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One of the earliest activities expressed within the developing central nervous system is a widely propagating wave-like activity, which we referred to as the depolarization wave. Despite considerable consensus concerning the global features of the activity, its physiological role is yet to be clarified. The depolarization wave is expressed during a specific period of functional synaptogenesis, and this developmental profile has led to the hypothesis that the wave plays some roles in synaptic network organization. In the present study, we tested this hypothesis by inhibiting the depolarization wave in ovo and examining its effects on the development of functional synapses in vagus nerve-related brainstem nuclei of the chick embryo. Chronic inhibition of the depolarization wave had no significant effect on the developmental time course, amplitude, and spatial distribution of monosynaptic excitatory postsynaptic potentials in the first-order nuclei of the vagal sensory pathway (the nucleus of the tractus solitarius (NTS) and the contralateral non-NTS region), but reduced polysynaptic responses in the higher-order nucleus (the parabrachial nucleus). These results suggest that the depolarization wave plays an important role in the initial process of functional synaptic expression in the brainstem, especially in the higher-order nucleus of the cranial sensory pathway., (Copyright © 2017 IBRO. Published by Elsevier Ltd. All rights reserved.)

The central issue in developmental neuroscience is when and how neural synaptic networks are established and become functional within the central nervous system (CNS). Investigations of the neural network organization have been hampered because conventional electrophysiological means have some technical limitations. In this study, the multiple-site optical recording technique with a voltage-sensitive dye was employed to survey the developmental organization of the vagal system in the mouse embryo. Stimulation of the vagus nerve in E11-E14 mouse embryos elicited optical responses in areas corresponding to the vagal sensory and motor nuclei. Postsynaptic responses in the first-order sensory nucleus, the nucleus of the tractus solitarius (NTS), were identified from E11, suggesting that sensory information becomes transferred to the brain at this stage. In addition to the NTS, optical responses were identified in the rostral and contralateral brainstem regions, which corresponded to second/higher order nuclei of the vagus nerve including the parabrachial nucleus (PBN). Postsynaptic responses in the second/higher-order nuclei were detected from E12, suggesting that polysynaptic networks were functional at this stage. We discuss the results of our optical mapping, comparing them with previous findings obtained in the chick and rat embryos, and suggest some fundamental principles in the functional organization of synaptic networks in the embryonic brain., (© 2016 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

The multiple-site optical recording technique with a voltage-sensitive dye, NK2761, was used to survey the functional organization of neuronal networks related to the vagus nerve (N.X) in the E16-stage rat brainstem. When we stimulated N.X, in addition to the responses in the vagal sensory nucleus (nucleus of the tractus solitarius (NTS)) on the stimulated side, other response areas were bilaterally detected. Characteristics of the optical signals in these areas suggested that they correspond to neural activity in the second/higher-ordered nucleus of the vagal pathway. The first area was located at the level of the pons. Based upon morphological information, we suggest that this area corresponds to the parabrachial nucleus (PBN), which receives inputs from the NTS. The second area was located between the NTS and the PBN. We suggest that this area is the A5 noradrenergic group. These results suggest that the N.X-related neural networks are established similarly to the adult pattern from an early developmental stage., (Copyright © 2012 Elsevier Ireland Ltd. All rights reserved.)

To regulate the autonomic function, the vagus nerve transfers various sensory information from peripheral organs, and appropriate motor reflexes are produced in the neural circuit. The functional development of the vagal pathway during the early phase of embryonic development has long been unclear. Optical recording with voltage-sensitive dyes has provided a new approach to the analysis of the functional development of the embryonic central nervous system. In this review, we present recent progress in optical studies on the vagal pathway in the embryonic chick and rat brainstems. The topics include how neural excitability is initially expressed in the motor and sensory nuclei [e.g. the dorsal motor nucleus of the vagus nerve (DMNV) and the nucleus of the tractus solitarius (NTS)] and how synapse networks are formed in the primary and higher-ordered sensory nuclei [e.g. the parabrachial nucleus (PBN)]. We also refer to the functional development of the glossopharyngeal nuclei and compare the developmental steps with those of the vagal nuclei., (Copyright © 2011 Elsevier B.V. All rights reserved.)

