Rebound depolarization in single units of the ventral cochlear nucleus: a contribution to grouping by common onset?

Simultaneous grouping by common onset time is believed to be a powerful cue in auditory perception; components that start or stop roughly at the same time are judged as far more likely to have originated from the same source. Here we report a simple experiment designed to simulate a complex psychophysical paradigm first described by Darwin and Sutherland [(1984) Grouping frequency components of vowels. When is a harmonic not a harmonic? Quarterly J of Experimental Psychology: Hum Exp Psychol 36(A):193-208]. It is possible to change the perception of the vowel /I/ to /epsilon/ by manipulating the harmonics around the first formant (F1). Increasing the amplitude of one harmonic around F1 caused the perception of the vowel to change from /I/ to /epsilon/. Extending the increased component before the vowel could, however, greatly reduce this change. The role of neural adaptation in this effect was questioned by repeating the experiment but this time using a 'captor' tone which was switched on with the asynchronous harmonic and off when the vowel started. This time the vowel percept did change in a fashion analogous to the effect of an increase in the amplitude of the fourth harmonic (which is close to F1). This effect was explained by assuming that the captor had grouped with the leading portion of the asynchronous component enabling the remainder of the asynchronous component to be grouped with the remainder of the components. We propose a relatively low-level neuronal explanation for this grouping effect: the captor reduces the neural response to the leading segment of the asynchronous component by activating across-frequency suppression, either from the cochlea, or acting via a wideband inhibitor in the ventral cochlear nucleus. The reduction in neural response results in a release from adaptation with the offset of the captor terminating the inhibition, such that the response to the continuation of that component is now enhanced. Using a simplified paradigm we show that both primary-like and chopper units in the ventral cochlear nucleus of the anesthetized guinea pig may show a rebound in excitation when a captor is positioned so as to stimulate the suppressive sidebands in its receptive field. The strength of the rebound was positively correlated with the strength of the suppression. These and other results are consistent with the view that low-level mechanisms underlie the psychophysical captor effect.

Age-related changes in auditory spatial properties of the guinea pig superior colliculus.

The map of auditory space located in the deep layers of the guinea pig superior colliculus (SC) is a complex computational representation of the auditory azimuth surrounding the animal. The map undergoes a protracted developmental profile during the first postnatal month and remains plastic until well into adulthood. However, there are no data concerning the state of the collicular auditory space map in much older animals. Multi-unit responses to broadband noise stimuli presented around the azimuthal plane under anechoic conditions were recorded from the deep SC of guinea pigs of a variety of ages, up to 44 months. The data obtained show that the map remains stable up to the age of approximately 36 months. However, after this age, the map shows rapid deterioration and at 42 months, multi-unit responses did not show features consistent with a normal map. It appears that deficits accruing within the central auditory system with increasing age, combine to overcome the ability of the mechanisms of plasticity responsible for space map maintenance to keep pace with the changes, resulting in degraded SC spatial tuning with increasing age.

The effects of monocular enucleation on the representation of auditory space in the superior colliculus of the guinea-pig.

Multi-unit responses, to free-field auditory stimuli, in the superior colliculus were investigated in guinea-pigs following earlier removal of one eye. Enucleation resulted in disruption of the normal tuning parameter values and of the topographical precision of auditory responses in the SC both ipsilateral and contralateral to the enucleated eye. These data demonstrate that monocular enucleation prevents the normal development of the superior collicular auditory space map.

The effect of dark-rearing, strobe-rearing and acute visual cortex removal on the visual responses in the superficial superior colliculus of the guinea-pig.

Extracellular multi-unit responses to visual stimuli were recorded in the cells of the superficial layers of the superior colliculus (SC) in four groups of adult guinea-pigs: a control group, a strobe-reared group, a dark-reared group and a group with the ipsilateral visual cortex removed acutely. Single unit visual responses were also recorded in a control and a dark-reared group. When guinea-pigs were either strobe or dark-reared from birth, the number of directionally selective responses in the superficial SC decreased significantly. Acute removal of the visual cortex had no affect on the number of directionally selective cells recorded in the SC. The correlation between azimuthal visual receptive field and rostrocaudal position of the recording electrode in the SC was not significantly different from the control group following strobe, dark-rearing or acute visual cortex removal. These data imply that, during early development, visual information is necessary for directional selectivity of the visual responses in the superficial SC. However, the map of visual azimuthal space is essentially unperturbed by visual restriction (in the form of dark or strobe-rearing) or acute visual cortex removal.

Hair cell loss in the aged guinea pig cochlea.

The various effects of ageing on the auditory system, collectively termed presbycusis, are being studied across a wide range of animal species, including humans. One contributing factor to presbycusis is thought to be losses of the sensory hair cells in the cochlea. In this study, hair cell counts were obtained from cochleas of pigmented guinea pigs (Cavia porcellus) at ages ranging from 11 days to 4 years 7 months, using scanning electron microscopy to visualize the organ of Corti. Representative samples of the basal, middle and apical turn of the cochlea were photographed for analysis. Hair cell loss was observed, even in young animals. However, the loss was greater in the aged animals, but was not distributed evenly throughout the length of the cochlea. No significant loss of hair cells was seen in the basal (high frequency) or middle turn of the cochlea of the aged animals. In the apical (low frequency) turn, there was a significant loss of hair cells in all rows of outer hair cells (up to around 20%), and was most severe in the third row. There was no loss of apical inner hair cells in the aged animals.

