“…Second, most of these neurons in bats have onset temporal patterns (Metzner and Radtke-Schuller, 1987; Covey and Casseday, 1991; Portfors and Wenstrup, 2001) that correspond closely to the inputs required to create the transient, onset-type facilitation observed in most IC neurons (Gans et al, 2009). Third, the level-tolerant response latencies of most VNLLc neurons (Covey and Casseday, 1991) are consistent with the observation that delay tuning in most IC facilitated neurons does not change with increasing sound level (Macias et al, 2012). These features of VNLLc neurons strengthen the conclusion that they provide the critical glycinergic inputs underlying combination-sensitive facilitation in IC (Figure 12).…”
“…Between 23 and 62% of tested IC neurons display facilitation, while 24–41% of tested neurons show inhibition without facilitation (Mittmann and Wenstrup, 1995; Portfors and Wenstrup, 1999; Leroy and Wenstrup, 2000; Nataraj and Wenstrup, 2005, 2006; Macias et al, 2012). The numbers reported in these studies likely vary due to different testing methods and neuronal populations sampled, and may also differ as a result of the different sub-species of mustached bats that were studied.…”
Section: Combination-sensitive Response Properties In the Inferior Comentioning
confidence: 99%
“…An input neuron with a phasic temporal pattern is consistent with these observations. Macias et al (2012) showed that most IC facilitated neurons have delay tuning that is relatively invariant with level of the high-frequency signal. This suggests that inputs to delay-tuned neurons have level-invariant response latencies.…”
Section: Combination-sensitive Response Properties In the Inferior Comentioning
confidence: 99%
“…Thus, in physiological studies of the mustached bat, auditory midbrain responses show the full range of frequency interactions, the low-frequency inhibition at 0 ms, and the range of best delays of facilitation that have been observed in auditory thalamus or cortex (Portfors and Wenstrup, 2003; Hagemann et al, 2011; Wenstrup and Portfors, 2011; Macias et al, 2012). Anatomical studies in this species show that combination-sensitive regions of the IC project to comparable regions of the medial geniculate body (MGB, Frisina et al, 1989; Wenstrup et al, 1994; Wenstrup and Grose, 1995) and that the combination-sensitive regions in MGB project to the appropriate cortical combination-sensitive areas (Pearson et al, 2007).…”
Section: Processing Of Combination-sensitive Responses Beyond the Midmentioning
This review describes mechanisms and circuitry underlying combination-sensitive response properties in the auditory brainstem and midbrain. Combination-sensitive neurons, performing a type of auditory spectro-temporal integration, respond to specific, properly timed combinations of spectral elements in vocal signals and other acoustic stimuli. While these neurons are known to occur in the auditory forebrain of many vertebrate species, the work described here establishes their origin in the auditory brainstem and midbrain. Focusing on the mustached bat, we review several major findings: (1) Combination-sensitive responses involve facilitatory interactions, inhibitory interactions, or both when activated by distinct spectral elements in complex sounds. (2) Combination-sensitive responses are created in distinct stages: inhibition arises mainly in lateral lemniscal nuclei of the auditory brainstem, while facilitation arises in the inferior colliculus (IC) of the midbrain. (3) Spectral integration underlying combination-sensitive responses requires a low-frequency input tuned well below a neuron's characteristic frequency (ChF). Low-ChF neurons in the auditory brainstem project to high-ChF regions in brainstem or IC to create combination sensitivity. (4) At their sites of origin, both facilitatory and inhibitory combination-sensitive interactions depend on glycinergic inputs and are eliminated by glycine receptor blockade. Surprisingly, facilitatory interactions in IC depend almost exclusively on glycinergic inputs and are largely independent of glutamatergic and GABAergic inputs. (5) The medial nucleus of the trapezoid body (MNTB), the lateral lemniscal nuclei, and the IC play critical roles in creating combination-sensitive responses. We propose that these mechanisms, based on work in the mustached bat, apply to a broad range of mammals and other vertebrates that depend on temporally sensitive integration of information across the audible spectrum.