Using voltage-sensitive dye recording, we traced the ontogenetic expression of neural excitability related to the glossopharyngeal nerve (N. IX) and the vagus nerve (N. X) in the embryonic chick brainstem. At the 3.5-day embryonic stage, by averaging optical signals, we succeeded in recording very small action potential-related optical responses (DeltaI/I<10(-4)) induced by N. IX stimulation in the nucleus of the glossopharyngeal nerve (motor nucleus). This suggests that glossopharyngeal excitability in the motor nucleus is first generated no later than this developmental stage. On the other hand, action potential-related optical responses induced by N. X stimulation were first detected at the 4-day embryonic stage. Comparison with morphogenesis indicated that glossopharyngeal and vagal motoneurons acquire their excitability and send their axons to the periphery soon after they leave the proliferative pool.

Multiple-site optical recording with a fast voltage-sensitive dye, absorption dye NK2761, was used to study the developmental organization of functional synaptic networks in the vagal pathway. Glutamatergic excitatory postsynaptic potentials (EPSPs) evoked by vagus nerve stimulation was first detected from the nucleus of the tractus solitarius (NTS) at embryonic day 7 (E7) in chick embryos and E15 in rat embryos, when morphological differentiation of pre- and postsynaptic neurons is incomplete. When extracellular Mg2+ was removed, small EPSPs were elicited at E6 in chick embryos and E14 in rat embryos. These results suggest that synaptic function mediated by N-methyl-D-aspartate (NMDA) receptors is latently generated 1 day before the expression of glutamatergic EPSP. Functional synapses related to the glossophyaryngeal nerve appear to be generated at the same time as the vagus nerve, but their spatial distribution was different from that of the vagus nerve. We further investigated the development of second synaptic pathways from the NTS to higher centers, and found that neuronal circuits from the NTS are already generated when the primary afferents form functional synapses with NTS neurons.

The multiple-site optical recording technique with a voltage-sensitive dye, NK2761, was used to survey functional organization of neural circuits related to the vagus nerve in the embryonic chick brainstem. When we stimulated the vagus nerve, in addition to the responses in the vagal sensory nucleus (nucleus of the tractus solitarius (NTS)) and motor nucleus (dorsal motor nucleus of the vagus nerve (DMNV)) on the stimulated side, another response area was discriminated at the level of the pons/rostral medulla on the contralateral side. Characteristics of the contralateral optical signals suggested that they correspond to the neural activity in the second/higher-ordered nucleus of the vagal pathway, possibly the parabrachial nucleus, which receives inputs from the NTS. Blockade of non-N-methyl-d-aspartate (NMDA) receptors abolished the responses on the contralateral side, together with the postsynaptic firing in the NTS, suggesting the significance of non-NMDA receptor function in sensory information transfer via the NTS. The responses on the contralateral side were first detected from the 7-day-old embryonic stage, when the glutamatergic excitatory postsynaptic potentials were first expressed in the NTS. The results suggest that the synaptic pathway from the NTS to the contralateral nucleus is already generated when the primary vagal afferents make functional synapses on NTS neurons.

We investigated the functional organization of the glossopharyngeal and vagal motor nuclei during embryogenesis using multiple-site optical recording with a fast voltage-sensitive dye. Intact brain stem preparations with glossopharyngeal and vagus nerves were dissected from 4- to 8-day-old chick embryos. Electrical responses evoked by glossopharyngeal/vagus nerve stimulation were optically recorded from many loci of the stained preparations. In 4- to 6-day-old preparations, action potential-related fast spikelike signals were detected from the nucleus of the glossopharyngeal nerve and the dorsal motor nucleus of the vagus nerve. Contour line maps of the signal amplitude showed multiple-peak patterns, suggesting that the neurons and/or their activity were not uniformly distributed within the nuclei at early developmental stages. As development proceeded from 4 to 6 days, the peaks fused with each other and the number of peaks decreased gradually. In most 7- and 8-day-old preparations, only a single peak was identified in the nuclei, and the distribution of the signal amplitude formed a layered pattern surrounding the peak-signal area. These results suggest that functional organization of the motor nuclei in the embryonic hindbrain changes dynamically with development, resulting in a rearrangement of functional nuclear cores from multiple-peaks to a single peak.

