagal lobe of channel catfish. The map of the oropharyngeal epithelium is distorted so that the gill arches are rotated through an angle of 90° along the transverse plane, and the dorsally mapped region of the gill rakers is tilted posteriorly in the sagittal plane of the vagal lobe. Multiunit recording studies fail to provide definitive boundaries of adjacently mapped domains of oropharyngeal structures. Gustatory receptive fields of neurons in the vagal lobe correspond to their location on the topological map obtained by tactile stimulation of the oropharyngeal epithelium. A few single unit recordings indicate restricted receptive fields and different response patterns of taste, tactile, and proprioceptive neurons in the vagal lobe of catfish.     

Overlapping taste and tactile maps of the oropharynx in the vagal lobe of the channel catfish, Ictalurus punctatus

Microelectrode mapping experiments indicate an ipsilateral representation of the oropharynx and a well-defined, bilateral input from the proximal portion of the maxillary barbels and snout region within the vagal lobe of channel catfish. The map of the oropharyngeal epithelium is distorted so that the gill arches are rotated through an angle of 90 degrees along the transverse plane, and the dorsally mapped region of the gill rakers is tilted posteriorly in the sagittal plane of the vagal lobe. Multiunit recording studies fail to provide definitive boundaries of adjacently mapped domains of oropharyngeal structures. Gustatory receptive fields of neurons in the vagal lobe correspond to their location on the topological map obtained by tactile stimulation of the oropharyngeal epithelium. A few single unit recordings indicate restricted receptive fields and different response patterns of taste, tactile, and proprioceptive neurons in the vagal lobe of catfish.     

Analysis of tonic activity in sympathetic ganglionic neurons in rabbits

Tonic activity of neurons of the rabbit superior cervical ganglion was studied by intracellular recording. This activity was compared with responses of these same neurons to single stimulation of preganglionic fibers. Neurons of the ganglion were found to have two synaptic inputs — principal and accessory. The principal input consists of one preganglionic fiber, the discharge in which evokes an action potential in the neuron with a high safety factor. The accessory input consists of two or more converging fibers, and the neuron discharges only as a result of summation of EPSPs evoked by activation of these fibers.     

Labellar lobe spreading in the blowfly: Regulation by taste and satiety

Summary Lobe spreading behavior was studied by recording electromyograms from the muscles which spread the labellar lobes, the retractors of the furca (RF) inPhormia regina. RF responses and lobe spreading could be elicited by stimulating labellar, but not tarsal, taste hairs with sucrose (Fig. 3). RF activity was important to spread the lobes at the beginning of a meal, but was not necessary for continued feeding (Fig. 4).Temporal summation between sugar receptor spikes was necessary to elicit RF responses. Central response decrement occurs independently for different labellar hairs and may participate in the termination of motor responses.RF responses were more probable and more intense when either the sucrose concentration of the stimulus or the number of hairs stimulated was increased (Fig. 7). Stimulation with NaCl had no effect on the response to simultaneous sucrose stimulation of other hairs (Table 1).Feeding caused decreases in the probability and intensity of motor responses, but did not alter chemosensory responses (Figs. 8 and 9). Section of either the recurrent or median abdominal nerves prevented this postingestional inhibition of lobe spreading (Fig. 9).These results are discussed with regard to the possible role that regulation of lobe spreading may play in the control of food intake.This work was supported by United States Public Health Service Training Grant 5T01 GM 00457-13S2 and by a grant from the National Science Foundation to Dr. Vincent G. Dethier. I wish to thank Dr. Dethier for his support and encouragement.     

A chemosensory gene family encoding candidate gustatory and olfactory receptors in Drosophila

A novel family of candidate gustatory receptors (GRs) was recently identified in searches of the Drosophila genome. We have performed in situ hybridization and transgene experiments that reveal expression of these genes in both gustatory and olfactory neurons in adult flies and larvae. This gene family is likely to encode both odorant and taste receptors. We have visualized the projections of chemosensory neurons in the larval brain and observe that neurons expressing different GRs project to discrete loci in the antennal lobe and subesophageal ganglion. These data provide insight into the diversity of chemosensory recognition and an initial view of the representation of gustatory information in the fly brain.     

