Lateral Paragigantocellular Reticular Nuclei Research Articles

Analysis of projections from the cochlear nucleus to the lateral paragigantocellular reticular nucleus in the rat.

Golgi-staining, retrograde and anterograde tract-tracing, and a two-color immunoperoxidase technique have been employed, at the light- and electron-microscopic levels, to analyze the auditory projections from the cochlear nucleus (CN) to the lateral paragigantocellular reticular nucleus (LPGi) in the rat. We have found that the auditory input originates predominantly in the posteroventral and cochlear root nuclei. The auditory axons terminate in the cell-poor, ventral portion of the LPGi, which is strongly invaded by beaded dendritic profiles, originating from parent cell bodies located in the dorsal half of the LPGi. Ultrastructural analysis has revealed that the anterogradely labeled auditory axons form functional synapses preferentially with dendritic shafts. These axo-dendritic contacts are apparently excitatory in nature. By means of a sequential two-color immunoperoxidase staining method, we have further characterized potential postsynaptic neurons in the LPGi. Black-stained auditory fibers intermingle with brown-stained serotonergic or adrenergic neurons. Varicose auditory axons are often closely apposed to immunoreactive dendritic profiles of serotonergic and adrenergic neurons, indicating the presence of possible synaptic contacts of auditory terminal fibers with these transmitter-classified cells. The monosynaptic auditory in- put from the CN may modulate the activity of B3 sero-tonergic and C1 adrenergic cells in the LPGi and may thus induce adaptive changes in response to acoustic stimuli.

Efferent connections from the external cuneate nucleus to the medulla oblongata in the gerbil

The present study revealed the efferent projections from the external cuneate nucleus (ECN) to various medullary nuclei in the gerbl as demonstrated in fresh living brainstem slices by using in vitro anterogradely tracing with the dextran-tetramethyl-rhodamine-biotin. The tracer-labelled ECN axon terminals were observed (1) in most of the vital autonomic-related nuclei: the nucleus solitary tractus, nucleus ambiguus, rostroventrolateral reticular nucleus and C 2 adrenergic area, (2) in the reticular formation: the medullary, parvocellular, intermediate, gigantocellular, dorsal paragigantocellular and lateral paragigantocellular reticular nuclei and medullary linear nucleus, and (3) in sensory nuclei: the cuneate nucleus, spinal trigeminal nuclei caudalis and interpolaris, paratrigeminal nucleus, medial and spinal vestibular nuclei, inferior olive and prepositus hypoglossal nucleus. These new findings are discussed in relation to possible roles of the ECN in cardiovascular, respiratory and sensorimotor controls.

Localization of the antinociceptive and antianalgesic effects of neurotensin within the rostral ventromedial medulla.

Triple microinjections of neurotensin (10 nmol each), which occupied a large volume of tissue within the nucleus reticularis gigantocellularis (Gi), produced an inhibition of the tail-flick reflex in awake rats. This effect was less than that previously observed by a single injection (10 nmol) into the nucleus raphe magnus (RMg) (see ref. [25]). Bilateral injections of neurotensin (10 nmol each) into the nucleus reticularis paragigantocellularis lateralis (LPGi) had no effect. The neurotensin antagonist [D-Trp11]-neurotensin (3 pmol) was previously found to enhance morphine, but not beta-endorphin antinociception from the periaqueductal gray (PAG) when injected into the RMg. A similar enhancement of morphine, but not beta-endorphin antinociception from the PAG was observed in the current study by [D-Trp11]-neurotensin injections into the bilateral LPGi, bilateral Gi, or medial Gi. These data suggest that neurotensinergic projections from the PAG function in an antianalgesic manner throughout the RVM during morphine, but not beta-endorphin antinociception. The antinociceptive effect of neurotensin, on the other hand, is more localized.

