Retrospective Valuation of Experienced Outcome Encoded in Distinct Reward Representations in the Anterior Insula and Amygdala.

Our ability to evaluate an experience retrospectively is important because it allows us to summarize its total value, and this summary value can then later be used as a guide in deciding whether the experience merits repeating, or whether instead it should rather be avoided. However, when an experience unfolds over time, humans tend to assign disproportionate weight to the later part of the experience, and this can lead to poor choice in repeating, or avoiding experience. Using model-based computational analyses of fMRI recordings in 27 male volunteers, we show that the human brain encodes the summary value of an extended sequence of outcomes in two distinct reward representations. We find that the overall experienced value is encoded accurately in the amygdala, but its merit is excessively marked down by disincentive anterior insula activity if the sequence of experienced outcomes declines temporarily. Moreover, the statistical strength of this neural code can separate efficient decision-makers from suboptimal decision-makers. Optimal decision-makers encode overall value more strongly, and suboptimal decision-makers encode the disincentive markdown (DM) more strongly. The separate neural implementation of the two distinct reward representations confirms that suboptimal choice for temporally extended outcomes can be the result of robust neural representation of a displeasing aspect of the experience such as temporary decline.SIGNIFICANCE STATEMENT One of the numerous foibles that prompt us to make poor decisions is known as the "Banker's fallacy," the tendency to focus on short-term growth at the expense of long-term value. This effect leads to unwarranted preference for happy endings. Here, we show that the anterior insula in the human brain marks down the overall value of an experience as it unfolds over time if the experience entails a sequence of predominantly negative temporal contrasts. By contrast, the amygdala encodes overall value accurately. These results provide neural indices for the dichotomy of decision utility and experienced utility popularized as Thinking fast and slow by Daniel Kahneman.

Dopamine Modulates Adaptive Prediction Error Coding in the Human Midbrain and Striatum.

Learning to optimally predict rewards requires agents to account for fluctuations in reward value. Recent work suggests that individuals can efficiently learn about variable rewards through adaptation of the learning rate, and coding of prediction errors relative to reward variability. Such adaptive coding has been linked to midbrain dopamine neurons in nonhuman primates, and evidence in support for a similar role of the dopaminergic system in humans is emerging from fMRI data. Here, we sought to investigate the effect of dopaminergic perturbations on adaptive prediction error coding in humans, using a between-subject, placebo-controlled pharmacological fMRI study with a dopaminergic agonist (bromocriptine) and antagonist (sulpiride). Participants performed a previously validated task in which they predicted the magnitude of upcoming rewards drawn from distributions with varying SDs. After each prediction, participants received a reward, yielding trial-by-trial prediction errors. Under placebo, we replicated previous observations of adaptive coding in the midbrain and ventral striatum. Treatment with sulpiride attenuated adaptive coding in both midbrain and ventral striatum, and was associated with a decrease in performance, whereas bromocriptine did not have a significant impact. Although we observed no differential effect of SD on performance between the groups, computational modeling suggested decreased behavioral adaptation in the sulpiride group. These results suggest that normal dopaminergic function is critical for adaptive prediction error coding, a key property of the brain thought to facilitate efficient learning in variable environments. Crucially, these results also offer potential insights for understanding the impact of disrupted dopamine function in mental illness.SIGNIFICANCE STATEMENT To choose optimally, we have to learn what to expect. Humans dampen learning when there is a great deal of variability in reward outcome, and two brain regions that are modulated by the brain chemical dopamine are sensitive to reward variability. Here, we aimed to directly relate dopamine to learning about variable rewards, and the neural encoding of associated teaching signals. We perturbed dopamine in healthy individuals using dopaminergic medication and asked them to predict variable rewards while we made brain scans. Dopamine perturbations impaired learning and the neural encoding of reward variability, thus establishing a direct link between dopamine and adaptation to reward variability. These results aid our understanding of clinical conditions associated with dopaminergic dysfunction, such as psychosis.

Imbalance in habitual versus goal directed neural systems during symptom provocation in obsessive-compulsive disorder.

