Differences in intensity and arrival time of sounds at the two ears, interaural intensity and time differences (IID, ITD), are the chief cues for sound localization. Both cues are initially processed in the superior olivary complex (SOC), which projects to the dorsal nucleus of the lateral lemniscus (DNLL) and the auditory midbrain. Here we present basic response properties of low-frequency (< 2 kHz) DNLL neurons and their binaural sensitivity to ITDs and IIDs in the anesthetized gerbil. We found many neurons showing binaural properties similar to those reported for SOC neurons. IID-properties were similar to that of the contralateral lateral superior olive (LSO). A majority of cells had an ITD sensitivity resembling that of either the ipsilateral medial superior olive (MSO) or the contralateral LSO. A smaller number of cells displayed intermediate types of ITD sensitivity. In neurons with MSO-like response ITDs that evoked maximal discharges were mostly outside of the range of ITDs the gerbil naturally experiences. The maxima of the first derivative of their ITD-functions (steepest slope), however, were well within the physiological range of ITDs. This finding is consistent with the concept of a population rather than a place code for ITDs. Moreover, we describe several other binaural properties as well as physiological and anatomical evidence for a small but significant input from the contralateral MSO. The large number of ITD-sensitive low-frequency neurons implicates a substantial role for the DNLL in ITD processing and promotes this nucleus as a suitable model for further studies on ITD-coding.
Psychophysical forward masking is an increase in threshold of detection of a sound (probe) when it is preceded by another sound (masker). This is reminiscent of the reduction in neuronal responses to a sound following prior stimulation. Studies in the auditory nerve and cochlear nucleus using signal detection theory techniques to derive neuronal thresholds showed that in centrally projecting neurons, increases in masked thresholds were significantly smaller than the changes measured psychophysically. Larger threshold shifts have been reported in the inferior colliculus of awake marmoset. The present study investigated the magnitude of forward masking in primary auditory cortical neurons of anaesthetised guinea-pigs. Responses of cortical neurons to unmasked and forward masked tones were measured and probe detection thresholds estimated using signal detection theory methods. Threshold shifts were larger than in the auditory nerve, cochlear nucleus and inferior colliculus. The larger threshold shifts suggest that central, and probably cortical, processes contribute to forward masking. However, although methodological differences make comparisons difficult, the threshold shifts in cortical neurons were, in contrast to subcortical nuclei, actually larger than those observed psychophysically. Masking was largely attributable to a reduction in the responses to the probe, rather than either a persistence of the masker responses or an increase in the variability of probe responses.
The spike activity of neuromodulatory dorsal unpaired median (DUM) neurons was analyzed during a pilocarpine-induced motor pattern in the locust. Paired intracellular recordings were made from these octopaminergic neurons during rhythmic activity in hindleg motor neurons evoked by applying pilocarpine to an isolated metathoracic ganglion. This motor pattern is characterized by two alternating phases: a levator phase, during which levator, flexor, and common inhibitor motor neurons spike, and a depressor phase, during which depressor and extensor motor neurons spike. Three different subpopulations of efferent DUM neurons could be distinguished during this rhythmical motor pattern according to their characteristic spike output. DUM 1 neurons, which in the intact animal do not innervate muscles involved in leg movements, showed no change apart from a general increase in spike frequency. DUM 3 and DUM 3,4 neurons produced the most variable activity but received frequent and sometimes pronounced hyperpolarizations that were often common to both recorded neurons. DUM 5 and DUM 3,4,5 neurons innervate muscles of the hindleg and showed rhythmical excitation leading to bursts of spikes during rhythmic activity of the motor neurons, which innervate these same muscles. Sometimes the motor output was coordinated across both sides of the ganglion so that there was alternating activity between levators of both sides. In these cases, the spikes of DUM 5 and DUM 3,4,5 neurons and the hyperpolarization of DUM 3 and DUM 3,4 neurons occurred at particular phases in the motor pattern. Our data demonstrate a central coupling of specific types of DUM neurons to a rhythmical motor pattern. Changes in the spike output of these particular efferent DUM neurons parallel changes in the motor output. The spike activity of DUM neurons thus may be controlled by the same circuits that determine the action of the motor neurons. Functional implications for real walking are discussed.