Long ascending fibers in the dorsal column of a teleost fish: a disynaptic pathway connecting sense organs to cerebellum]

In spite of the generally accepted opinion that long ascending proprioceptive and tactile fibers do not occur in the spinal dorsal columns of teleost fish, it was demonstrated with degeneration and axonal transport tracing methods that such dorsal column fibers exist in the teleost fish Gnathonemus petersii. These fibers are in fact common spinal afferent fibers originating in spinal ganglion cells. They connect the peripheral sense organs with the lateral funicular nuclei (Fl2) in which the dorsal column fibers terminate, directly through the dorsal columns. In contrast to the dorsal column nuclei of higher vertebrates, the Fl2 nuclei do not project to the diencephalic thalamus but to the caudal lobe and the second lobe (C2) of the corpus cerebelli. Thus, sense organs and cerebellum are connected by a disynaptic pathway. Since the caudal lobe projects directly to the electrosensory lobe, that is, to the target of electrosensory afferents, the presence of a disynaptic pathway in G. petersii suggests the existence of a proprioceptive control of the electrosensory input.     

The brain structure of selected trypanorhynch tapeworms

The brain architecture in four species of tapeworms from the order Trypanorhyncha has been studied. In all species, the brain consists of paired anterior and lateral lobes, and an unpaired central lobe. The anterior lobes connect by dorsal and ventral semicircular commissures; the central and lateral lobes connect by a median and an X-shaped crisscross commissure. In the center of the brain, five well-developed compact neuropils are present. The brain occupies a medial position in the scolex pars bothrialis. The ventral excretory vessels are situated outside the lateral lobes of the brain; the dorsal excretory vessels are located inside the brain and dorsal to the median commissure. The brain gives rize four anterior proboscis nerves and four posterior bulbar nerves with myelinated giant axons (GAs). The cell bodies of the GAs are located within the X-commissure and in the bulbar nerves. Highly developed serotonergic neuropils are present in the anterior and lateral lobes; numerous 5-HT neurons are found in the brain lobes including the central unpaired lobe. The X-cross commissure consists of the α-tub-immunoreactive and 5-HT-IR neurites. Eight ultrastructural types of neurons were found in the brain of the three species investigated. In addition, different types of synapses were present in the neuropils. Glial cells ensheath the brain lobes, the neuropils, the GAs, and the bulbar nerves. Glia cell processes form complex branching patterns of thin cytoplasmic sheets sandwiched between adjacent neural processes and filling the space between neurons. Multilayer myelin-like envelopes and a mesaxon-like structure have been found in Trypanorhyncha nervous system. We compared the brain architecture of Trypanorhyncha with that of an early basal cestode taxon, that is, Diphyllobothriidea, and present a hypothesis about the homology of the anterior brain lobes in order Trypanorhyncha; and the lateral lobes and median commissure are homologous brain structures within Eucestoda.     

Recurrent facial taste neurons of sea catfish Plotosus japonicus: morphology and organization in the ganglion

This study investigated the morphology of the recurrent facial taste neurons and their organization in the recurrent ganglion of the sea catfish Plotosus japonicus. The recurrent ganglion is independent of the anterior ganglion, which consists of trigeminal, facial and anterior lateral line neurons that send peripheral fibres to the head region. The recurrent taste neurons are round or oval and bipolar, with thick peripheral and thin central fibres, and completely wrapped by membranous layers of satellite cells. Two peripheral nerve branches coursing to the trunk or pectoral fin originate from the recurrent ganglion. The results presented here show that the trunk and pectoral‐fin neurons are independently distributed to form various sizes of groups, and the groups are intermingled throughout the ganglion. No distinct topographical relationship of the two nerve branches occurs in the ganglion. Centrally, the trunk and pectoral‐fin branches project somatotopically in the anterolateral and intermediate medial regions of the trunk tail lobule of the facial lobe, respectively.     