CNS cell groups projecting to pancreatic parasympathetic preganglionic neurons

The CNS cell groups that project to the pancreatic parasympathetic preganglionic neurons were identified by the viral retrograde transneuronal labeling method. Pseudorabies virus (PRV) was injected into the pancreas of C8 spinal rats and after 6 days survival, the animals were perfused and their brains processed for immunohistochemical detection of PRV. Parasympathetic preganglionic neurons of the dorsal vagal nucleus were retrogradely labeled with PRV. Several CNS cell groups consistently contained transneuronally labeled neurons. In the medulla oblongata, labeled neurons were found in the nucleus tractus solitarius, area postrema, paratrigeminal nucleus, lateral paragigantocellular reticular nucleus, raphe pallidus and obscurus nuclei, C3 region and scattered cells in the ventral medullary reticular formation. In the pons, the A5 cell group, Barrington's nucleus and the subcoeruleus region contained labeled neurons. Only an occasional labeled cell was identified in the parabrachial nucleus. In the midbrain, almost no labeling was found except for an occasional neuron in the central gray matter. In the diencephalon, labeling was found in the paraventricular hypothalamic nucleus (PVN) as well as in the lateral hypothalamic nucleus at two levels (one at the level of the PVN and the other at the level of the subthalamic nucleus). in addition, the perifornical and dorsal hypothalamic nuclei contained labeled neurons. A few cells were found in the peripheral part of the dorsomedial hypothalamic nucleus. No labeling was seen in the ventromedial hypothalamic nucleus. In the telencephalon, the central amygdaloid nucleus and the bed nucleus of the stria terminalis were labeled.

CNS innervation of airway-related parasympathetic preganglionic neurons: a transneuronal labeling study using pseudorabies virus

The CNS cell groups that innervate the tracheal parasympathetic preganglionic neurons were identified by the viral retrograde transneuronal labeling method. Pseudorabies virus (PRV) was injected into the tracheal wall of C8 spinal rats and after 4 days survival, brain tissue sections from these animals were processed for immunohistochemical detection of PRV. Retrogradely labeled parasympathetic preganglionic neurons were seen in three sites in the medulla: the compact portion of the nucleus ambiguus, the area ventral to the nucleus ambiguus, and the rostralmost portion of the medial nucleus tractus solitarius (NTS); this labeling pattern correlated well with the retrograde cell body labeling seen following cholera toxin ß-subunit injections in the tracheal wall. PRV transneuronally labeled neurons were found throughout the CNS with the most abundant labeling concentrated in the ventral medulla oblongata. Labeled neurons were identified along the ventral medullary surface, and in nearby areas including the parapyramidal, retrotrapezoid, gigantocellular and lateral paragigantocellular reticular nuclei as well as the caudal raphe nuclei (raphe pallidus, obscurus, and magnus). Serotonin (5-HT) neurons of the caudal raphe complex (B1–B3 cell groups) and ventromedial medulla were labeled as well as a few C1 adrenergic neurons. The A5 cell group was the major noradrenergic area labeled although a small number of locus coeruleus neurons were also labeled. Several NTS regions contained labeled cells including the commissural, intermediate, medial, central, ventral, and ventrolateral subnuclei. PRV infected neurons were present in the Kölliker-Fuse and Barrington's nuclei. In the rostral mesencephalon, the precommissural nucleus of the dorsal periventricular gray matter was labeled. Labeling was present in the dorsal, lateral and paraventricular hypothalamic nuclei. In summary, the airway parasympathetic preganglionic neurons are innervated predominantly by a network of lower brainstem neurons that lie in the same regions known to be involved in respiratory and cardiovascular regulation. These findings are discussed in relationship to some of the potential CNS mechanisms that may be operative in airway disorders as well as potentially involved in certain fatal respiratory conditions such as Ondine's curse and sudden infant death syndrome (SIDS).

Studies of brain structures involved in diffuse noxious inhibitory controls in the rat: the rostral ventromedial medulla.

