Ocular dominance stripe formation by regenerated isogenic double temporal retina in Xenopus laevis

In lower vertebrates such as frogs and fish, long ocular dominance stripes with anterior-posterior (A-P) orientation can be produced by causing both eyes to innervate one optic tectum during the course of development. Similar experiments on adult animals usually produce patches rather than stripes. During development, new retinal fibers from the nasal retina segregate into appropriate stripes at the growing edge of the posterior (P) tectum while new temporal fibers segregate at the non-growing anterior (A) tectal edge. Fiber segregation into long A-P oriented stripes might depend upon a template produced by new nasal fibers initiating stripe orientation in the vicinity of new tectal cells; new nasal fibers would orient to the nascent (posterior) edge of the template while temporal fibers would orient to the anterior (non-growing) end of the template. To test the dependence of stripe formation on the matching of nascent retinal cells with nascent tectal cells, we compared stripe orientation in animals with isogenic double nasal innervation and isogenic double temporal innervation of the tectum. In double nasal innervation, the oldest retinal cells innervate the anterior tectum; new fibers from the entire retinal periphery always innervate the newest tectal cells at the posterior tectum. Stripes are oriented A-P, consistent with a maturation front model. In contrast, the oldest retinal cells innervate the newest (posterior) tectal cells in double temporal innervation of the tectum; the growing retinal periphery innervates the non-growing anterior tectum. Stripes are also oriented A-P, indicating that the production of long stripes does not depend upon maturation front matching of nascent retinal fibers and nascent tectal cells.     

Further serial transmission electron microscopy studies of auditory hair cell innervation in lizards and in a snake

Auditory hair cells of three lizard and one snake species were studied by serial transmission electron microscopy (TEM) sections of two unidirectional hair cells (UHC) and two bidirectional hair cells (BHC) and by nonserial section montages of each entire papilla cut at 2-microns intervals across the papillar width. The unidirectional hair cell region of the agamid lizard, Acanthosaura crucigera, lacked efferent innervation. Another agamid lizard, Agama agama, studied by nonserial section only, also lacked efferent innervation to the UHC. Afferent innervation to both the UHC and BHC of Acanthosaura was primarily exclusive (each nerve fiber innervates only one hair cell), although an occasional nerve fiber innervated two hair cells. Both the UHC and the BHC of the anguid, Celestus costatus, were exclusively innervated. Both hair cell types of the varanid, Varanus exanthematicus, were nonexclusively innervated (all afferent nerve fibers innervate two or more hair cells). The auditory papilla of the colubrid snake, Elaphe obsoleta, has only one type of hair cell and each is nonexclusively innervated. The numbers of afferent and efferent nerve fibers and of afferent synapses are presented in tabular form.     

Neuromuscular organization and regional EMG activity of the pectoralis in the pigeon

In order to improve our understanding of the neuromuscular control of the most massive avian flight muscle, we studied the innervation pattern of the pigeon pectoralis. Nine primary branches from the rostral trunk and nine to ten branches from the caudal trunk of the pectoral nerve were identified by microdissection in ten pigeons. The region of muscle that each branch innervates was delineated by nerve stimulation studies (ten pigeons) and six regions were confirmed by glycogen depletion (ten pigeons). In pigeons, branches from the rostral nerve innervate the anterior 3/5 of the sternobrachialis (SB) head of the pectoralis and branches from the caudal trunk innervate the posterior 1/2 of the SB and all of the throacobrachials (TB). In the SB, individual branches of the rostral pectoral nerve innervate wedge-shaped muscle regions (each approximately 1.3 cm wide), collectively forming a fan shaped arrangement along the sternal carina. Adjacent muscle regions partially overlap at their boundaries. Within the thoracobrachialis (TB) head of the pectrolis, muscle regions are wider. There is a region in mid-SB-where the innervation territories of the rostral and caudal nerves oferlap. Electromyographic (EMG) activity patterns were recorded within ten of the identified muscle regions during take-off, level flapping flight, and landing. Onset of EMG activity and EMG intensity within various muscle regions exhibits significant differences both within a wingbeat cycle and among different modes of flight. The innervation pattern of the pectoralis presents the anatomical substrate for neuromuscular compartmentalization and differential EMG activity within the pectoralis may reflect sensory-motor partitioning. The extent to which the neuromuscular compartmentalization of the pectoralis corresponds to its ability to produce an array of force vectors to the wing awaits further more detailed biomechanical studies. © 1993 Wiley-Liss, Inc.     