Intrusive thoughts and compulsive urges to perform stereotyped behaviours are typical symptoms of obsessive-compulsive disorder. Emerging evidence suggests a cognitive bias towards habit formation at the expense of goal-directed performance in obsessive-compulsive disorder. In this study, we test this hypothesis using a novel individualized ecologically valid symptom provocation design: a live provocation functional magnetic resonance imaging paradigm with synchronous video-recording of behavioural avoidance responses. By pairing symptom provocation with online avoidance responses on a trial-by-trial basis, we sought to investigate the neural mechanisms leading to the compulsive avoidance response. In keeping with the model of habit formation in obsessive-compulsive disorder, we hypothesized that this disorder would be associated with lower activity in regions implicated in goal-directed behaviours and higher activity in regions implicated in habitual behaviours. Fifteen patients with obsessive-compulsive disorder and 15 healthy control volunteers participated in this functional magnetic resonance imaging study. Online stimuli were individually tailored to achieve effective symptom provocation at neutral, intermediate and strong intensity levels. During the symptom provocation block, the participant could choose to reject or terminate the provoking stimuli resulting in cessation of the symptom provocation. We thus separately analysed the neural correlates of symptom provocation, the urge to avoid, rejection and relief. Strongly symptom-provoking conditions evoked a dichotomous pattern of deactivation/activation in patients, which was not observed either in control conditions or in healthy subjects: a deactivation of caudate-prefrontal circuits accompanied by hyperactivation of subthalamic nucleus/putaminal regions. This finding suggests a dissociation between regions engaged in goal-directed and habitual behaviours. The putaminal hyperactivity during patients' symptom provocation preceded subsequent deactivation during avoidance and relief events, indicating a pivotal role of putamen in regulation of behaviour and habit formation in obsessive-compulsive disorder. Effective connectivity analysis identified the ventromedial prefrontal cortex/orbitofrontal cortex as the main structure in this circuitry involved in the modulation of compulsivity in obsessive-compulsive disorder. These findings suggest an imbalance in circuitry underlying habitual and goal-directed action control, which may represent a fundamental mechanism underlying compulsivity in obsessive-compulsive disorder. Our results complement current models of symptom generation in obsessive-compulsive disorder and may enable the development of future therapeutic approaches that aim to alleviate this imbalance.

Danish reading span data from 283 hearing-aid users, including a sub-group analysis of their relationship to speech-in-noise performance.

OBJECTIVE: This study provides descriptive statistics of the Danish reading span (RS) test for hearing-impaired adults. The combined effect of hearing loss, RS score, and age on speech-in-noise performance in different spatial settings was evaluated in a subset of participants. DESIGN: Data from published and unpublished studies were re-analysed. Data regarding speech-in-noise performance with co-located or spatially separated sound sources were available for a subset of participants. STUDY SAMPLE: RS scores from 283 hearing-impaired participants were extracted from past studies, and 239 of these participants had completed a speech-in-noise test. RESULTS: RS scores (mean = 41.91%, standard deviation = 11.29%) were related to age (p <0.01), but not pure-tone average (PTA) (p = 0.29). Speech-in-noise performance for co-located sound sources was related to PTA and RS score (both p < 0.01, adjusted R-squared = 0.226). Performance for spatially separated sounds was related to PTA (p < 0.01, adjusted R-squared = 0.10) but not RS score (p = 0.484). We found no differences between the standardized coefficients of the two regression models. CONCLUSIONS: The distribution of RS scores indicated a high test difficulty. We found that age should be controlled when RS scores are compared across populations. The experimental setup of the speech-in-noise test may influence the relationship between performance and RS score.

Dopamine modulates the neural representation of subjective value of food in hungry subjects.

Although there is a rich literature on the role of dopamine in value learning, much less is known about its role in using established value estimations to shape decision-making. Here we investigated the effect of dopaminergic modulation on value-based decision-making for food items in fasted healthy human participants. The Becker-deGroot-Marschak auction, which assesses subjective value, was examined in conjunction with pharmacological fMRI using a dopaminergic agonist and an antagonist. We found that dopamine enhanced the neural response to value in the inferior parietal gyrus/intraparietal sulcus, and that this effect predominated toward the end of the valuation process when an action was needed to record the value. Our results suggest that dopamine is involved in acting upon the decision, providing additional insight to the mechanisms underlying impaired decision-making in healthy individuals and clinical populations with reduced dopamine levels.

Choice mechanisms for past, temporally extended outcomes.