1. Previous electrophysiological, pharmacological and anatomical evidence had suggested a possible participation of rostral ventromedial medulla (RVM) in the supraspinal part of the loop underlying diffuse noxious inhibitory controls (DNICs). In order to test this hypothesis, two experimental series were performed during which DNICs were compared in control sham-operated rats and rats with lesions of the RVM and of an adjacent candidate for such a role, the nucleus gigantocellularis reticularis (Gi). 2. In the first experimental series, lesions were induced, in anaesthetized animals, by injections of quinolinic acid (0.3-0.8 microliter of a 360 nmol/microliter solution) into the RVM or Gi. In the control animals (n = 10), the vehicle alone (artificial cerebrospinal fluid) was injected. Histological lesion reconstructions were performed after each electrophysiological experiment. Three groups of animals were considered: in the first group (n = 5), the lesion was centered on the RVM, including the two caudal thirds of nucleus raphe magnus (NRM) and adjacent reticular areas; in the second group (n = 5), the lesions extended more rostrally and involved the rostral pole of NRM; in the third group (n = 5), the lesion extended more laterally and dorsally and included nucleus reticularis gigantocellularis pars alpha (GiA), nucleus reticularis paragigantocellularis lateralis (LPGi) and the Gi. In each case, all the neuronal cell bodies within the lesioned area were destroyed. 3. In the second experimental series, electrolytic lesions of the total rostrocaudal extent of the NRM (n = 5) were induced, in anesthetized animals, by passing cathodal current (5 mA, 8 s). In the control animals (n = 5), the electrode was lowered but current was not applied. 4. One week after lesioning, the animals were anaesthetized, paralysed, artificially ventilated and recordings were made from convergent neurones in trigeminal nucleus caudalis. These neurones gave responses due to activation of A and C fibres when percutaneous electrical stimuli were applied to their receptive fields. DNICs were triggered by immersion of each paw in a 50 degrees C water-bath. Both the general properties of the convergent neurones and the inhibitions of the C fibre-evoked responses produced by these heterotopic noxious stimuli were compared in the different groups of animals. 5. The sizes of receptive field, spontaneous activities, thresholds for C fibre-evoked responses and responses to C fibre activation were not different in the control and lesioned animals. The percentage inhibitions of the C fibre-evoked responses both during and in the 44s following the conditioning periods were also very similar in the different groups of animals.(ABSTRACT TRUNCATED AT 400 WORDS)

The origins of supraspinal projections to the cervical and lumbar spinal cord at different stages of development in the gray short-tailed Brazilian opossum, Monodelphis domestica

We have used the retrograde transport of Fast blue (FB) to study the origins of supraspinal projections to the lumbar and cervical spinal cord at different stages of development in the Brazilian, short-tailed opossum, Monodelphis domestica. Monodelphis was chosen for study because its young are born in a very immature state, 14–15 days after copulation, making it possible to manipulate its nervous system in an embryonic state without intra-uterine surgery. When injections of FB were made into the lumbar cord at postnatal day (PD) 1, neurons were labeled within several areas of the reticular formation (the retroambiguus nucleus, the ventral and dorsal reticular nuclei of the medulla, the gigantocellular reticular nucleus, the lateral paragigantocellular reticular nucleus, and the pontine reticular nucleus), the presumptive coeruleus complex, and the lateral vestibular nucleus. In many cases, labeled neurons were also found within the caudal raphe and the presumptive interstitial nucleus of the medial longitudinal fasciculus. The results of immunocytochemical studies provided evidence for catecholaminergic and serotoninergic neurons in the brainstem at PD1 and for axons of both phenotypes in the spinal cord. By PD3, labeled neurons were found within the ventral gigantocellular and ventral pontine nuclei of the reticular formation, the spinal trigeminal nucleus, and the presumptive paraventricular nucleus of the hypothalamus. When injections were made at PD4, neurons were also labeled within the medial and inferior vestibular nuclei, the red nucleus, the mesencephalic nucleus of the trigeminal nerve, the presumptive nucleus of Edinger-Westphal and the lateral hypothalamus. By at least PD7, the pattern of supraspinal labeling was similar to that obtained at older ages and in the adult animal. When FB was injected into the cervical cord at PD1, neurons were labeled in all of the areas labeled by lumbar injections at the same age and in larger numbers. In addition, labeled neurons were found within the ventral gigantocellular and spinal trigeminal nuclei. When cervical injections were made at PD15, labeled neurons were found within the deep cerebellar nuclei and amygdala and by PD17 they were also present within the superior colliculus and cerebral cortex. In some cases, cortical labeling was present outside the areas labeled by comparable injections in adult animals. Our results indicate that: (1) restricted areas of the brainstem innervate the spinal cord at birth in Monodelphis and some of the neurons which contribute to that innervation are monoaminergic, (2) several areas of the brain do not project to the spinal cord until well after birth and (3) the development of descending spinal pathways occurs asynchronously and according to a predictable sequence.