The auditory brainstem response of aged guinea pigs.

The auditory brainstem response (ABR) technique was used to investigate potential dysfunctions in the auditory brainstem of the pigmented guinea pig (Cavia porcellus) associated with biological ageing. Animals aged from 58 days to 4 years 3 months were tested. ABRs were recorded at stimulation intensities from 85 dB HL to -10 dB HL. The auditory thresholds were found to undergo marked elevations in old animals, by an average of 32 dB. From the traces obtained, four positive deflection waves were reliably recorded. The latency of each of the four waves was evaluated at different stimulation intensities in guinea pigs of different ages. Although there was a trend for the latencies to increase in old age, these differences were not statistically significant. Similarly, there were no significant age-related changes in the inter-peak intervals. The latency/intensity functions of the four waves produced parallel curves. However, the curve from the old age group was shifted to the right, by an average of 35 dB, indicative of conductive hearing loss. There was no evidence of retro-cochlear hearing loss. Therefore, it appears that the threshold elevations in the old animals can be accounted for by conductive hearing loss, presumably in the middle ear. In 24% of the old animals tested, no ABR could be elicited. It would appear that these animals had suffered severe sensorineural hearing loss.

Spatial receptive fields of inferior colliculus neurons to auditory apparent motion in free field.

We examined responses from 91 single-neurons in the inferior colliculus (IC) of anesthetized guinea pigs to auditory apparent motion in the free field. Apparent motion was generated by presenting 100-ms tone bursts, separated by 50-ms silent intervals, at consecutive speaker positions in an array of 11 speakers, positioned in an arc +/-112.5 degrees around midline. Most neurons demonstrated discrete spatial receptive fields (SRFs) to apparent motion in the clockwise and anti-clockwise directions. However, SRFs showed marked differences for apparent motion in opposite directions. In virtually all neurons, mean best azimuthal positions for SRFs to opposite directions occurred at earlier positions in the motion sweep, producing receptive fields to the two directions of motion that only partially overlapped. Despite this, overall spike counts to the two directions were similar for equivalent angular velocities. Responses of 28 neurons were recorded to stimuli with different duration silent intervals between speaker presentations, mimicking different apparent angular velocities. Increasing the stimulus OFF time increased neuronal discharge rates, particularly at later portions of the apparent motion sweep, and reduced the differences in the SRFs to opposite motion directions. Consequently SRFs to both directions broadened and converged with decreasing motion velocity. This expansion was most obvious on the outgoing side of the each SRF. Responses of 11 neurons were recorded to short (90 degrees ) partially overlapping apparent motion sweeps centered at different spatial positions. Nonoverlapping response profiles were recorded in 9 of the 11 neurons tested and confirmed that responses at each speaker position were dependent on the preceding response history. Together these data are consistent with the suggestion that a mechanism of adaptation of excitation contributes to the apparent sensitivity of IC neurons to auditory motion cues. In addition, the data indicate that the sequential activation of an array of speakers to produce apparent auditory motion may not be an optimal stimulus paradigm to separate the temporal and spatial aspects of auditory motion processing.

Visual movement and pattern are important for the development of a map of auditory space in the guinea pig superior colliculus.

Previous data have indicated that, if guinea pigs are deprived of all visual information during a crucial period early in development (26-30 days after birth), the map of auditory space in the superior colliculus (SC) is completely disrupted. In the experiments reported here, multi-unit auditory receptive fields were recorded in the SC of two groups of anaesthetised guinea pigs that had been exposed to different forms of visual deprivation. One group was reared in a movement-free environment (strobe-reared) and the other group was reared in a pattern-free environment (their eyes covered with light-diffusing masks). Both groups experienced visual restriction during the crucial period for auditory space map development. In both experimental groups, the multi-unit auditory receptive fields were broad and all spatial tuning parameter values were significantly greater than the equivalent values from a control group of normal animals. In the pattern- and motion-deprived groups, a significant correlation existed between the rostro-caudal position of the recording electrode in the SC and the peak response angle of the receptive field, thus showing a degree of topographic organisation of the auditory receptive fields in the SC. However, the topographic order was less precise than that displayed by the control group of animals. These results indicate that, during development, both visual pattern and movement are important for the refinement of the SC auditory space map in the guinea pig.

Neurotransmitter involvement in development and maintenance of the auditory space map in the guinea pig superior colliculus.