Accurate retrospection is critical in many decision scenarios ranging from investment banking to hedonic psychology. A notoriously difficult case is to integrate previously perceived values over the duration of an experience. Failure in retrospective evaluation leads to suboptimal outcome when previous experiences are under consideration for revisit. A biologically plausible mechanism underlying evaluation of temporally extended outcomes is leaky integration of evidence. The leaky integrator favours positive temporal contrasts, in turn leading to undue emphasis on recency. To investigate choice mechanisms underlying suboptimal outcome based on retrospective evaluation, we used computational and behavioural techniques to model choice between perceived extended outcomes with different temporal profiles. Second-price auctions served to establish the perceived values of virtual coins offered sequentially to humans in a rapid monetary gambling task. Results show that lesser-valued options involving successive growth were systematically preferred to better options with declining temporal profiles. The disadvantageous inclination towards persistent growth was mitigated in some individuals in whom a longer time constant of the leaky integrator resulted in fewer violations of dominance. These results demonstrate how focusing on immediate gains is less beneficial than considering longer perspectives.

The mutual roles of temporal glimpsing and vocal characteristics in cocktail-party listening.

At a cocktail party, listeners must attend selectively to a target speaker and segregate their speech from distracting speech sounds uttered by other speakers. To solve this task, listeners can draw on a variety of vocal, spatial, and temporal cues. Recently, Vestergaard et al. [J. Acoust. Soc. Am. 125, 1114-1124 (2009)] developed a concurrent-syllable task to control temporal glimpsing within segments of concurrent speech, and this allowed them to measure the interaction of glottal pulse rate and vocal tract length and reveal how the auditory system integrates information from independent acoustic modalities to enhance recognition. The current paper shows how the interaction of these acoustic cues evolves as the temporal overlap of syllables is varied. Temporal glimpses as short as 25 ms are observed to improve syllable recognition substantially when the target and distracter have similar vocal characteristics, but not when they are dissimilar. The effect of temporal glimpsing on recognition performance is strongly affected by the form of the syllable (consonant-vowel versus vowel-consonant), but it is independent of other phonetic features such as place and manner of articulation.

Evidence for early specialized processing of speech formant information in anterior and posterior human auditory cortex.

Many speech sounds, such as vowels, exhibit a characteristic pattern of spectral peaks, referred to as formants, the frequency positions of which depend both on the phonological identity of the sound (e.g. vowel type) and on the vocal-tract length of the speaker. This study investigates the processing of formant information relating to vowel type and vocal-tract length in human auditory cortex by measuring electroencephalographic (EEG) responses to synthetic unvoiced vowels and spectrally matched noises. The results revealed specific sensitivity to vowel formant information in both anterior (planum polare) and posterior (planum temporale) regions of auditory cortex. The vowel-specific responses in these two areas appeared to have different temporal dynamics; the anterior source produced a sustained response for as long as the incoming sound was a vowel, whereas the posterior source responded transiently when the sound changed from a noise to a vowel, or when there was a change in vowel type. Moreover, the posterior source appeared to be largely invariant to changes in vocal-tract length. The current findings indicate that the initial extraction of vowel type from formant information is complete by the level of non-primary auditory cortex, suggesting that speech-specific processing may involve primary auditory cortex, or even subcortical structures. This challenges the view that specific sensitivity to speech emerges only beyond unimodal auditory cortex.

Location and acoustic scale cues in concurrent speech recognition.

Location and acoustic scale cues have both been shown to have an effect on the recognition of speech in multi-speaker environments. This study examines the interaction of these variables. Subjects were presented with concurrent triplets of syllables from a target voice and a distracting voice, and asked to recognize a specific target syllable. The task was made more or less difficult by changing (a) the location of the distracting speaker, (b) the scale difference between the two speakers, and/or (c) the relative level of the two speakers. Scale differences were produced by changing the vocal tract length and glottal pulse rate during syllable synthesis: 32 acoustic scale differences were used. Location cues were produced by convolving head-related transfer functions with the stimulus. The angle between the target speaker and the distracter was 0 degrees, 4 degrees, 8 degrees, 16 degrees, or 32 degrees on the 0 degrees horizontal plane. The relative level of the target to the distracter was 0 or -6 dB. The results show that location and scale difference interact, and the interaction is greatest when one of these cues is small. Increasing either the acoustic scale or the angle between target and distracter speakers quickly elevates performance to ceiling levels.

Effects of voicing in the recognition of concurrent syllables.

This letter reports a study designed to measure the benefits of voicing in the recognition of concurrent syllables. The target and distracter syllables were either voiced or whispered, producing four combinations of vocal contrast. Results show that listeners use voicing whenever it is present either to detect a target syllable or to reject a distracter. When the predictable effects of audibility were taken into account, limited evidence remained for the harmonic cancellation mechanism thought to make rejecting distracter syllables more effective than enhancing target syllables.