GABAergic circuitry in the rostral ventral medulla of the rat and its relationship to descending antinociceptive controls

This study used postembedding immunocytochemistry to examine the organization of GABA-immunoreactive synapse in the rostral ventral medulla (RVM) of the rat. To determine whether the outflow neurons of the RVM are under GABAergic control, we examined the distribution of GABA-immunoreactive synapses upon bulbospinal projection neurons that were labelled by retrograde transport of wheatgerm agglutinin-HRP from the cervical spinal cord. To study the possible convergence of GABAergic and periaqueductal gray (PAG) synaptic inputs to RVM neurons, we also made lesions in the PAG and examined the relationship between degenerating PAG axons and GABA-immunoreactive terminals. Approximately 45% of all synapses in the RVM, which includes the midline nucleus raphe magnus and the nucleus reticularis paragigantocellularis lateralis, were GABA-immunoreactive. The vast majority of GABA-immunoreactive terminals contained round, clear, and pleomorphic vesicles and made symmetrical axodendritic synapses; axoaxonic synapses were not found. Almost 50% of the retrogradely labeled dendrites in the NRM were postsynaptic to GABA-immunoreactive terminals. Several examples of convergence of degenerating PAG terminals and GABAergic terminals onto the same unlabelled dendrite were also found. These data indicate that the projection neurons of the RVM are under profound GABAergic inhibitory control. The results are discussed with regard to the hypothesis that the analgesic action of narcotics and electrical stimulation of the midbrain PAG involves the regulation of tonic GABAergic inhibitory controls that are exerted upon spinally-projecting neurons of the nucleus raphe magnus.

Direct projection from the dorsal hypothalamic area to the nucleus raphe pallidus: a study using anterograde transport with Phaseolus vulgaris leucoagglutinin in the rat.

A hypothalamic projection to the nucleus raphe pallidus of the medulla was examined using the anterograde tracing technique based on Phaseolus vulgaris leucoagglutinin (PHA-L) in the rat. After the iontophoretic application of PHA-L to the dorsal hypothalamic area, labeled fibers that finally ended in the nucleus raphe pallidus were observed descending through the most medial part of the ventral tegmental area and the nucleus reticularis tegmenti pointis to reach the medial aspect of the pyramid. Many varicose fibers forming a loose plexus were observed in the nucleus raphe pallidus, especially ventrally. The ventral surface of the pyramid and the most ventral region of the nucleus reticularis paragigantocellularis lateralis (PGCL) contained labeled varicose fibers. At the electron microscopic level, the labeled profiles in the nucleus raphe pallidus were small-sized unmyelinated axons and axon terminals. Labeled axon terminals containing spherical synaptic vesicles formed synapses on spine-like protrusions or small-sized dendritic shafts. These results strongly indicate that neurons in the dorsal hypothalamic area have a direct connection with neurons in the nucleus raphe pallidus and the ventral part of the PGCL. The possible involvement of this pathway in cardiovascular regulation was discussed.

Brainstem projections to the major respiratory neuron populations in the medulla of the cat

Efferent and afferent connections of the dorsal and ventral respiratory groups in the medulla of the cat were mapped by axonal transport of wheat germ agglutinin conjugated to horseradish peroxidase. Injections of wheat germ agglutinin-horseradish peroxidase into the dorsal respiratory group and the three principal subdivisions of the ventral respiratory group (caudal, rostral, and Bötzinger Complex) revealed extensive interconnections between these regions and with a limited number of other brainstem neuron populations. Major neuron populations with efferent projections to the regions of the dorsal and ventral respiratory groups include the parabrachial nuclear complex (medial parabrachial, lateral parabrachial, and Kölliker-Fuse nuclei), subregions of the lateral paragigantocellular reticular nucleus, subregions of the lateral and magnocellular tegmental fields, inferior central and postpyramidal nuclei of the raphe, and sensory trigeminal nuclei. A previously unidentified neuron population with extensive efferent projections to the dorsal and ventral respiratory groups was found near the ventral surface of the rostral medulla; we refer to this group as the retrotrapezoid nucleus. The results suggest that the dorsal and ventral respiratory groups form an extensively interconnected neuronal system receiving convergent inputs from the same brainstem nuclear groups, consistent with the hypothesis that the dorsal and ventral groups are primarily sites for integration of sensory and premotor respiratory drive inputs. Neuron populations in the rostral ventrolateral medulla with projections to both the dorsal and ventral respiratory groups, particularly the retrotrapezoid nucleus and neighboring subregions of the lateral paragigantocellular reticular nucleus, are candidate sites for participation in respiratory rhythmogenesis or other critical functions of the brainstem respiratory control system such as intracranial chemoreception.