“…Second, most of these neurons in bats have onset temporal patterns (Metzner and Radtke-Schuller, 1987; Covey and Casseday, 1991; Portfors and Wenstrup, 2001) that correspond closely to the inputs required to create the transient, onset-type facilitation observed in most IC neurons (Gans et al, 2009). Third, the level-tolerant response latencies of most VNLLc neurons (Covey and Casseday, 1991) are consistent with the observation that delay tuning in most IC facilitated neurons does not change with increasing sound level (Macias et al, 2012). These features of VNLLc neurons strengthen the conclusion that they provide the critical glycinergic inputs underlying combination-sensitive facilitation in IC (Figure 12).…”
“…Between 23 and 62% of tested IC neurons display facilitation, while 24–41% of tested neurons show inhibition without facilitation (Mittmann and Wenstrup, 1995; Portfors and Wenstrup, 1999; Leroy and Wenstrup, 2000; Nataraj and Wenstrup, 2005, 2006; Macias et al, 2012). The numbers reported in these studies likely vary due to different testing methods and neuronal populations sampled, and may also differ as a result of the different sub-species of mustached bats that were studied.…”
Section: Combination-sensitive Response Properties In the Inferior Comentioning
confidence: 99%
“…An input neuron with a phasic temporal pattern is consistent with these observations. Macias et al (2012) showed that most IC facilitated neurons have delay tuning that is relatively invariant with level of the high-frequency signal. This suggests that inputs to delay-tuned neurons have level-invariant response latencies.…”
Section: Combination-sensitive Response Properties In the Inferior Comentioning
confidence: 99%
“…Thus, in physiological studies of the mustached bat, auditory midbrain responses show the full range of frequency interactions, the low-frequency inhibition at 0 ms, and the range of best delays of facilitation that have been observed in auditory thalamus or cortex (Portfors and Wenstrup, 2003; Hagemann et al, 2011; Wenstrup and Portfors, 2011; Macias et al, 2012). Anatomical studies in this species show that combination-sensitive regions of the IC project to comparable regions of the medial geniculate body (MGB, Frisina et al, 1989; Wenstrup et al, 1994; Wenstrup and Grose, 1995) and that the combination-sensitive regions in MGB project to the appropriate cortical combination-sensitive areas (Pearson et al, 2007).…”
Section: Processing Of Combination-sensitive Responses Beyond the Midmentioning
This review describes mechanisms and circuitry underlying combination-sensitive response properties in the auditory brainstem and midbrain. Combination-sensitive neurons, performing a type of auditory spectro-temporal integration, respond to specific, properly timed combinations of spectral elements in vocal signals and other acoustic stimuli. While these neurons are known to occur in the auditory forebrain of many vertebrate species, the work described here establishes their origin in the auditory brainstem and midbrain. Focusing on the mustached bat, we review several major findings: (1) Combination-sensitive responses involve facilitatory interactions, inhibitory interactions, or both when activated by distinct spectral elements in complex sounds. (2) Combination-sensitive responses are created in distinct stages: inhibition arises mainly in lateral lemniscal nuclei of the auditory brainstem, while facilitation arises in the inferior colliculus (IC) of the midbrain. (3) Spectral integration underlying combination-sensitive responses requires a low-frequency input tuned well below a neuron's characteristic frequency (ChF). Low-ChF neurons in the auditory brainstem project to high-ChF regions in brainstem or IC to create combination sensitivity. (4) At their sites of origin, both facilitatory and inhibitory combination-sensitive interactions depend on glycinergic inputs and are eliminated by glycine receptor blockade. Surprisingly, facilitatory interactions in IC depend almost exclusively on glycinergic inputs and are largely independent of glutamatergic and GABAergic inputs. (5) The medial nucleus of the trapezoid body (MNTB), the lateral lemniscal nuclei, and the IC play critical roles in creating combination-sensitive responses. We propose that these mechanisms, based on work in the mustached bat, apply to a broad range of mammals and other vertebrates that depend on temporally sensitive integration of information across the audible spectrum.
“…A neuron was considered to respond to a given echo delay-echo level combination whenever it fired above 50% of the maximum response observed in the DRF. The criteria used here to define a response is the same as that used in previous studies to define the borders of DRFs 6,7,22,24 .…”
Echolocating bats use the time from biosonar pulse emission to the arrival of echo (defined as echo delay) to calculate the space depth of targets. In the dorsal auditory cortex of several species, neurons that encode increasing echo delays are organized rostrocaudally in a topographic arrangement defined as chronotopy. Precise chronotopy could be important for precise target-distance computations. Here we show that in the cortex of three echolocating bat species (Pteronotus quadridens, Pteronotus parnellii and Carollia perspicillata), chronotopy is not precise but blurry. In all three species, neurons throughout the chronotopic map are driven by short echo delays that indicate the presence of close targets and the robustness of map organization depends on the parameter of the receptive field used to characterize neuronal tuning. The timing of cortical responses (latency and duration) provides a binding code that could be important for assembling acoustic scenes using echo delay information from objects with different space depths.
It has been reported previously that in the inferior colliculus of the bat Molossus molossus, neuronal duration tuning is ambiguous because the tuning type of the neurons dramatically changes with the sound level. In the present study, duration tuning was examined in the auditory cortex of M. molossus to describe if it is as ambiguous as the collicular tuning. From a population of 174 cortical 104 (60 %) neurons did not show duration selectivity (all-pass). Around 5 % (9 units) responded preferentially to stimuli having longer durations showing long-pass duration response functions, 35 (20 %) responded to a narrow range of stimulus durations showing band-pass duration response functions, 24 (14 %) responded most strongly to short stimulus durations showing short-pass duration response functions and two neurons (1 %) responded best to two different stimulus durations showing a two-peaked duration-response function. The majority of neurons showing short- (16 out of 24) and band-pass (24 out 35) selectivity displayed "O-shaped" duration response areas. In contrast to the inferior colliculus, duration tuning in the auditory cortex of M. molossus appears level tolerant. That is, the type of duration selectivity and the stimulus duration eliciting the maximum response were unaffected by changing sound level.
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