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Brainstem reticular nuclei that project to the cerebellum in rats: a retrograde tracer study.

The nuclear origins of projections from the brainstem reticular formation to the cerebellum were examined using four retrograde tracer substances: horseradish peroxidase, wheat germ agglutinin-horseradish peroxidase conjugate, Fluoro-Gold, and rhodamine beads. Tracer injections were made into each of the three major longitudinal zones of the cerebellar cortex (vermis, paravermal hemisphere, and lateral hemisphere) as well as into the various deep cerebellar nuclei. Counts of retrogradely labeled cells were done on a large sample of select cases. The data generated by these cell counts indicate that the strongest reticulocerebellar projections arise from the three specialized pre-cerebellar reticular nuclei: the lateral reticular nucleus, the medullary paramedian reticular nucleus, and the reticulotegmental nucleus. The presumed noradrenergic locus coeruleus (A6 cell group) was also densely packed with retrogradely labeled neurons. However, strong reticulocerebellar projections also arose from other presumed catecholamine cell groups such as those in the ventrolateral medulla (the A1/C1 complex) and the caudal pons (A5). Substantial cerebellar projections originated from most of the various presumed serotonergic brainstem raphe cell groups (particularly raphe obscurus in the medulla), as well as from the presumed cholinergic Ch5 cell group (the pedunculopontine pars compactus nucleus). Labeled cells were also seen in several nonaminergic isodendritic reticular nuclei thought to be involved in visuomotor activity (e.g. paragigantocellularis dorsalis, raphe interpositus, and the pontine dorsomedial tegmental area), as well as in the lateral reticular zone of the medulla and lower pons (reticularis dorsalis and parvocellularis). Tracer injections into the deep nuclei produced relatively greater numbers of labeled neurons in large-celled medial reticular nuclei associated with skeletomotor activity, such as gigantocellularis, magnocellularis, and pontis caudalis. Reticular nuclei conspicuous in their lack of projections to the cerebellum included reticularis ventralis in the medulla, pontis oralis, and both subdivisions of the midbrain reticular formation (cuneiformis and subcuneiformis). As a whole, the various isodendritic reticular nuclei project most strongly to midline cerebellar structures (vermal cortex or fastigial nuclei), less strongly to the paravermal cortex or interposed nuclei, and least strongly to the lateral cortex or dentate nucleus. Within individual reticular nuclei, the morphology of labeled neurons is identical to that reported previously by this laboratory subsequent to spinal or cortical HRP injections, thus strengthening this laboratory's hypothesis that the various brainstem reticular nuclei can be distinguished on the basis of neuronal morphology.

Animals↗

The role of the cortex in play fighting by rats: developmental and evolutionary implications.

Play is a distinctive behavior of young mammals, especially mammals with a well-developed forebrain. For this reason it is thought that there may be a relation between forebrain evolution and highly elaborated play behavior. This study investigated the contribution of the cortex to play behavior by comparing play in control and neonatally decorticated rats (Rattus norvegicus). Play fighting in rats involves the combination of attack by one rat and defense by the recipient, with pinning arising when specific patterns of defense are used. Whether paired with another decorticate or with an intact pairmate, decorticates attacked pairmates as frequently as did intacts, and they were just as likely to defend against playful attacks as were intacts. Where decorticates differed from intacts was on a measure of pinning, in which one rat stands over a supine partner, decorticate rats displayed a reduction of 50% relative to control rats during the juvenile stage in which play is most pronounced (days 25 to 40). Juvenile decorticate rats adopted types of defensive responses which were less likely to result in the pinning configuration. Thus, a reduced pinning frequency reflects an altered pattern of defense, not a reduced level of play fighting. Rather, the decorticate patterns of defense were typical of those defensive responses displayed by adult rats. That is, decorticate juveniles exhibit a precociously mature pattern of playful defense. As intact controls mature, they come to resemble the decorticates in their defensive responses, and hence the difference in pinning frequency between decorticate and intact pairs diminishes. This suggests that the cortex may inhibit the escalation of defense in juveniles and thus promote prolonged ventral-ventral contact during play fighting. The results further suggest that the cortex is involved in the development of adult behavioral skills by facilitating juvenile play.