Bulbospinal thyrotropin-releasing hormone projections to the intermediolateral cell column: A double fluorescence immunohistochemical-retrograde tracing stdy in the rat

Whereas the neurochemistry of the peripheral autonomic nervous system has been well characterized, less is known concerning the neurotransmitters utilized by medullary projections onto sympathetic preganglionic neurons residing in the thoracolumbar spinal intermediolaleral cell column. Retrograde transport of rhodamine-labeled fluorescent microspheres following discrete microinjection into the thoracic intermediolateral cell column was combined with immunohistochemistry to determine neuroanatomic location of thyrotropin-releasing hormone-immunoreactive neurons which project to the intermediolateral cell column in the rat. The ventromedial group of raphe nuclei including the nucleus raphe pallidus. obscurus, and magnus possessed the greatest number of medullary thyrotropin-releasing hormone-immunoreactive neurons which also contained rhodamine-labeled microspheres. High numbers of intermediolateral cell column-projecting thyrotropin-releasing hormone-immunoreactive neurons were also observed in nucleus reticularis paragigantocellularis lateralis and magnocellularis, the lateral reticular nucleus, and the superficial ventral (arcuate) medullary surface. Despite the observations that nucleus reticularis gigantocellularis, paramedianus, and ventralis pars beta project to the intermediolateral cell column, double-labeled cells were not observed in these nuclei. Furthermore, whereas the nucleus reticularis magnocellularis and gigantocellularis, and the lateral reticular nucleus displayed strong ipsilateral predominance in projecting to the intermediolateral cell column, other medullary reticular and raphe nuclei displayed bilateral projections. The present findings support the hypothesis that thyrotropin-releasing hormone-containing neurons in the ventral medulla project to the intermediolateral cell column and may influence sympathetic preganglionic neurons.

The distribution of substance P-, enkephalin-and dynorphin-immunoreactive neurons in the medulla of the rat and their contribution to bulbospinal pathways

This study examined the medullary distribution of peptide-containing neurons at the origin of bulbospinal pathways in the rat. Antisera directed against substance P, methionine-enkephalin-arg-glγ-leu and dynorphin B were used on sections in which spinally projecting brainstem neurons had been identified by the retrograde transport of a protein-gold complex that was injected into the spinal cord. Both the relative numbers and distribution of the different peptide-immunoreactive spinally projecting neurons differed. Methionine-enkephalin-immunoreactive neurons were twice as numerous as the substance P-immunoreactive cells and seven times more numerous than the dynorphin B-positive neurons. The methionine-enkephalin cells were found in all medullary raphénuclei, and in the ventromedial and ventrolateral medullary reticular formation. Caudally, the methionine-enkephalin cells were concentrated laterally; more rostrally, they were located more medially. Three major loci of methionine-enkephalin-immunoreactive cells were found: (1) the nucleus reticularis paragigantocellularis lateralis, at levels caudal to the facial nucleus, (2) the B3 cell group (nucleus raphémagnus and the nucleus reticularis magnocellularis, pars alpha) and the most rostral part of the B1 and B2 cell groups (nuclei raphépallidus and obscurus), (3) a dense cluster of cells that flanks the dorsal surface of the dorsal accessory olive (referred to as the nucleus interfascicularis hypoglossi, pars dorsalis). Substance P-like cells were seen in all raphénuclei except for the most anterior portion of the B3 cell group. Substance P-immunoreactive cells were also seen in both the ventromedial (nuclei reticularis ventralis and magnocellularis) and ventrolateral medulla (nucleus reticularis paragigantocellularis lateralis). Finally there was a dense concentration of substance P neurons in the nucleus interfascicularis hypoglossi, pars ventralis. The distribution of dynorphin-immunoreactive neurons differed significantly from that of methionine-enkephalin and substance P. Dynorphin cells were almost exclusively found in the ventrolateral medulla (nucleus reticularis paragigantocellularis lateralis), at all Ievels between the lateral reticular nucleus and the caudal pole of the facial nucleus. The proportion of each of these peptidergic-immunoreactive cells at the origin of bulbospinal pathways differed considerably. Substance P spinally projecting neurons were more numerous than methionine-enkephalin spinally projecting neurons. The majority of the substance P projection neurons were found in the Bl and B2 cell groups, in the adjacent ventromedial medulla and in the nucleus interfascicularis hypoglossi, pars ventralis. A few substance P projection neurons were also found in the caudal reaches of the B3 cell group. Methionine-enkephalin projection neurons were located in the rostral B1/B2 cell groups, B3 cell group and in the nucleus interfascicularis hypoglossi, pars dorsalis. None were found in caudal B1/B2 cell groups. Methionine-enkephalin projecting neurons were thus located rostral to the majority of the double-labelled substance P cells. Only scattered projection neurons were found among the methionine-enkephalin- and substance P-immunoreactive cells of the nucleus reticularis paragigantocellularis lateralis. In contrast to the large numbers of double-labelled methionine-enkephalin and substance P cells, no spinally projecting dynorphin neurons were found. We suggest that there are differential methionine-enkephalin 5-hydroxytryptamine inhibitory and substance P/5 hydroxytryptamine excitatory influences that issue from different medullary areas and are, respectively, exerted upon dorsal horn nociceptive neurons, and visceral/somatic motoneurons and intermediolateral preganglionic neurons.

