Carl H. Parsons

Training-induced plasticity of auditory localization in adult mammals

Accurate auditory localization relies on neural computations based on spatial cues present in the sound waves at each ear. The values of these cues depend on the size, shape, and separation of the two ears and can therefore vary from one individual to another. As with other perceptual skills, the neural circuits involved in spatial hearing are shaped by experience during development and retain some capacity for plasticity in later life. However, the factors that enable and promote plasticity of auditory localization in the adult brain are unknown. Here we show that mature ferrets can rapidly relearn to localize sounds after having their spatial cues altered by reversibly occluding one ear, but only if they are trained to use these cues in a behaviorally relevant task, with greater and more rapid improvement occurring with more frequent training. We also found that auditory adaptation is possible in the absence of vision or error feedback. Finally, we show that this process involves a shift in sensitivity away from the abnormal auditory spatial cues to other cues that are less affected by the earplug. The mature auditory system is therefore capable of adapting to abnormal spatial information by reweighting different localization cues. These results suggest that training should facilitate acclimatization to hearing aids in the hearing impaired.

The health promoting school in Europe: conceptualising and evaluating the change

This paper presents a holistic model of the health promoting school and a panoramic framework for evalution. The framework is board, acknowle dging the range of national settings in which health promoging schools are being developed, and draws attention to the emphasis on context and process rather than outcomes. The conceptual model is applied to the European Network of Health Promoting Schools (ENHPS), where the same forces may be at work in relation to the innovation but operate in different ways and with different degrees of strength. The paper recommends a selective approach to evaluation which will serve specific audiences and decision making needs.

Improved auditory spatial acuity in visually deprived ferrets

We have examined the effects on auditory spatial acuity in the horizontal plane of depriving ferrets of patterned visual cues by binocular eyelid suture in infancy or for a comparable period in adulthood. Minimum audible angles (MAAs) were measured for 500‐, 100‐ and 40‐ms broadband noise bursts at the midline and at 45° to one side. A logistic regression analysis revealed no consistent difference between the midline MAAs of normal and infant lid‐sutured ferrets. However, the lateral field MAAs of the infant‐deprived group were significantly smaller and showed less inter‐subject variability than those of normal‐sighted ferrets. The animals deprived in adulthood were tested in the lateral field only, firstly 6 months after binocular eyelid suture and again after a further 10 months. For the first test, the MAAs achieved by these animals with 500‐ and 100‐ms noise bursts were significantly smaller than the normal values and no different from those of the infant‐deprived group. A significant improvement in performance at the two shortest stimulus durations (100 and 40 ms) was observed when the adult‐deprived animals were re‐tested. Their second‐test MAAs did not differ from those of the infant‐deprived group at any of the three stimulus durations used, and both groups achieved significantly better scores than the normal‐sighted control animals. These results show that prolonged visual deprivation in both juvenile and adult ferrets can lead to a significant improvement in auditory spatial acuity in the lateral sound field. This is consistent with reports that congenitally blind humans can localize peripheral sounds more accurately than normal controls.

An investigation of the role of auditory cortex in sound localization using muscimol-releasing Elvax.

Lesion studies suggest that primary auditory cortex (A1) is required for accurate sound localization by carnivores and primates. In order to elucidate further its role in spatial hearing, we examined the behavioural consequences of reversibly inactivating ferret A1 over long periods, using Elvax implants releasing the GABAA receptor agonist muscimol. Sub‐dural polymer placements were shown to deliver relatively constant levels of muscimol to underlying cortex for >5 months. The measured diffusion of muscimol beneath and around the implant was limited to 1 mm. Cortical silencing was assessed electrophysiologically in both auditory and visual cortices. This exhibited rapid onset and was reversed within a few hours of implant removal. Inactivation of cortical neurons extended to all layers for implants lasting up to 6 weeks and throughout at least layers I–IV for longer placements, whereas thalamic activity in layer IV appeared to be unaffected. Blockade of cortical neurons in the deeper layers was restricted to ≤ 500 µm from the edge of the implant, but was usually more widespread in the superficial layers. In contrast, drug‐free Elvax implants had little discernible effect on the responses of the underlying cortical neurons. Bilateral implants of muscimol–Elvax over A1 produced significant deficits in the localization of brief sounds in horizontal space and particularly a reduced ability to discriminate between anterior and posterior sound sources. The performance of these ferrets gradually improved over the period in which the Elvax was in place and attained that of control animals following its removal. Although similar in nature, these deficits were less pronounced than those caused by cortical lesions and suggest a specific role for A1 in resolving the spatial ambiguities inherent in auditory localization cues.

Sound localization behavior in ferrets : comparison of acoustic orientation and approach-to-target responses

Auditory localization experiments typically either require subjects to judge the location of a sound source from a discrete set of response alternatives or involve measurements of the accuracy of orienting responses made toward the source location. To compare the results obtained by both methods, we trained ferrets by positive conditioning to stand on a platform at the center of a circular arena prior to stimulus presentation and then approach the source of a broadband noise burst delivered from 1 of 12 loudspeakers arranged at 30 degrees intervals in the horizontal plane. Animals were rewarded for making a correct choice. We also obtained a non-categorized measure of localization accuracy by recording head-orienting movements made during the first second following stimulus onset. The accuracy of the approach-to-target responses declined as the stimulus duration was reduced, particularly for lateral and posterior locations, although responses to sounds presented in the frontal region of space and directly behind the animal remained quite accurate. Head movements had a latency of approximately 200 ms and varied systematically in amplitude with stimulus direction. However, the final head bearing progressively undershot the target with increasing eccentricity and rarely exceeded 60 degrees to each side of the midline. In contrast to the approach-to-target responses, the accuracy of the head orienting responses did not change much with stimulus duration, suggesting that the improvement in percent correct scores with longer stimuli was due, at least in part, to re-sampling of the acoustical stimulus after the initial head turn had been made. Nevertheless, for incorrect trials, head orienting responses were more closely correlated with the direction approached by the animals than with the actual target direction, implying that at least part of the neural circuitry for translating sensory spatial signals into motor commands is shared by these two behaviors.