Aggression↗

Partition of function in the morphological subdivisions of the lateral geniculate nucleus of the tammar wallaby (Macropus eugenii).

Extra-cellular recordings from single cells in the dorsal lateral geniculate nucleus (dLGN) of the tammar wallaby, Macropus eugenii, were made to find out whether the stratification of the nucleus could be correlated with the receptive field properties of units. Retinofugal fibres terminate in the lateral geniculate nucleus of the wallaby in nine interleaved eye-specific layers. These may be grouped into a lateral alpha segment of six laminae and a medial beta segment of three, in which the cells are less densely packed. Ninety percent of the geniculate neurons recorded from in the alpha segment gave brisk responses to stimulation of their receptive fields. Cells with sluggish responses predominated in the beta segment, but there was also a sizable minority of cells with brisk responses that were indistinguishable from those recorded in the alpha segment. In contrast, other response properties were rarely differentiated in individual layers. Thus, in most layers, the numbers of cells with transient or sustained responses were not significantly different, and this was also true for cells with ON- or OFF- responses. For each of these response pairings, however, the numbers of one type (ON- and transient) predominated in every layer. The accumulation of this laminar distinction lead to significant differences in the alpha and beta segments and in the nucleus as a whole. We conclude that cells in the individual layers of the dLGN of the tammar wallaby show no evidence of having receptive field properties in common that might correlate with separate functional streams. There is a functional segregation of receptive field properties between the alpha and beta segments. The organization of these two segments resembles that of the A and the C layers of the dLGN in cats and, possibly, the magnocellular and koniocellular components of the dLGN in primates. These broad similarities in functional partition of the dLGN of different species suggests that this aspect of the organization of the nucleus is independent of lifestyle.

Animals↗

Morphology, distribution and innervation of the lateral-line receptors of the Florida gar, Lepisosteus platyrhincus.

The lateral-line system of gars consists of superficial neuromasts, which are arranged in lines termed pit lines, canal neuromasts and spiracular organs, which are located within diverticula of the hyoid gill pouch. Both canal and superficial neuromasts possess polarized hair cells whose directional sensitivity parallels the long axis of their respective lines. However, the apical surfaces of canal neuromasts are larger and possess far more hair cells than do those of superficial neuromasts, but superficial neuromasts have longer kinocilia and, presumably, longer cupulae. The receptors of the lateral-line system are innervated by three pairs of cranial nerves: anterior, middle and posterior lateral-line nerves. The anterior lateral-line nerves innervate neuromasts of the supraorbital, infraorbital and preoperculo-mandibular canals as well as dorsally located anterior pit lines, cheek (horizontal, vertical and mandibular) and gular pit lines of superficial neuromasts and the spiracular organ. The middle lateral-line nerves innervate dorsally located middle pit lines and a single neuromast in each temporal canal. The posterior lateral-line nerves innervate dorsally located posterior pit lines, neuromasts of the supratemporal commissures and all remaining postotic and trunk neuromasts. The ganglion of the anterior lateral-line nerve is divided into dorsal and ventral subganglia; the single ganglion of the middle lateral line nerve has no recognizable subdivisions, and the ganglion of the posterior lateral-line nerve consists of rostral and caudal subganglia. Analysis of the roots of these nerves and review of the embryonic origin of their ganglia as well as comparisons with cranial nerves in other anamniotes suggest that the anterior and posterior lateral-line nerves of gars may represent the fusion of four to five separate lateral-line nerves at some stage in vertebrate phylogeny. Thus, with the addition of the middle lateral-line nerve, and the possible existence of a ventral lateral-line nerve of the trunk, it is possible that the earliest jawed vertebrates possessed six or even seven pairs of lateral-line nerves.

Afferent Pathways↗

The primary projections of the lateral-line nerves of the Florida gar, Lepisosteus platyrhincus.