The interaction of vocal characteristics and audibility in the recognition of concurrent syllables.

In concurrent-speech recognition, performance is enhanced when either the glottal pulse rate (GPR) or the vocal tract length (VTL) of the target speaker differs from that of the distracter, but relatively little is known about the trading relationship between the two variables, or how they interact with other cues such as signal-to-noise ratio (SNR). This paper presents a study in which listeners were asked to identify a target syllable in the presence of a distracter syllable, with carefully matched temporal envelopes. The syllables varied in GPR and VTL over a large range, and they were presented at different SNRs. The results showed that performance is particularly sensitive to the combination of GPR and VTL when the SNR is 0 dB. Equal-performance contours showed that when there are no other cues, a two-semitone difference in GPR produced the same advantage in performance as a 20% difference in VTL. This corresponds to a trading relationship between GPR and VTL of 1.6. The results illustrate that the auditory system can use any combination of differences in GPR, VTL, and SNR to segregate competing speech signals.

Adaptive Prediction Error Coding in the Human Midbrain and Striatum Facilitates Behavioral Adaptation and Learning Efficiency.

Effective error-driven learning benefits from scaling of prediction errors to reward variability. Such behavioral adaptation may be facilitated by neurons coding prediction errors relative to the standard deviation (SD) of reward distributions. To investigate this hypothesis, we required participants to predict the magnitude of upcoming reward drawn from distributions with different SDs. After each prediction, participants received a reward, yielding trial-by-trial prediction errors. In line with the notion of adaptive coding, BOLD response slopes in the Substantia Nigra/Ventral Tegmental Area (SN/VTA) and ventral striatum were steeper for prediction errors occurring in distributions with smaller SDs. SN/VTA adaptation was not instantaneous but developed across trials. Adaptive prediction error coding was paralleled by behavioral adaptation, as reflected by SD-dependent changes in learning rate. Crucially, increased SN/VTA and ventral striatal adaptation was related to improved task performance. These results suggest that adaptive coding facilitates behavioral adaptation and supports efficient learning.

Evidence accumulation in obsessive-compulsive disorder: the role of uncertainty and monetary reward on perceptual decision-making thresholds.

The compulsive behaviour underlying obsessive-compulsive disorder (OCD) may be related to abnormalities in decision-making. The inability to commit to ultimate decisions, for example, patients unable to decide whether their hands are sufficiently clean, may reflect failures in accumulating sufficient evidence before a decision. Here we investigate the process of evidence accumulation in OCD in perceptual discrimination, hypothesizing enhanced evidence accumulation relative to healthy volunteers. Twenty-eight OCD patients and thirty-five controls were tested with a low-level visual perceptual task (random-dot-motion task, RDMT) and two response conflict control tasks. Regression analysis across different motion coherence levels and Hierarchical Drift Diffusion Modelling (HDDM) were used to characterize response strategies between groups in the RDMT. Patients required more evidence under high uncertainty perceptual contexts, as indexed by longer response time and higher decision boundaries. HDDM, which defines a decision when accumulated noisy evidence reaches a decision boundary, further showed slower drift rate towards the decision boundary reflecting poorer quality of evidence entering the decision process in patients under low uncertainty. With monetary incentives emphasizing speed and penalty for slower responses, patients decreased the decision thresholds relative to controls, accumulating less evidence in low uncertainty. These findings were unrelated to visual perceptual deficits and response conflict. This study provides evidence for impaired decision-formation processes in OCD, with a differential influence of high and low uncertainty contexts on evidence accumulation (decision threshold) and on the quality of evidence gathered (drift rates). It further emphasizes that OCD patients are sensitive to monetary incentives heightening speed in the speed-accuracy tradeoff, improving evidence accumulation.

Progressive associative phonagnosia: a neuropsychological analysis.