Innervation pattern of the gill arches and gills of the carp (Cyprinus carpio)

The innervation pattern of the respiratory gill arches of the carp (Cyprinus carpio) is described. The gill region is innervated by the branchial branches of the glossopharyngeal and vagal nerves. Each branchial nerve divides at the level of or just distal to the epibranchial ganglion into: 1) a pretrematic branch, 2) a dorsal pharyngeal branch, and 3) a posttrematic branch. The dorsal pharyngeal branch innervates the palatal organ in the roof of the buccal cavity. The pretrematic and posttrematic branches innervate the posterior and anterior halves, respectively, of the gill arches bordering a gill slit. Each branch splits into an internal and an external part. The internal bundle innervates the buccal side of the gill arch, including the gill rakers. The external bundle terminates in the gill filaments. The epibranchial motor branch, a small nerve bundle containing only motor fibers, circumvents the ganglion and anastomoses distally with the posttrematic branch. The detailed course and branching patterns of these branches are described.     

Neural response directionality correlates of hair cell orientation in a teleost fish

The otolithic end organs in the ears of teleost fishes play important roles in hearing. Although previous studies have shown that afferent fibers innervating otolithic organs are directionally sensitive to acoustic stimulation, no study has demonstrated that directionality of the otolithic afferent neurons derives directly from morphological polarity of the hair cells that they innervate. In this study we investigated whether or not there exists such a structure and function relationship in one of the otolithic organs, the saccule, by using intracellular and extracellular tracing, histochemistry, and confocal imaging techniques. We observed a variety of morphologies of dendritic terminals of saccular ganglion neurons. Arbor innervation areas of these saccular neurons ranged from 893 microm2 to 21,393 microm2, and the number of dendritic endings fell into a range between 10 and 54. We found that the response directionality of saccular ganglion neurons correlates significantly with the morphological polarization of the hair cells in the regions that they innervate. Therefore, we provide direct evidence to support the hypothesis that fish are able to encode directional information about a sound source, particularly in elevation, using arrays of hair cells in the otolithic organs that are oriented specifically along the sound propagation axis.     

Cranial nerves in the Australian lungfish,Neoceratodus forsteri,and in fossil relatives (Osteichthyes: Dipnoi)

Three systems, two sensory and one protective, are present in the skin of the living Australian lungfish, Neoceratodus forsteri, and in fossil lungfish, and the arrangement and innervation of the sense organs is peculiar to lungfish. Peripheral branches of nerves that innervate the sense organs are slender and unprotected, and form before any skeletal structures appear. When the olfactory capsule develops, it traps some of the anterior branches of cranial nerve V, which emerged from the chondrocranium from the lateral sphenotic foramen. Cranial nerve I innervates the olfactory organ enclosed within the olfactory capsule and cranial nerve II innervates the eye. Cranial nerve V innervates the sense organs of the snout and upper lip, and, in conjunction with nerve IX and X, the sense organs of the posterior and lateral head. Cranial nerve VII is primarily a motor nerve, and a single branch innervates sense organs in the mandible. There are no connections between nerves V and VII, although both emerge from the brain close to each other. The third associated system consists of lymphatic vessels covered by an extracellular matrix of collagen, mineralised as tubules in fossils. Innervation of the sensory organs is separate from the lymphatic system and from the tubule system of fossil lungfish.     