Gars, like most other ray-finned fishes, possess lines of superficial and canal neuromasts that form highly ordered spatial arrays on the head and trunk. These neuromasts are innervated by three pairs of cranial nerves: anterior, middle and posterior lateral-line nerves. Application of horseradish peroxidase to the roots of these nerves indicates that the afferent fibers of each nerve terminate throughout the rostrocaudal extent of the ipsilateral octavolateralis column, composed of medial and caudal octavolateralis nuclei, as well as rostrally in the eminentia granularis of the cerebellum. The afferents of each nerve terminate within different regions of these nuclei, thus preserving a rostrocaudal somatotopy of the peripheral receptors. The individual rami of these nerves innervate peripheral receptors in a dorsoventral sequence; however, application of horseradish peroxidase to these rami indicates an almost total overlap of the terminal fields of adjacent rami and, thus, little or no preservation of dorsoventral somatotopy of the peripheral receptors. Similarly, the terminal fields of fibers that innervate canal and superficial neuromasts overlap extensively, and there is no evidence for separate maps of canal and superficial neuromasts. Furthermore, the position of the terminal fields of superficial neuromasts is not consistent with a hypothesis that these receptors may have evolved into electroreceptors in some teleosts. However, these experiments do indicate that superficial neuromasts, like canal neuromasts, are innervated by efferent fibers of rostral and caudal efferent nuclei, but there is no evidence of a diencephalic efferent nucleus as occurs in teleosts.

Afferent Pathways↗

Electrocorticograms of hippocampal and dorsal cortex of two reptiles: comparison with possible mammalian homologs.

To compare the ongoing electrical activity in possibly homologous structures of reptiles and mammals, the electrographic activity (micro-EEG) from major parts of the cortex of unanesthetized turtles (Pseudemys) and geckos (Gekko) was recorded with and without acute and chronic stimuli, physostigmine and atropine. Electrodes were placed in the medial cortex (MC) and in the dorsal cortex (DC), the possible homologs of the mammalian hippocampus and transitional or/and isocortex, respectively. The resting corticograms (1-50 Hz) are different in the two cortical areas. Both are wide-band; power falls steadily with frequency above a single maximum about 2 Hz. The MC has a nonrhythmic, low-voltage activity with occasional superimposed large sharp waves (LSWs), generally biphasic, 100-300 microV and lasting 0.25-0.75 s. The DC has smaller amplitudes (ca. 3-6 dB) at all frequencies and fewer LSWs. Reptilian LSWs are reminiscent of mammalian hippocampal sharp waves or spikes, a correlate of decreased arousal. The immobility-related rhythmic slow activity (theta), so characteristic of the hippocampus in a number of mammals, was not found in the cortex of either species of reptile under a variety of conditions. We cannot exclude the possibility of movement-related theta waves. Physostigmine injection does not produce theta, although it acts like an arousing stimulus, producing a disappearance of the LSWs and a substantial increase in the amplitude of the frequencies 12-24 Hz; these changes were more obvious in the DC. Atropine reversed the effects of physostigmine. Theta may represent a trait of the more highly differentiated hippocampal field of mammals. The condition represented by these reptiles, in which the EEG differs between parts of the pallium but without theta or reciprocal changes in the MC and DC, may be an earlier evolutionary stage. A distinctive reptilian EEG is not recognizable in Pseudemys and Gekko, but a number of differences from the EEG in familiar mammals are shared by these two neurologically quite different reptiles.

Animals↗

Comparative cytoarchitectonic analysis of some visual pretectal nuclei in teleosts.

The posterior pretectal nucleus, which in Osteoglossum receives second order visual input and projects to the inferior lobe of the hypothalamus, was identified and characterized in species from all major groups of non-neoteleost teleosts. The hypothesis that the posterior pretectal nucleus in these species is homologous to both the pars intermedius of the superficial pretectal nucleus and nucleus glomerulosus in acanthopterygians is supported by multiple similarities in relative position and cytoarchitecture. Nucleus corticalis, which receives retinal input and projects to the posterior pretectal nucleus (or to nucleus glomerulosus), was identified in species belonging to three of the four major teleost radiations. Both the posterior pretectal nucleus and nucleus corticalis are plesiomorphic for teleosts. The presence of glomeruli in the posterior pretectal nucleus and nucleus glomerulosus in esocids and acanthopterygians, respectively, and the presence of two nuclei, the pars intermedius and nucleus glomerulosus, in acanthopterygians, as opposed to one nucleus, the posterior pretectal nucleus, are apomorphies.

Animals↗

Evolution of homologous vocal control traits.