How plastic is spatial hearing

The location of a sound source is derived by the auditory system from spatial cues present in the signals at the two ears. These cues include interaural timing and level differences, as well as monaural spectral cues generated by the external ear. The values of these cues vary with individual differences in the shape and dimensions of the head and external ears. We have examined the neurophysiological consequences of these intersubject variations by recording the responses of neurons in ferret primary auditory cortex to virtual sound sources mimicking the animal’s own ears or those of other ferrets. For most neurons, the structure of the spatial response fields changed significantly when acoustic cues measured from another animal were presented. This is consistent with the finding that humans localize less accurately when listening to virtual sounds from other subjects. To examine the role of experience in shaping the ability to localize sound, we have studied the behavioural consequences of altering binaural cues by chronically plugging one ear. Ferrets raised and tested with one ear plugged learned to localize as accurately as control animals, which is consistent with previous findings that the representation of auditory space in the midbrain can accommodate abnormal sensory cues during development. Adaptive changes in behaviour were also observed in adults, particularly if they were provided with regular practice in the localization task. Together, these findings suggest that the neural circuits responsible for sound localization can be recalibrated throughout life.

Tracking the expression of excitatory and inhibitory neurotransmission-related proteins and neuroplasticity markers after noise induced hearing loss

Excessive exposure to loud noise can damage the cochlea and create a hearing loss. These pathologies coincide with a range of CNS changes including reorganisation of frequency representation, alterations in the pattern of spontaneous activity and changed expression of excitatory and inhibitory neurotransmitters. Moreover, damage to the cochlea is often accompanied by acoustic disorders such as hyperacusis and tinnitus, suggesting that one or more of these neuronal changes may be involved in these disorders, although the mechanisms remain unknown. We tested the hypothesis that excessive noise exposure increases expression of markers of excitation and plasticity, and decreases expression of inhibitory markers over a 32-day recovery period. Adult rats (n = 25) were monaurally exposed to a loud noise (16 kHz, 1/10th octave band pass (115 dB SPL)) for 1-hour, or left as non-exposed controls (n = 5). Animals were euthanased at either 0, 4, 8, 16 or 32 days following acoustic trauma. We used Western Blots to quantify protein levels of GABAA receptor subunit α1 (GABAAα1), Glutamic-Acid Decarboxylase-67 (GAD-67), N-Methyl-D-Aspartate receptor subunit 2A (NR2A), Calbindin (Calb1) and Growth Associated Protein 43 (GAP-43) in the Auditory Cortex (AC), Inferior Colliculus (IC) and Dorsal Cochlear Nucleus (DCN). Compared to sham-exposed controls, noise-exposed animals had significantly (p<0.05): lower levels of GABAAα1 in the contralateral AC at day-16 and day-32, lower levels of GAD-67 in the ipsilateral DCN at day-4, lower levels of Calb1 in the ipsilateral DCN at day-0, lower levels of GABAAα1 in the ipsilateral AC at day-4 and day-32. GAP-43 was reduced in the ipsilateral AC for the duration of the experiment. These complex fluctuations in protein expression suggests that for at least a month following acoustic trauma the auditory system is adapting to a new pattern of sensory input.

Auditory Learning as a Cause and Treatment of Central Dysfunction

Conductive hearing loss produced by middle ear disease (MED) is very prevalent in the first 5 years of childhood. Both MED in children and prolonged ear plugging in animals lead to a binaural hearing impairment that persists beyond the duration of the peripheral impairment. However, after cessation of the MED, or removal of the ear plug, binaural hearing gradually improves. We suggest here that this improvement is a passive form of auditory learning. We also show that active auditory learning, through repetition of discrimination tasks, can accelerate performance increments, both after hearing loss and in unimpaired individuals. A more detailed understanding of auditory learning holds out the prospect of improving rehabilitation strategies for the language- and hearing-impaired.

TVEI and the appeal of the really useful curriculum

Abstract The TVEI Pilot projects were particularly generously funded. For designated cohorts of students, covering the whole range of ability, there were novel opportunities for a work related curriculum taught in a more negotiated way which contrasted with both the content and the processes of the ‘traditional’ elitist curriculum. It was forced upon the education service in the name of the industrial lobby and served, inadvertently, the interests of less academic students. With the passing of TVEI we see the imposition, by a political/administrative alliance of the elitist content and subject matter of the National Curriculum.

The politics, ethics and diplomacy of reporting educational evaluation

Abstract To be effective in any formative sense, evaluation has to be received, acknowledged and acted upon. Numerous writers have dealt with the politics of evaluation and the ethics of evaluation. This paper introduces the notion of audience sensitivity and the ‘payability’ of the evaluation report. In particular it explores the tensions amongst the three demands for confidentiality, the right to know and palatability. A number of practical dilemmas, most drawn from experience, are presented to highlight the need for sensitive decision‐making and compromise if the potential beneficial impact of the evaluation is to be maximised.