Comparative study of otolith organs between two species of upside-down swimming catfish Synodontis nigriventris showing converse swimming postures

To clarify whether the unique postural control of the upside‐down swimming catfish (Synodontis nigriventris, family Mochokidae) is related to the histological characteristics of the otolith organs, we performed light microscopic observation of the utricle, the saccule and the lagena. The histological aspects of the otolith organs were compared between S. nigriventris and Synodontis multipunctatus, which belong to the same genus. S. multipunctatus usually shows upside‐up swimming posture except for feeding behaviour near water surface. As controls, we additionally used a miniature catfish, Corydoras paleatus and goldfish, Carassius auratus, which shows upside‐up swimming posture. We concluded that the structural aspects of the otolith organs did not cause the unique postural control of S. nigriventris. Light microscopic observation clarified the following aspects: (1) The utricle of S. nigriventris was located at the anterior region of the otocyst and under the semicircular canals, and the saccule and the lagena were located at the posteroventral region of the otocyst like those of S. multipunctatus and the other two fishes. (2) The hair cells of the utricle were arranged on the horizontal plane of the fishes with a variation in cell size at the ventral and ventrolateral sites in S. nigriventris, S. multipunctatus and the other two fishes. (3) The hair cells of the saccule and lagena of S. nigriventris, S. multipunctatus and C. auratus presented perpendicular to the horizontal plane of the fish. (4) Region‐specific differences in the size and shape of the hair cells of S. nigriventris were observed along the three‐dimensional axes of the otolith organs like those of S. multipunctatus and the other two fishes. It is unlikely that the unique postural control of upside‐down catfish is related to the localization of the utricle, the saccule and the lagena and the distribution of the different types of hair cell of the otolith organs. Furthermore, the distribution of the hair cells suggests that the otolith organs in S. nigriventris can detect three‐dimensional postural changes like the organs of other fishes showing generally observed upside‐up swimming posture.     

The membranous labyrinth and its innervation inChaetodon trifasciatus (Pisces,Teleostei, Chaetodontidae)

Synopsis In the butterflyfishChaetodon trifasciatus, the labyrinth is characterized by its elevated form and especially the size of the vertical canals, the almost circular form of the horizontal canal and its posterior opening not directly in the utriculus but in the common pillar of the two vertical canals. There is an almost complete separation between utriculus and sacculus which are only linked by a virtual pore. The lagena, which is medially situated to the posterior part of the sacculus, is separated from it by an incomplete vertical wall. There are two maculae neglectae, the anterior macula being situated in the pore separating utriculus from sacculus and filling this pore, the posterior in a gutter of the floor of the utriculus. A long and narrow endolymphatic canal, originating from the sacculus close to the communication with the utriculus, follows the common pillar of the two vertical canals and widens into an endolymphatic sac at the top of the membranous labyrinth. The innervation of the labyrinth is made by the acoustic ganglion, which is connected to the brain by two roots and elongated into three parts: the anterior part innervates the anterior and horizontal cristae and the utricular and saccular maculae; the middle part innervates the macula sacculae and the macula neglecta 1; the posterior part innervates the macula neglecta II, the macula lagenae and the posterior crista. The important size of the vertical canals and the almost circular form of the horizontal canal may reflect very precise locomotory aptitudes.     

Sensory surface of the saccule and lagena in the ears of ostariophysan fishes

The inner ears of a few fishes in the teleost superorder Ostariophysi are structurally unlike those of most other teleosts. Scanning electron microscopy was used to determine if other ostariophysans share these unusual features. Examined were the families Cyprinidae, Characidae, and Gymnotidae (all of the series Otophysi), and Chanidae (of the sister series Anotophysi), representing the four major ostariophysan lineages, the auditory organs of which have not yet been well described. Among the Otophysi, the saccular and lagenar otolith organs are similar to those reported for other ostariophysans. The lagena is generally the larger of the two organs. The saccular sensory epithelium (macula) contains long ciliary bundles on the sensory hair cells in the caudal region, and short bundles in the rostral region. The saccule and the lagena each have hair cells organized into two groups having opposing directional orientations. In contrast, Chanos, the anotophysan, has a saccular otolith larger than the lagenar otolith, and ciliary bundles that are more uniform in size over most of its saccular macula. Most strikingly, its saccular macula has hair cells organized into groups oriented in four directions instead of two, in a pattern very similar to that in many nonostariophysan teleosts. We suggest that the bi-directional pattern seen consistently in the Otophysi is a derived development related to particular auditory capabilities of these species.     