Evolutionary neurobiologists want to know how neuronal properties (or traits) have been modified to subserve adaptive changes in behavioral phenotypes. Homology can provide a conceptual framework to distinguish the separate contributions of phylogenetic factors and current adaptive modifications to extant traits and behaviors. In this essay, a suite of nine vocal/sonic motor traits are compared in two orders of teleost fishes, the Batrachoidiformes and Scorpaeniformes. Only three of the traits are modified among Scorpaeniformes, the more advanced group. The large number of conserved characters among the study species suggests their sonic motor systems are homologs. This conclusion is consistent with the known phylogeny of teleosts and further implies that homologous sonic motor traits are more extensively modified among more recently evolved members (in this case the Scorpaeniformes) of the teleostean lineage. Since homology implies a common ontogenetic history for any trait, modifications thereof can potentially be linked to changes in identifiable developmental events, which themselves are homologs. Several hypotheses are proposed to account for the origins of modified sonic traits. The further demonstration that modified traits of the sonic motor system are in fact adaptations sets the stage for behavioral ecological studies that attempt to understand why the modified traits underlie behavioral changes that increase an individual's fitness.

Animal Communication↗

Evaluation of species-specific biochemical variation as a means for assessing homology in neuronal populations.

The phylogenetic history of neurons, as derived from the establishment of homologies, has been thought to be useful for furthering the understanding of nervous system function and behavior. The sensitivity of current biochemical and molecular techniques has been heralded as the ultimate means to establish neuronal phenotype and, thus in, turn to assess neuronal homology among species. Studies delineated in this review define the caveats associated with over reliance on such an approach, even when an in-depth understanding of the biochemical and genetic makeup of homologous neuronal populations is available.

Animals↗

The molecular evolution of neuropeptides: prospects for the '90s.

Three distinct opioid precursors have been identified in the central nervous system of mammals: proopiomelanocortin (POMC), proenkephalin, and prodynorphin. These precursors are derived from separate genes, synthesized in distinct neurons, and yield unique sets of opioid end products. This review will discuss the general mechanisms involved in the biosynthesis of neuropeptide precursors and consider the roles of posttranscriptional and posttranslational processing mechanisms in the generation of multiple sets of end products from a single gene. In addition, techniques that can be used for isolating and characterizing neuropeptide genes, mRNAs, and end products will be reviewed. These introductory comments will serve as the framework for a discussion of the phylogeny of the opioid precursors in the major groups of non-mammalian vertebrates.

Animals↗

Ontogeny and phylogeny: a re-evaluation of conceptual relationships and some applications.

The history of biology reveals numerous interpretations of the relationship between ontogeny and phylogeny, among them Garstang's critical realization that phylogeny can only be the history of changes in an ancestral ontogeny. One extrapolation of Garstang's conclusion is that the study of phylogeny should involve comparisons of ontogenies, in order to allow a reconstruction of the sequence of changes that led to the present variation in these ontogenies as well as the mechanisms underlying these changes. It is suggested that this can be done by recognizing homologous developmental stages among various ontogenies and subjecting these stages to a cladistic analysis. Some of the problems involving such analysis are explored in the course of examining the phylogeny of pharyngeal pouch ontogenies as a model system. It is concluded that sequences of ontogenetic stages are conserved (von Baerian recapitulation) but that terminal (Haeckelian recapitulation) and nonterminal alterations in the ancestral ontogeny are frequent. Both terminal and nonterminal alterations occur with approximately equal frequency in the case of pharyngeal pouches, but non-terminal deletion is rare to nonexistent. Examination of two additional morphological cases in which ontogeny has been utilized to infer phylogenetic changes-ontogenetic parcellation of neural systems and craniate head organization-reveals a number of problems from the point of view of a cladistical analysis of ontogenesis. Ontogenetic parcellation is essentially a recasting of Haeckel's biogenetic law and is rejected on the grounds that ontogenies do not parallel phylogenies, and only primitive and derived characters, not organisms, can be recognized. It is argued that the phylogenetic significance of transient neural characters can be determined only within a cladistical context and that insufficient ontogenetic data presently exist to evaluate most characters. Finally, the history of current models of craniate head organization is traced, and this information, along with that from more recent developmental studies, necessitates rejection of the Goodrich and Jarvik-Bjerring models. A new model is presented that purports to incorporate current ideas of somitomere and neuromere distribution, but it is deemed inadequate because, like previous models, it is based on information gleaned primarily from the ontogeny of a single species, whereas the model sought is a model of the ontogeny of ancestral craniates. To arrive at such a model, we must generate a morphotype, based on shared primitive features of cephalic ontogenies of all craniate radiations, which involves a cladistic analysis of these ontogenies.