Neurotransmitter involvement in development and maintenance of the auditory space map in the guinea pig superior colliculus. J. Neurophysiol. 80: 2941-2953, 1998. The mammalian superior colliculus (SC) is a complex area of the midbrain in terms of anatomy, physiology, and neurochemistry. The SC bears representations of the major sensory modalites integrated with a motor output system. It is implicated with saccade generation, in behavioral responses to novel sensory stimuli and receives innervation from diverse regions of the brain using many neurotransmitter classes. Ethylene-vinyl acetate copolymer (Elvax-40W polymer) was used here to deliver chronically neurotransmitter receptor antagonists to the SC of the guinea pig to investigate the potential role played by the major neurotransmitter systems in the collicular representation of auditory space. Slices of polymer containing different drugs were implanted onto the SC of guinea pigs before the development of the SC azimuthal auditory space map, at approximately 20 days after birth (DAB). A further group of animals was exposed to aminophosphonopentanoic acid (AP5) at approximately 250 DAB. Azimuthal spatial tuning properties of deep layer multiunits of anesthetized guinea pigs were examined approximately 20 days after implantation of the Elvax polymer. Broadband noise bursts were presented to the animals under anechoic, free-field conditions. Neuronal responses were used to construct polar plots representative of the auditory spatial multiunit receptive fields (MURFs). Animals exposed to control polymer could develop a map of auditory space in the SC comparable with that seen in unimplanted normal animals. Exposure of the SC of young animals to AP5, 6-cyano-7-nitroquinoxaline-2,3-dione, or atropine, resulted in a reduction in the proportion of spatially tuned responses with an increase in the proportion of broadly tuned responses and a degradation in topographic order. Thus N-methyl--aspartate (NMDA) and non-NMDA glutamate receptors and muscarinic acetylcholine receptors appear to play vital roles in the development of the SC auditory space map. A group of animals exposed to AP5 beginning at approximately 250 DAB produced results very similar to those obtained in the young group exposed to AP5. Thus NMDA glutamate receptors also seem to be involved in the maintenance of the SC representation of auditory space in the adult guinea pig. Exposure of the SC of young guinea pigs to gamma-aminobutyric acid (GABA) receptor blocking agents produced some but not total disruption of the spatial tuning of auditory MURFs. Receptive fields were large compared with controls, but a significant degree of topographical organization was maintained. GABA receptors may play a role in the development of fine tuning and sharpening of auditory spatial responses in the SC but not necessarily in the generation of topographical order of the these responses.

Plasticity in the superior collicular auditory space map of adult guinea-pigs.

In the guinea-pig the development and maintenance of the superior collicular (SC) auditory space map requires both auditory and visual experience. This paper reports the results of experiments in which adult animals (of different ages) were dark-reared for 4 weeks to define the extent of the period of susceptibility to visual deprivation of the SC auditory map. Dark-rearing for 4 weeks from 100, 150 or 200 days after birth caused degradation of spatial tuning and topography of multi-unit auditory responses in the SC. In contrast, animals reared in the dark from 250 days after birth showed spatially tuned auditory responses similar to those seen in normal animals. These data suggest that the SC auditory space map in the adult guinea-pig remains susceptible to visual deprivation up to 200 days after birth.

Urodynamic variables cannot be used to classify the severity of detrusor instability.

OBJECTIVE: To explore the relationship between subjective severity of symptoms of detrusor instability (DI) on presentation, outcome after treatment for DI and initial diagnostic urodynamic variables, with the aim of identifying a urodynamic variable which might, by predicting a favourable outcome from treatment, classify the severity of DI. PATIENTS AND METHODS: Women with a urodynamically proven diagnosis of DI were recruited prospectively for the study. Data on disease symptoms and variables from their diagnostic cystometrogram were collected. All women were then treated and their outcome at 6 weeks after treatment compared with the initial urodynamic variables. Data on severity of symptoms were compared with initial urodynamic variables to explore any differences in these variables attributable to symptom severity. RESULTS: Of 300 women studied (mean age 54 years, SD 16), 290 were treated with oxybutynin and bladder retraining. At 6 weeks, 82 women had their treatment outcome classified as worse/no change; 218 women had improved. When good or poor outcome was compared with the urodynamic results, there was no significant difference between the groups. Likewise, the severity of symptoms did not relate to the values of urodynamic variables. CONCLUSIONS: There was no statistically significant relationship between reported severity of symptoms and urodynamic variables, and no relationship between the urodynamic variables used and response to treatment. Therefore, using these values it is not possible to predict a favourable outcome from treatment or to use them to classify disease severity.

Post-crucial period effects of visual experience and deprivation on the guinea-pig superior collicular map of auditory space.

There were two primary aims of this study. First, to observe if the map of auditory space in the superior colliculus (SC) of the guinea-pig could recover after periods of normal visual experience following visual deprivation during the crucial period. Second, to determine whether any degradation of the space map was observed when the animal was visually deprived for different lengths of time after the crucial period. Animals deprived of visual experience during the crucial period and then allowed normal experience did show a limited ability to construct a SC auditory space map. Whereas visual deprivation following normal auditory and visual experience during the crucial period caused a profound degradation, of both spatial tuning and topography, of auditory receptive fields in the SC. Additional data indicate that the SC auditory space map remains vulnerable to visual deprivation until at least 100 days after birth.