Structural variation in the inner ears of four deep-sea elopomorph fishes

Deep-sea fishes have evolved in dark or dimly lit environments devoid of the visual cues available to shallow-water species. Because of the limited opportunity for visual scene analysis by deep-sea fishes, it is reasonable to hypothesize that the inner ears of at least some such species may have evolved structural adaptations to enhance hearing capabilities in lieu of vision. As an initial test of this hypothesis, scanning electron microscopy was used to examine the structure of the inner ears of four deep-sea elopomorph species inhabiting different depths: Synaphobranchus kaupii, Synaphobranchus bathybius, Polyacanthonotus challengeri, and Halosauropsis macrochir. The shape of the sensory epithelia and hair cell ciliary bundle orientation of the saccule, lagena, and utricle, the three otolithic organs associated with audition and vestibular function, are described. The saccules of all four species have a common, alternating ciliary bundle orientation pattern. In contrast, the lagena exhibits more interspecific diversity in shape and ciliary bundle orientation, suggesting that it has special adaptations in these species. The macula neglecta, a sensory epithelium of unknown function, is present in all four species.     

Anatomical partitioning of three multiarticular human muscles.

To examine neuromuscular partitioning within human muscles, the innervation patterns and muscle fiber architecture of the flexor carpi radialis (FCR), extensor carpi radialis longus (ECRL) and lateral gastrocnemius (LG) muscles were examined. Consistent patterns of innervation between specimens were found within each of the three muscles. The nerve to the FCR clearly innervates three major architectural divisions of the muscle. The ECRL is innervated by two different muscle nerves. Branches of these nerves innervate at least two distinct anatomical subvolumes. However, the subvolumes of the ECRL defined by muscle architecture are not totally congruent with those defined by the innervation pattern. In the LG, the single muscle nerve branches into two main divisions, and these subsequently divide into branches which supply the three heads. However, each head does not receive a completely private nerve. These results indicate that human muscles are partitioned in a manner roughly similar to the divisions of the same muscles in cats and rats, but with less congruency of architecture and innervation.     

Comparative light and electron microscopic study of the auditory organs of two species of fishes (pisces): Hyphessobrycon simulans (Ostariophysi) and Poecilia reticulata (Acanthopterygii).

Hyphessobrycon simulans has a Weberian apparatus for transmission of sound energy to the auditory organ, whereas Poecilia reticulata does not. The fine structure of the auditory organs is identical in the two species. The better hearing - expressed by large bandwidth and high sensitivity - typical of the Ostariophysi - seems to be based exclusively on the presence of the Weberian apparatus. The sensory epithelium of the saccule and the lagena is made up of hair (sensory) cells and supporting cells. The vertically orientated macula sacculi is divided into a dorsal and a ventral cell area with oppositely arranged hair-cell kinocilia. The sagitta takes up the center of the saccule and shows only three small sites with connections to the otolithic membrane. Remarkably, the dorsal sensory cells are connected to the ventral part of the otolith, but the ventral cells are connected to the dorsal part. The macula of the lagena also comprises a dorsal and a ventral cell area with oppositely arranged hair cells. The sensory cells in all maculae are of type II. They exhibit a striking apical cell protrusion, the cuticular villus. It is partially fused with the kinocilium in the contact zones and joined to the otolithic membrane. The cuticular villus probably stabilizes the long kinocilia.     

The ears of butterflyfishes (Chaetodontidae): ‘hearing generalists' on noisy coral reefs?

Analysis of the morphology of all three otolithic organs (sacculus, lagena and utriculus), including macula shape, hair cell morphology, density, orientation pattern, otolith morphology and the spatial relationships of the swimbladder and ear, reveals that butterflyfishes in the genera Chaetodon (which has anterior swimbladder horns) and Forcipiger (which lacks anterior swimbladder horns) both demonstrate the ear morphology typical of teleosts that lack otophysic connections, fishes that have traditionally been considered to be 'hearing generalists'.     