There are few detailed studies of impaired voice recognition, or phonagnosia. Here we describe two patients with progressive phonagnosia in the context of frontotemporal lobar degeneration. Patient QR presented with behavioural decline and increasing difficulty recognising familiar voices, while patient KL presented with progressive prosopagnosia. In a series of neuropsychological experiments we assessed the ability of QR and KL to recognise and judge the familiarity of voices, faces and proper names, to recognise vocal emotions, to perceive and discriminate voices, and to recognise environmental sounds and musical instruments. The patients were assessed in relation to a group of healthy age-matched control subjects. QR exhibited severe impairments of voice identification and familiarity judgments with relatively preserved recognition of difficulty-matched faces and environmental sounds; recognition of musical instruments was impaired, though better than recognition of voices. In contrast, patient KL exhibited severe impairments of both voice and face recognition, with relatively preserved recognition of musical instruments and environmental sounds. Both patients demonstrated preserved ability to analyse perceptual properties of voices and to recognise vocal emotions. The voice processing deficit in both patients could be characterised as associative phonagnosia: in the case of QR, this was relatively selective for voices, while in the case of KL, there was evidence for a multimodal impairment of person knowledge. The findings have implications for current cognitive models of voice recognition.

Timbre-independent extraction of pitch in newborn infants.

The ability to separate pitch from other spectral sound features, such as timbre, is an important prerequisite of veridical auditory perception underlying speech acquisition and music cognition. The current study investigated whether or not newborn infants generalize pitch across different timbres. Perceived resonator size is an aspect of timbre that informs the listener about the size of the sound source, a cue that may be important already at birth. Therefore, detection of infrequent pitch changes was tested by recording event-related brain potentials in healthy newborn infants to frequent standard and infrequent pitch-deviant sounds while the perceived resonator size of all sounds was randomly varied. The elicitation of an early negative and a later positive discriminative response by deviant sounds demonstrated that the neonate auditory system represents pitch separately from timbre, thus showing advanced pitch processing capabilities.

Auditory size-deviant detection in adults and newborn infants.

Auditory size perception refers to the ability to make accurate judgements of the size of a sound source based solely upon the sound emitted from the source. Electro-physiological and behavioural data were collected to test whether sound-source size parameters are detected from task-irrelevant sequences in adults and newborn infants. The mismatch negativity (MMN) obtained from adults indexed automatic detection of changes in size for voices, musical instruments and animal calls, regardless of whether the acoustic change indicated larger or smaller sources. Neonates detected changes in the size of a musical instrument. The data are consistent with the notion that auditory size-deviant detection in humans is an innate automatic process. This conclusion is compatible with the theory that the ability to assess the size of sound sources evolved because it provided selective advantage of being able to detect larger (more competent) suitors and larger (more dangerous) predators.

Self-report outcome in new hearing-aid users: Longitudinal trends and relationships between subjective measures of benefit and satisfaction.

This study focussed on self-report outcome in new hearing-aid users. The objectives of the experiment were changes in self-report outcome over time, and relationships between different subjective measures of benefit and satisfaction. Four outcome inventories and a questionnaire on auditory lifestyle were administered to 25 hearing-aid users repeatedly after hearing-aid fitting, and assessments took place one week, four weeks, and 13 weeks after hearing-aid provision. The results showed that, for first-time users who used their hearing aids more than four hours per day, self-reported outcome increased over 13 weeks in some scales, although there was no change in amplification during this time. Furthermore, it was found that, for data collected immediately post-fitting, some subscales were much less face valid than for data collected later. This result indicates that the way in which hearing-aid users assess outcome changes over time. The practical consequence of the results is that early self-report outcome assessment may be misleading for some self-report outcome schemes.

Dead regions in the cochlea: implications for speech recognition and applicability of articulation index theory.

Dead regions in the cochlea have been suggested to be responsible for failure by hearing aid users to benefit from apparently increased audibility in terms of speech intelligibility. As an alternative to the more cumbersome psychoacoustic tuning curve measurement, threshold-equalizing noise (TEN) has been reported to enable diagnosis of dead regions. The purpose of the present study was first to assess the feasibility of the TEN test protocol, and second, to assess the ability of the procedure to reveal related functional impairment. The latter was done by a test for the recognition of low-pass-filtered speech items. Data were collected from 22 hearing-impaired subjects with moderate-to-profound sensorineural hearing losses. The results showed that 11 subjects exhibited abnormal psychoacoustic behaviour in the TEN test, indicative of a possible dead region. Estimates of audibility were used to assess the possible connection between dead-region candidacy and ability to recognize low-pass-filtered speech. Large variability was observed with regard to the ability of audibility to predict recognition scores for both dead-region and no-dead-region subjects. Furthermore, the results indicate that dead-region subjects might be better than no-dead-region subjects at recognizing speech of marginal audibility.