1. Multiple-site optical recording with the voltage-sensitive absorption dye NK2761 was used to investigate functiogenesis of the neural circuits in the embryonic central nervous system (CNS). In the present review, we emphasize the functional development of glossopharyngeal nerve (N. IX)- and vagus nerve (N. X)-related neural circuits in the embryonic chick brainstem. 2. In the nucleus tractus solitarius (NTS), glutamatergic excitatory post-synaptic potentials (EPSPs) evoked by stimulation of N. IX/N. X were first detected at Embryonic Day (E) 7, when morphological differentiation of pre- and post-synaptic neurons is incomplete. The removal of extracellular Mg2+ elicited small EPSPs at E6, suggesting that synaptic function mediated by N-methyl-d-aspartate (NMDA) receptors (glutamatergic receptors) is latently generated 1 day before the expression of glutamatergic EPSPs. In the nucleus of the glossopharyngeal nerve and the dorsal motor nucleus of the vagus nerve, action potentials were detected from E3.5 and E4, respectively. Similar chronological sequences were observed in the rat embryo and in other cranial nerve-related nuclei and were used to construct basic profiles of embryonic EPSPs in the vertebrate CNS. 3. We further investigated the development of the secondary synaptic pathways from the NTS to higher centres and found that neuronal circuits from the NTS are already generated when the primary afferents form functional synapses with NTS neurons. 4. A widely spreading depolarization wave was found in the embryonic CNS. This wave was triggered by multisensory inputs and spontaneous activities and was suggested to be mediated by a dual network of chemical synapses and gap junctions. It was also accompanied by a Ca2+ wave, indicating its nurturing effects on CNS development. [ABSTRACT FROM AUTHOR]

Multiple-site optical recording of neural activity was performed in the nucleus of the tractus solitarius (NTS) of the chick embryo with stimulation of the glossopharyngeal nerve (N. IX) and vagus nerve (N. X). We measured the amplitudes of the optical signals related to glutamate-mediated excitatory postsynaptic responses, and calculated the ratio of the signal evoked by simultaneous N. IX/N. X stimulation to the signal obtained after mathematical summation of the individual N. IX and N. X responses. The ratio was significantly lower than 100% in the rostral region of the NTS, in which postsynaptic responses were elicited by both N. IX and N. X stimulations. This result means that there is a convergence of visceral inputs via the N. IX and N. X in the embryonic chick NTS. The existence of the convergence suggests that the NTS performs complex integration of information from multiple sensory inputs from the early stages of embryogenesis. [Copyright &y& Elsevier]

To examine whether there are any differences in functional organization between the glossopharyngeal nerve (N. IX)- and vagus nerve (N. X)-projecting areas in the nucleus of the tractus solitarius (NTS), we performed optical recording of neural responses evoked by N. IX stimulation in 5- to 9-day-old embryonic chick brain stem preparations and compared the results with those in our previous studies concerning the N. X-related NTS. First, we investigated DL-2-amino-5-phosphonovaleric acid (APV)/Mg2+ sensitivity of the glutamatergic excitatory postsynaptic potentials (EPSPs) in the N. IX-related NTS. In 7- to 9-day-old preparations, we found regional differences in the degree of both the APV-induced reduction and Mg2+-free-induced enhancement of the EPSPs. We constructed developmental maps of spatial patterns of the APV- and Mg2+-sensitive components and showed that functional expression of the N-methyl-D-aspartate (NMDA) receptor dynamically changed during development. Second, we studied initial expression of synaptic functions in the N. IX-related NTS. In 6-day-old preparations, although action potentials alone were usually detected in normal Ringer solution, small EPSPs were elicited in a Mg2+-free solution. This result suggests that the NMDA receptor-mediated synaptic function is latently generated in the N. IX-related NTS at the 6-day-old embryonic stage and that external Mg2+ regulates the onset of synaptic functions. Developmental patterns of APV/Mg2+ sensitivity and the stage of initial expression of the glossopharyngeal EPSP were similar to those of the N. X, suggesting that the developmental sequence of the synaptic function in the NTS is the same for the N. IX- and N. X-related NTS. [ABSTRACT FROM AUTHOR]