Structure of the nervous system of Myzostoma cirriferum (Annelida) as revealed by immunohistochemistry and cLSM analyses

The nervous systems of juvenile and adult Myzostoma cirriferum Leuckart, 1836, were stained with antisera against 5-HT (5-hydroxytryptamine, serotonin), FMRFamide, and acetylated alpha-tubulin in combination with the indirect fluorescence technique and analyzed by confocal laser scanning microscopy. The central nervous system consists of two small cerebral ganglia, connected by a dorsal commissure, a ventral nerve mass, and a pair of long circumesophageal connectives joining the former to the latter. The two neuropil cords within the ventral nerve mass curve outward and are joined to one another anteriorly and posteriorly. They are connected by 12 commissures, forming a ladder-like system. A single median nerve runs along the midventral axis. In addition to the circumesophageal connectives, 11 peripheral nerves arise from each main cord. The first innervates the anterior body region. The others form five groups of two nerves each, the first and thicker nerve of which is the parapodial nerve, innervating the parapodium and two corresponding cirri. Except for those in the most posterior group, the second nerves innervate the lateral organs and the body periphery. Serotonergic perikarya are arranged in six more or less distinct clusters, the first lying in front of and the other five between the main nerve cords. The distribution pattern of the FMRFamidergic perikarya is less clear and the somata lie between and outside the cords. One pair of dorsolateral longitudinal nerves was visualized by tubulin staining. Peripheral nerves and the commissures, in particular, demonstrate a segmental organization of the nervous system of M. cirriferum. Furthermore, their arrangement indicates that the body consists of six segments, the first of which is identifiable only by the first pair of peripheral nerves, the first two commissures, and the anteriormost ventral ganglion. The nervous system M. cirriferum thus exhibits several structures also found in the basic plan of the polychaete nervous system.     

The eye muscles and their innervation inChaetodon trifasciatus (Pisces,Teleostei, Chaetodontidae)

Synopsis InChaetodon trifasciatus, the large eye has the form of a thick disk rather than that of a globe. A deep cutaneous groove surrounds the eyeball, probably allowing rapid eye movements. The form and innervation of the three pairs of extraocular muscles are described. Each muscle is made of two types of fascicles of fibres, thick and thin. There is neither an anterior nor posterior myodome. The skull attachment of the obliques and of the inferior rectus is made on the thin sagittal ethmoidal membranous septum while that of the other recti occurs on osseous pieces of the skull. The attachment on the eyeball is made on the cartilaginous sclera. The ratio of the lengths of the antagonist muscles, superior vs. inferior oblique, superior vs. inferior rectus and medial vs. lateral rectus, is about 1.43:1. The three oculomotor nerves (III: common oculomotor, IV: trochlear and VI: abducens) as well as the ciliary system are described. For the following reasons, an analogy between the lateral rectus ofChaetodon trifasciatus and the lateral rectus + retractor bulbi of other vertebrates is indicated: (1) the nucleus of nerve III (which innervates four muscles) has four sectors, while that of IV (which innervates only the superior oblique) is made of one sector; (2) nerve VI consists of two roots corresponding to two groups of nerve cells of its motor nucleus and (3) in other vertebrates, nerve VI innervates both the lateral rectus and the retractor bulbi.     