1 Although barbiturates, like other general anaesthetics, depress excitatory synaptic transmission in the central nervous system (CNS), the underlying cellular mechanisms remain unresolved. They may increase the likelihood that an action potential will fail to invade every branch of the axonal arbour, thereby decreasing the synaptic drive to the postsynaptic neurons. Alternatively, they may inhibit calcium entry into the presynaptic terminals, thus reducing transmitter release. 2 To resolve these issues, we have used two-photon microscopy to monitor calcium transients evoked by action potentials in axons, axonal varicosities (synaptic boutons) and fine axon collaterals of hippocampal CA1 neurons. 3 Pentobarbitone (75 -300 mM) did not block the invasion of the axonal arbour or the synaptic boutons, but it did reduce the amplitude of the calcium transients recorded from the axons in a concentration-dependent manner. At 150 mM, pentobarbitone reduced the transients to 7874% of the control. 4 Pentobarbitone depressed the calcium transients recorded from the synaptic boutons in a concentration-dependent manner. When 150 mM pentobarbitone was applied, the calcium transients recorded from the boutons were 5373% of the control. This concentration of pentobarbitone also reduced the amplitude and frequency of the spontaneous excitatory postsynaptic potentials to 5474 and 42717% of the control, respectively. 5 The local anaesthetic procaine (500 mM) had no significant effect on action potential invasion of axon collaterals, even though it reduced the action potential amplitude by 25%. 6 This data are consistent with the notion that the pentobarbitone-induced depression of presynaptic calcium transients contributes to its depressant effect on excitatory synaptic transmission in the CNS.
The segmental ganglia of the locust contain efferent neuromodulatory neurones with cell bodies at the dorsal midline and axons that supply muscles and other tissue on both sides of the body. These are the dorsal unpaired median (DUM) neurones. Intracellular recordings were made from pairs of known metathoracic efferent DUM neurones in locusts in which all nerves were intact and in isolated metathoracic ganglia. The 19 metathoracic, efferent DUM neurones were identified according to the nerve roots through which their axons emerge from the ganglion. The synaptic potentials in these DUM neurones have been analysed to investigate how these neurones are activated and how their spikes are controlled. The degree of correlation between the synaptic potentials in particular pairs of neurones was quantified using a correlation analysis. This allowed the population of DUM neurones to be divided into three subsets that also map onto an anatomical grouping based on the distribution of their axons in the lateral nerves: (i) DUM1 neurones (DUMDL and DUM1b); (ii) DUM3 and DUM3,4 neurones; and (iii) DUM3,4,5, DUM5b neurones and DUMETi. Individual neurones within each subset showed strong correlations between their synaptic potentials, in both intact locusts and isolated ganglia, and tended to spike at the same time. Neurones in different subsets had few synaptic potentials in common and tended to spike independently. The persistence of common synaptic potentials in neurones of the three subsets in isolated ganglia indicates that they are derived from neurones within the metathoracic ganglion. The DUM neurones that had many common synaptic potentials in a quiescent locust responded in similar ways to mechanosensory stimulation of different parts of the body. DUM3,4, 5 and DUM5 neurones gave the clearest and most consistent responses to stimulation of mechanoreceptors on either hind leg. DUM3 and DUM3, 4 neurones responded variably, but usually with a hyperpolarisation. DUM1 neurones were rarely excited by mechanosensory stimuli but, like the preceding group, their responses were dependent upon whether the locust was moving its legs. These results lend further support to the idea that there is a subdivision of action amongst this population of DUM neurones, with those supplying the same targets being driven by the same presynaptic local neurones.
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