Synaptic Inhibition Influences the Temporal Coding Properties of Medial Superior Olivary Neurons. An in vitro Study

The medial superior olive (MSO) functions as a coincidence detector for interaural time and phase differences by integrating excitatory synaptic inputs. Recent studies demonstrating glycinergic projections to MSO neurons suggest that coincidence detection results from the temporal integration of both EPSPs and IPSPs. We examined the impact of synaptic inhibition on the temporal coding properties of gerbil MSO neurons in vitro with intracellular recordings and electrical stimulation. For low-level bilateral electric stimulation, the EPSPs summated to produce an action potential in 73% of MSO neurons if they occurred within 50–500 microseconds of one another. Synaptic inhibition became more prominent at higher stimulus amplitudes in 73% of MSO neurons, and could block an evoked action potential if the stimuli to each pathway were delivered within 250 microseconds of one another. The glycine receptor antagonist strychnine influenced the response to simulated interaural time differences. In the presence of strychnine, interstimulus delays that originally resulted in full action potential suppression were sufficient to evoke an action potential. For trains of stimuli, as stimulus intensity increased (spatial summation), or as stimulus repetition rate increased to 100–500 Hz (temporal summation), there was a decrease in the number of stimulus pulses that evoked an action potential. In the presence of strychnine, MSO neurons generated a greater percentage of action potentials to the stimulus trains. When stimulus trains were delivered bilaterally, MSO neurons fired a greater number of action potentials at specific interstimulus time differences, and were selectively inhibited at other time differences.(ABSTRACT TRUNCATED AT 250 WORDS)</jats:p

Asymmetric Excitatory Synaptic Dynamics Underlie Interaural Time Difference Processing in the Auditory System

In order to localize sounds in the environment, the auditory system detects and encodes differences in signals between each ear. The exquisite sensitivity of auditory brain stem neurons to the differences in rise time of the excitation signals from the two ears allows for neuronal encoding of microsecond interaural time differences

Afferent regulation of inhibitory synaptic transmission in the developing auditory midbrain

To determine whether afferent innervation regulates the strength of inhibitory connections in the gerbil auditory midbrain, both cochleas were surgically removed in postnatal day 7 animals, before sound-driven activity is first observed. Inhibitory synaptic currents were measured in a brain slice preparation 1–7 d after the ablations. Whole-cell and gramicidinperforated patch recordings were obtained from inferior colliculus neurons, and IPSCs were evoked by stimulation of the commissure of the inferior colliculus (CIC) or the ipsilateral lateral lemniscus (LL) in the presence of kynurenic acid. Deafferentation led to a 24 mV depolarizing shift in the IPSC equilibrium potential within 1 d of deafferentation. As a consequence, there was a large reduction of IPSC amplitude at a holding potential of �20 mV in neurons from bilaterally ablated animals. Furthermore, both afferent pathways displayed a 50

Ó Springer-Verlag 2003 Gain adjustment of inhibitory synapses in the auditory system

Abstract. A group of central auditory neurons residing in the lateral superior olivary nucleus (LSO) responds selectively to interaural level differences and may contribute to sound localization. In this simple circuit, ipsilateral sound increases firing of LSO neurons, whereas contralateral sound inhibits the firing rate via activation of the medial nucleus of the trapezoid body (MNTB). During development, individual MNTB fibers arborize within the LSO, but they undergo a restriction of their boutons that ultimately leads to mature topography. A critical issue is whether a distinct form of inhibitory synaptic plasticity contributes to MNTB synapse elimination within LSO. Whole-cell recording from LSO neurons in brain slices from developing gerbils show robust long-term depression (LTD) of the MNTBevoked IPSP/Cs when the MNTB was activated at a low frequency (1 Hz). These inhibitory synapses also display mixed GABA/glycinergic transmission during development, as assessed physiologically and immunohistochemically (Kotak et al. 1998). While either glycine or GABAA receptors could independently display inhibitory LTD, focal delivery of GABA, but not glycine, at the postsynaptic-locus induces depression. Furthermore, the GABAB receptor antagonist, SCH-50911, prevents GABA or synaptically induced depression. Preliminary evidence also indicated strengthening of inhibitory transmission (LTP) by a distinct pattern of inhibitory activity. These data support the idea that GABA is crucial for the expression inhibitory LTD and that this plasticity may underlie the early refinement of inhibitory synaptic connections in the LSO

Developmental Conductive Hearing Loss Reduces Modulation Masking Release

Hearing-impaired individuals experience difficulties in detecting or understanding speech, especially in background sounds within the same frequency range. However, normally hearing (NH) human listeners experience less difficulty detecting a target tone in background noise when the envelope of that noise is temporally gated (modulated) than when that envelope is flat across time (unmodulated). This perceptual benefit is called modulation masking release (MMR). When flanking masker energy is added well outside the frequency band of the target, and comodulated with the original modulated masker, detection thresholds improve further (MMR+). In contrast, if the flanking masker is antimodulated with the original masker, thresholds worsen (MMR−). These interactions across disparate frequency ranges are thought to require central nervous system (CNS) processing. Therefore, we explored the effect of developmental conductive hearing loss (CHL) in gerbils on MMR characteristics, as a test for putative CNS mechanisms. The detection thresholds of NH gerbils were lower in modulated noise, when compared with unmodulated noise. The addition of a comodulated flanker further improved performance, whereas an antimodulated flanker worsened performance. However, for CHL-reared gerbils, all three forms of masking release were reduced when compared with NH animals. These results suggest that developmental CHL impairs both within- and across-frequency processing and provide behavioral evidence that CNS mechanisms are affected by a peripheral hearing impairment

Central neural motor programs underlying short- and long-term patterns of \u3cem\u3eLimulus\u3c/em\u3e respiratory activity

Isolated, unstimulated abdominal ventral nerve cords ofLimulus display patterns of motor output characteristic of rhythmic gill ventilation and of gill cleaning (Fig. 2). The motor output may occur as long-term patterns of alternate gill cleaning and ventilation (Fig. 3), or alternate gill cleaning and tonic activity (Fig. 4). Two patterns of gill cleaning motor output, apparently corresponding to the left-leading and right-leading patterns observed in intact animals, persist in the isolated ventral cord (Fig. 4). These two patterns occur with the same rough alternation as in intact animals. Thus all the patterns of gill-plate movement in intact animals, except those involving swimming, have underlying motor programs that are expressed in isolated nerve cords in the absence of stimulation or of sensory feedback. These findings extend the concept of central pattern generators to include complex and long-duration patterns of stereotyped behavior such as those underlying the movements of theLimulus abdominal appendages