The stomatogastric nervous system of the house cricket Acheta domesticus L. I. The anatomy of the system and the innervation of the gut

The distribution of the ganglia and nerves of the stomatogastric nervous system and the innervation of the extrinsic and intrinsic muscles are described. Median unpaired frontal and hypocerebral ganglia and paired ingluvial ganglia are present. The anterior pharynx is innervated by branches of the frontal nerve and by the anterior and posterior pharyngeal nerves, originating from the frontal ganglion. The posterior pharyngeal nerves are linked to nerves innervating the posterior part of the pharynx which have their origin in the hypocerebral ganglion, the anterior portion of which has previously been regarded as part of the recurrent nerve. Paired esophageal nerves run the length of the esophagus and crop between the hypocerebral and and ingluvial ganglia, innervating the muscularis by serial side branches. From each ingluvial ganglion runs an ingluvial nerve which innervates the gizzard and a cecal nerve which innervates the midgut and its ceca. At the posterior end of the midgut there is a poorly developed nerve ring. Nerves running posteriorly from this nerve ring link the stomatogastric nervous system with the proctodeal innervation from the terminal abdominal ganglion. Multipolar peripheral neurons are present on the muscularis of the whole of the foregut, rather randomly distributed on the crop and gizzard but forming fairly definite groupings at some points on the pharynx. Though of varied appearance, these cells could not be divided into discrete morphological categories. Peripheral neurons on the midgut are of different and characteristic morphology, though a few cells of the same appearance as those of the foregut occur at the midgut-hindgut boundary. Nerve fibers on the gut almost invariably terminate on the fibers of the muscularis.     

Insect epidermis: polarity patterns after grafting result from divergent cell adhesions between host and graft tissue

Insect epidermal cells display planar polarity (i.e. polarity in the plane of the cell sheet) by secreting oriented cuticular denticles and bristles before each moult. We investigate how cell polarities in an abdominal segment are uniformly oriented towards the posterior of the animal. Recently we have shown for the cotton bug Dysdercus that, in 180 degrees-rotated grafts pretreated with colchicine, graft cells tend to adopt the orientation prevailing in surrounding host cells via an intermediate stage with outward oriented denticles (Nübler-Jung and Grau, 1987). Here we show that, in untreated grafts that were transposed along the anteroposterior segment axis, the denticles also always tend to point outwards. This independence of the polarity pattern from the direction of transposition is compatible neither with a gradient model for polarity control, nor with the assumption that epidermal cells orient according to the local sequence of distinctly differentiated cells. Instead we found that outward orientation of graft denticles correlates with an elongation of epidermal cells along a host-graft border with divergent cell adhesiveness. We therefore propose that outward orientation in a graft results from a combination of two factors: epidermal cells stretch along an interface with divergent cell adhesiveness, and they form a denticle perpendicular to their long axis. By analogy, the normal anteroposterior orientation of denticles in a segment may result because epidermal cells tend to elongate parallel to the segment boundary and to form denticles perpendicular to this mediolateral cell elongation, i.e. along the anteroposterior segment axis.     

Acoustic response properties of lagenar nerve fibers in the sleeper goby,<Emphasis Type="Italic"> Dormitator latifrons</Emphasis>

Auditory and vestibular functions of otolithic organs vary among vertebrate taxa. The saccule has been considered a major hearing organ in many fishes. However, little is known about the auditory role of the lagena in fishes. In this study we analyzed directional and frequency responses from single lagenar fibers of Dormitator latifrons to linear accelerations that simulate underwater acoustic particle motion. Characteristic frequencies of the lagenar fibers fell into two groups: 50 Hz and 80–125 Hz. We observed various temporal response patterns: strong phase-locking, double phase-locking, phase-locked bursting, and non-phase-locked bursting. Some bursting responses have not been previously observed in vertebrate otolithic nerve fibers. Lagenar fibers could respond to accelerations as small as 1.1 mm s^–2. Like saccular fibers, lagenar fibers were directionally responsive and decreased directional selectivity with stimulus level. Best response axes of the lagenar fibers clustered around the lagenar longitudinal axis in the horizontal plane, but distributed in a diversity of axes in the mid-sagittal plane, which generally reflect morphological polarizations of hair cells in the lagena. We conclude that the lagena of D. latifrons plays a role in sound localization in elevation, particularly at high stimulus intensities where responses of most saccular fibers are saturated.Abbreviations BRA best response axis/axes - BS best sensitivity - CF characteristic frequency - CV coefficient of variation - DI directionality index - ISIH inter-spike interval histogram - PSTH peri-stimulus time histogram - SR spontaneous rate