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1.  Interaural timing difference circuits in the auditory brainstem of the emu (Dromaius novaehollandiae) 
In the auditory system, precise encoding of temporal information is critical for sound localization, a task with direct behavioral relevance. Interaural timing differences are computed using axonal delay lines and cellular coincidence detectors in nucleus laminaris (NL). We present morphological and physiological data on the timing circuits in the emu, Dromaius novaehollandiae, and compare these results with those from the barn owl (Tyto alba) and the domestic chick (Gallus gallus). Emu NL was composed of a compact monolayer of bitufted neurons whose two thick primary dendrites were oriented dorsoventrally. They showed a gradient in dendritic length along the presumed tonotopic axis. The NL and nucleus magnocellularis (NM) neurons were strongly immunoreactive for parvalbumin, a calcium-binding protein. Antibodies against synaptic vesicle protein 2 and glutamic acid decarboxlyase revealed that excitatory synapses terminated heavily on the dendritic tufts, while inhibitory terminals were distributed more uniformly. Physiological recordings from brainstem slices demonstrated contralateral delay lines from NM to NL. During whole-cell patch-clamp recordings, NM and NL neurons fired single spikes and were doubly-rectifying. NL and NM neurons had input resistances of 30.0 ± 19.9 MΩ and 49.0 ± 25.6 MΩ, respectively, and membrane time constants of 12.8 ± 3.8 ms and 3.9 ± 0.2 ms. These results provide further support for the Jeffress model for sound localization in birds. The emu timing circuits showed the ancestral (plesiomorphic) pattern in their anatomy and physiology, while differences in dendritic structure compared to chick and owl may indicate specialization for encoding ITDs at low best frequencies.
PMCID: PMC2948976  PMID: 16435285
avian; nucleus laminaris; nucleus magnocellularis; dendrite; coincidence detection; sound localization
2.  Short-term synaptic plasticity and intensity coding 
Hearing research  2011;279(1-2):13-21.
Alterations in synaptic strength over short time scales, termed short-term synaptic plasticity, can gate the flow of information through neural circuits. Different information can be extracted from the same presynaptic spike train depending on the activity- and time-dependent properties of the plasticity at a given synapse. The parallel processing in the brain stem auditory pathways provides an excellent model system for investigating the functional implications of short-term plasticity in neural coding. We review recent evidence that short-term plasticity differs in different pathways with a special emphasis on the ‘intensity’ pathway. While short-term depression dominates the ‘timing’ pathway, the intensity pathway is characterized by a balance of short-term depression and facilitation that allows linear transmission of rate-coded intensity information. Target-specific regulation of presynaptic plasticity mechanisms underlies the differential expression of depression and facilitation. The potential contribution of short-term plasticity to different aspects of ‘intensity’-related information processing, such as interaural level/intensity difference coding, amplitude modulation coding, and intensity-dependent gain control coding, is discussed.
PMCID: PMC3210195  PMID: 21397676
3.  A rapid form of activity-dependent recovery from short-term synaptic depression in the intensity pathway of the auditory brainstem 
Biological Cybernetics  2011;104(3):209-223.
Short-term synaptic plasticity acts as a time- and firing rate-dependent filter that mediates the transmission of information across synapses. In the avian auditory brainstem, specific forms of plasticity are expressed at different terminals of the same auditory nerve fibers and contribute to the divergence of acoustic timing and intensity information. To identify key differences in the plasticity properties, we made patch-clamp recordings from neurons in the cochlear nucleus responsible for intensity coding, nucleus angularis, and measured the time course of the recovery of excitatory postsynaptic currents following short-term synaptic depression. These synaptic responses showed a very rapid recovery, following a bi-exponential time course with a fast time constant of ~40 ms and a dependence on the presynaptic activity levels, resulting in a crossing over of the recovery trajectories following high-rate versus low-rate stimulation trains. We also show that the recorded recovery in the intensity pathway differs from similar recordings in the timing pathway, specifically the cochlear nucleus magnocellularis, in two ways: (1) a fast recovery that was not due to recovery from postsynaptic receptor desensitization and (2) a recovery trajectory that was characterized by a non-monotonic bump that may be due in part to facilitation mechanisms more prevalent in the intensity pathway. We tested whether a previously proposed model of synaptic transmission based on vesicle depletion and sequential steps of vesicle replenishment could account for the recovery responses, and found it was insufficient, suggesting an activity-dependent feedback mechanism is present. We propose that the rapid recovery following depression allows improved coding of natural auditory signals that often consist of sound bursts separated by short gaps.
PMCID: PMC3257163  PMID: 21409439
Auditory nerve; Cochlear nucleus; Angularis; Magnocellularis; Short-term depression; Short-term facilitation; Vesicle cycling
5.  Microseconds Matter 
PLoS Biology  2010;8(6):e1000405.
This Primer focuses on detection of the small interaural time differences that underlie sound localization.
PMCID: PMC2893944  PMID: 20613856

Results 1-5 (5)