The chemosensory anterior dorsal fin in rocklings (Gaidropsarus and Ciliata, Teleostei, Gadidae): activity, fine structure and innervation

Observations and experiments on the behaviour of shore rocklings have shown that the modified and vibratile anterior dorsal fin can be involved in the detection of food but is not essential to foraging by the fish. The epidermis of the vibratile fin rays contains numerous chemosensory cells, of similar cytology in the two species studied. These chemosensory cells are compared with the gustatory cells of the taste buds borne on other fin rays. Synaptic modifications in both cases consist of densities on the apposed membranes, with a dense layer under the membrane of the neurite more distinct than that in the cell. Vesicles are not a feature of these synapses, although some of the sensory cell bases are vesicular. Denervation experiments have shown that the chemosensory cells of the vibratile rays are supplied by a facial nerve component. After denervation a small proportion of the sensory cells were found to have an association with spinal nerve fibres. The present status of solitary chemosensory cells in fishes is discussed.     

Nitric oxide synthase in the glossopharyngeal and vagal afferent pathway of a teleost, Takifugu niphobles

To examine the presence of nitric oxide synthase (NOS) in the sensory system of the glossopharyngeal and vagus nerves of teleosts, nicotinamide adenine dinucleotide phosphate diaphorase (NADPHd) activity and immunoreactivity for NOS were examined in the puffer fish Takifugu niphobles. The nitrergic sensory neurons were located in the ganglia of both the glossopharyngeal and the vagal nerves. In the vagal ganglion, positive neurons were found in the subpopulations for the branchial rami and the coelomic visceral ramus, but not for the posterior ramus or the lateral line ramus. In the medulla, nitrergic afferent terminals were found in the glossopharyngeal lobe, the vagal lobe, and the commissural nucleus. In the gill structure, the nitrergic nerve fibers were seen in the nerve bundles running along the efferent branchial artery of all three gill arches. These fibers appeared to terminate in the proximal portion of the efferent filament arteries of three gill arches. On the other hand, autonomic neurons innervating the gill arches were unstained. These results suggest that nitrergic sensory neurons in the glossopharyngeal and vagal ganglia project their peripheral processes through the branchial rami to a specific portion of the branchial arteries, and they might play a role in baroreception of this fish. A possible role for nitric oxide (NO) in baroreception is also discussed.     

Solitary chemosensory cells — taste,common chemical sense or what?

Summary Secondary solitary chemosensory cells (SCCs) occur scattered within the epidermis of lampreys, teleosts and ranid tadpoles. Counts in representative telost species revealed that SCC's outnumber chemosensory cells organized in taste buds. Therefore, SCCs may be considered the structural substrate of a basic and probably important vertebrate chemosense. However, detailed information on structure, innervation and function is only available from specialized fins in a few teleost species, where SCCs are sufficiently concentrated. The foremost research model has been the anterior dorsal fin (ADF) in rocklings, which contains millions of SCCs but no other specialized chemosensory elements. It has been shown that these ADF-SCCs are innervated from the recurrent facial nerve. Electrophysiological recordings revealed that there is virtually no overlap in stimulus spectrum between the ADF-SCCs and pelvic fin taste buds; SCC responses could only be triggered by dilutions of heterospecific fish body mucus. Results of behavioural experiments indicate that fish mucus is indeed a relevant stimulus. Therefore it is hypothesized that the biological role of the ADF-SCCs is predator avoidance rather than search for food. Whether these findings are valid for rockings only, or can be generalized for the scattered SCC systems in more than 20000 species of fish and in some amphibians, remains an open question. Further investigations on the function and biological roles of the SCC chemosense will be crucially important to improve our understanding of sensory perception and its evolution in aquatic vertebrates.     

Central projections of a homoeotic regenerate, antennapedia, in a stick insect, Carausius morosus (Phasmida)

Homoeotic appendages provide a system for the analysis of neural path-finding in which the appendage is mismatched with its segmented ganglion. Central projections of sensory neurons from homoeotic antennapedia regenerates induced by antennal amputation in the stick insect, Carausius morosus, are described. The majority of afferent axons project to the olfactory lobe as in the normal antennal nerve, but they do not give rise to compact glomeruli. Nor does the form of the projection resemble that of leg sensory nerves in thoracic ganglia. The projection of antennapedia regenerate neurons in Carausius resembles the antennapedia mutant of Drosophila except that some primary afferents bypass the olfactory lobe and take several courses through the brain, sometimes reaching distant contralateral areas. It appears that these wandering fibers, having bypassed the olfactory lobe, tend to follow established tracts and to arborize or to deviate at circumscribed synaptic areas. The behavioral evidence for sensory input from antennapedia regenerates is equivocal.     