Mapping study of serotoninergic input to diencephalic-projecting dorsal column neurons in the rat.

Several lines of evidence indicate that the processing of somatosensory information in the dorsal column nuclei (DCN) is subject to descending controls. Anatomical experiments have demonstrated projections to the DCN from the sensorimotor cerebral cortex and the reticular formation. Physiological studies have shown that the activity of DCN neurons can be altered following stimulation of the cerebral cortex, reticular formation, periaqueductal gray, or raphe nuclei. Recent biochemical and electrophysiological evidence suggests a serotoninergic modulation of DCN neurons. The present study identifies serotonin-containing contacts on cells in the DCN that project to the thalamus in the rat. Retrograde labeling of brainstem neurons by horseradish peroxidase demonstrated projections to the DCN from the nucleus reticularis paragigantocellularis lateralis and from several raphe nuclei, including nuclei raphe obscurus (RO), pallidus (RP), and magnus (RM). Double labeling with horseradish peroxidase and antibody for serotonin indicated that the RO, RP and RM are likely to be the sources of the serotoninergic projections to the DCN. Thus, the role of the serotoninergic output from the raphe nuclei includes modulation of activity in the DCN.

Efferent vestibular neurons: a study employing retrograde tracer methods in the pigeon (Columba livia).

Efferent neurons innervating the vestibular labyrinth and cochlea of the pigeon have been identified by means of a variety of retrograde tracers: [3H]-adenosine (Ad), horseradish peroxidase (HRP), Evan's Blue (EB) and Bisbenzimide (Bb). Discrete injections into individual cristae ampullares of the semicircular canals, into the macula utriculi, or into several of these end organs resulted in similar patterns of neuronal labelling. Efferent vestibular neurons were always found within a small portion of the nucleus reticularis pontis caudalis (RP), ventrolateral to the abducens nucleus on both sides. No systemic difference in the locations of labelled cells was found following injection into different sensory epithelia. Cell counts following injections into individual cristae did not differ significantly from those following injections into all three cristae. The injections into all cristae in both labyrinths yielded cell counts that were much lower than twice the number of cells labelled by injections into the three cristae on one side only. When HRP was injected into the right lateral canal crista and Ad into the right posterior canal crista, a high proportion of neurons was labelled with both compounds (61% of the HRP-labelled cells and 67% of the Ad-labelled cells). Injections of EB into all three cristae on the right side and Bb into all three cristae on the left side produced a smaller percentage of doubly labelled cells (10% of the EB-labelled cells and 6% of the Bb-labelled cells). It is concluded, therefore, that there is a considerable degree of collateralization within one labyrinth. Fewer collaterals of efferent neurons are directed to both labyrinths. Since each semicircular canal represents head rotation in one direction and one plane, it is unlikely that efferents which contact several different movement sensors can provide sensory motor control that is specific for directions and planes of head movements. Control injections of these tracers into the cochlea yielded labelled cells in a different reticular structure, the nucleus reticularis paragigantocellularis lateralis (Pgc), on both sides, as well as in the RP. It is proposed that the Pgc cells represent cochlear efferents, while the RP neurons are related to the macula lagenae, an otolithic organ of balance in the pigeon.