Animals↗

Origins of descending spinal pathways in prehensile birds: do parrots have a homologue to the corticospinal tract of mammals?

In mammals, the supraspinal descending projections that influence distal limb muscles are the rubrospinal and corticospinal tracts. The former, which is found in other vertebrates, shows greater somatotopy in mammals that are 'dextrous' (e.g. monkeys) than those that are not (e.g. opossums). Similarly, the corticospinal tract, which is found only in mammals, has more extensive connections (i.e. direct corticomotoneural) in mammals that are dextrous than in mammals that are not. Descending spinal pathways have been described in 'non-dextrous' avian species (chickens, ducks, geese and pigeons), and the purpose of this study was to determine if there are any differences in the origins of descending projections to the spinal cord in 'dextrous' or prehensile parrots (sulphur-crested cockatoo, Cacatua galerita, and eastern rosella, Platycerus eximius). True Blue or wheat germ agglutinin-horseradish peroxidase was injected into the lumbar or cervical spinal cord. The distribution of retrogradely labelled cells was similar to that previously reported for non-prehensile birds. We found no evidence of any direct spinal projections from the telencephalon (including any pathway homologous to the corticospinal tract of mammals), nor any specialized anatomical organization of the descending pathways that could account for the pedal dexterity of these species.

Animals↗

Auditory cortex of the long-eared hedgehog (Hemiechinus auritus). I. Boundaries and frequency representation.

The boundaries of the primary auditory cortex of the long-eared hedgehog, Hemiechinus auritus, were determined by single-cell recordings, myeloarchitecture and retrograde horseradish peroxidase labeling in the medial geniculate, using anesthetized animals. The auditory cortex is located on the lateral surface of the temporal cortex, medial to the rhinal fissure. Responses to pure tones revealed an orderly representation of best frequencies in the primary auditory cortex, with low frequencies represented rostrally and high frequencies caudally. A second auditory field caudal to the primary one was indicated.

Animals↗

Releasing (unclasping) in male American toads: a neural substrate in the lateral subtoral tegmentum.

Release signals (chirps and side vibrations) are given by a male toad being clasped by another toad. These signals trigger immediate releasing (i.e. unclasping) by the clasping toad. First, small electrolytic lesions in the lateral subtoral tegmentum of male American toads abolished releasing, i.e. these toads persisted in clasping release-signaling males. Second, stimulation of this area caused bilateral abduction of the forelimbs, as occurs in normal releasing. Third, neural responses, resulting from tactile stimulation of either thumbpad, could be recorded from this region. Supposedly, similar stimuli would be generated by the release vibrations of the sides given by a male being clasped. These data are consistent with the participation of the lateral subtoral tegmentum in the detection of release signals and in the triggering of releasing. Release signaling was probably an important step in the evolution of anuran vocalizations. The midbrain tegmentum may contain primitive circuits for the detection of stimuli eliciting anuran acoustic behaviors.

Animals↗

The diencephalon of the channel catfish, Ictalurus punctatus. II. Retinal, tectal, cerebellar and telencephalic connections.

In a companion paper, the nuclear organization of the diencephalon of the channel catfish, Ictalurus punctatus, was described and compared to that of other teleosts. The present paper describes the connections of the diencephalon with the retina, optic tectum, corpus of the cerebellum and telencephalon. The principal tracer employed is the indocarbocyanine dye DiI which diffuses along neuronal membranes in fixed tissues. Almost all of the nuclei that were recognized as distinct in the companion study are found to also exhibit distinct sets of connections. Most of these connections have not been described previously in catfishes or other teleosts. When combined with connectional data from the existing literature, the results of the present study allow one to recognize a great number of distinct pathways through the diencephalon of channel catfish, including several visual, auditory, gustatory, electrosensory and mechanosensory pathways to the telencephalon. Almost all of the species differences in diencephalic organization noted in the companion study can be accounted for by changes in one of the major sensory pathways. In contrast, the multimodal and integrative areas of the diencephalon appear to be relatively conservative. A comparison between the diencephalon of teleosts and that of other vertebrates suggests that the dorsal thalamus, the ventral thalamus and the posterior tuberculum are homologous, at least in part, to the dorsal thalamus, the zona incerta and the subthalamic nucleus of mammals, respectively. All three areas project to the telencephalon in both mammals and teleosts. In most vertebrates, however, the dorsal thalamus provides the dominant input to the telencephalon, whereas in teleosts the main telencephalic input derives instead from the posterior tuberculum.