Excitatory neural control of posterograde heartbeat by the frontal ganglion in the last instar larva of a lepidopteran, Bombyx mori

The frontal ganglion of the silkworm (Bombyx mori) gives rise to a visceral nerve, branches of which include a pair of anterior cardiac nerves and a pair of the posterior cardiac nerves. Forward-fill of the visceral nerve with dextran labeled with tetramethyl rhodamine shows the anterior cardiac nerves innervate the anterior region of the dorsal vessel. Back-fill of the anterior cardiac nerves with Co²⁺ and Ni²⁺ ions and the fluorescent dye reveals that the cell bodies of two motor neurons are located in the frontal ganglion. Injection of 5, 6-carboxyfluorescein into the cell body of an identified motor neuron shows that the neuron gives rise to an axon running to the visceral nerve. Unitary excitatory junctional potentials (EJPs) were recorded from a myocardial cell at the anterior end of the heart. They responded in a one-to-one manner to electrical stimuli applied to the visceral nerve, or to impulses generated by a depolarizing current injected into the cell body. EJPs induced by stimuli at higher than 0.5 Hz showed facilitation while those induced at higher than 2 Hz showed summation. Individual EJPs without summation, or a train of EJPs with summation, caused acceleration in the phase of posterograde heartbeat and heart reversal from anterograde heartbeat to posterograde heartbeat. It is likely that the innervation of the anterior region of the dorsal vessel by the motor neurons, through the anterior cardiac nerves is responsible for the control of heartbeat in Lepidoptera, at least in part.     

The vasopressin-like immunoreactive (VPLI) neurons of the locust,Locusta migratoria. I. Anatomy

Summary Antiserum to arginine-vasopressin has been used to characterise the pair of vasopressin-like immunoreactive (VPLI) neurons in the locust. These neurons have cell bodies in the suboesophageal ganglion, each with a bifurcating dorsal lateral axon which gives rise to predominantly dorsal neuropilar branching in every ganglion of the ventral nerve cord. There are extensive beaded fibre plexuses in most peripheral nerves of thoracic and abdominal ganglia, but in the brain, the peripheral plexuses are reduced while neuropilar branching is more extensive, although it generally remains superficial. An array of fibres runs centripetally through the laminamedulla chiasma in the optic lobes. Lucifer Yellow or cobalt intracellular staining of single VPLI cells in the adult suboesophageal ganglion shows that all immunoreactive processes emanate from these two neurons, but an additional midline arborisation (that was only partially revealed by immunostaining) was also observed. Intracellularly staining VPLI cells in smaller larval instars, which permits dye to reach the thoracic ganglia, confirms that there is no similar region of poorly-immunoreactive midline arborisation in these ganglia. It has been previously suggested that the immunoreactive superficial fibres and peripheral plexuses in ventral cord ganglia serve a neurohaemal function, releasing the locust vasopressin-like diuretic hormone, F₂. We suggest that the other major region of VPLI arborisation, the poorly immunoreactive midline fibres in the suboesophageal ganglion, could be a region where VPLI cells receive synaptic input. The function of the centripetal array of fibres within the optic lobe is still unclear.Abbreviations AVP arginine vasopressin - DIT dorsal intermediate tract - FLRF Phe-Leu-Arg-Phe - FMRF-amide Phe-Met-Arg-Phe-amide - LDT lateral dorsal tract - LVP lysine vasopressin - MDT median dorsal tract - MVT median ventral tract - SEM scanning electron microscopy - SOG suboesophageal ganglion - VIT ventral intermediate tract - VNC ventral nerve cord - VPLI vasopressin-like immunoreactive