Animals↗

Development and innervation of the paratympanic organ (Vitali organ) in chick embryos.

The paratympanic organ (Vitali organ) is a small sensory organ in the middle ear of birds. It possesses a sensory epithelium with hair cells similar to those of the inner ear. Injections of fluorescent carbocyanine tracers into the paratympanic organ of 9- to 11-day-old chick embryos labeled ganglion cells in the facial ganglia. Paratympanic nerve fibers enter the brainstem with the facial nerve but proceed to vestibular brainstem nuclei. A dorsal branch terminates in ventral areas of the cerebellum, while a ventral component projects to the descending vestibular nucleus, with some fibers turning medially into lateral parts of the medial vestibular nucleus. No fibers were labeled in the motor or sensory facial nucleus or in auditory brainstem nuclei. This projection pattern suggests a function of the paratympanic organ in equilibrium rather than audition. Projections similar to those of the paratympanic nerve have been reported for the lagenar nerve. Immunocytochemical techniques using an antiserum to gamma-aminobutyric acid (GABA) demonstrate that hair cells in the paratympanic organ develop GABA immunoreactivity at 5 days of incubation (E5), 2-4 days earlier than GABA immunoreactivity can be detected in hair cells of the inner ear, i.e. in the saccule (E6.5-7.0), the utricle (E7), the cristae (E8-9) and the cochlea (E9-9.5). Afferent fibers that are transiently GABAergic are rare in the paratympanic organ (1-2 fibers), though present from E6 to E7.5. The early onset of GABA immunoreactivity in the paratympanic organ may indicate that this organ matures (and possibly functions) earlier in ontogenetic development than its counterparts located in the inner ear. The present findings are consistent with the hypothesis that the paratympanic organ is homologous with the spiracular sense organ of fishes. The paratympanic organ of birds may represent a sense organ that is derived phylogenetically and ontogenetically from the lateral-line system.

Animals↗

Synodontid catfish: a new group of weakly electric fish. Behavior and anatomy.

Three species of synodontid catfish can produce weak biphasic electric discharges in either continuous or burst-like fashion. The peak-power frequency of these electric organ discharges is around 100 Hz. The electric organ appears to have originated from a sonic muscle: the organ lies dorsal to the swim bladder, contains modified muscle tissue, and is innervated from a motor nucleus located in the caudal medulla. It is possible that synodontid catfish use electric signals in communication and/or object detection.

Animals↗

Evolution of vertebrate motor systems for acoustic and electric communication: peripheral and central elements.

Among vertebrates, there are a number of different neuromuscular systems specialized for the production of acoustic and electric social communication signals. Each system involves distinct sets of striated muscles that are derived from paraxial mesodermal somites and are components of a peripheral effector or 'communication organ': the larynx and syrinx of tetrapods, the sonic swim bladder and electric organ of fishes. Each of these systems further exhibits species-typical characters ranging from the number of muscles involved to myofibrillar architecture. Given that communication uses striated muscle, its circuitry is seen to represent a modification of existing motor systems. Thus, it is not surprising that these systems share several features suggesting that common mechanical or developmental factors influence their evolution. Of prime importance in comparing central communication circuitry between fishes and tetrapods is that among tetrapods the sound-generating organs are a part of the ventilatory system and so their activity must be coordinated with other sets of neuromuscular units. Differences in the central pattern of circuitry are thus expected between fishes and tetrapods, as well as among tetrapods themselves, since ventilation mechanisms differ between taxa. Moreover, birds have independently evolved a vocal syrinx which is distinctly separate from their larynx.

Animal Communication↗