PsySuite: An android application designed to perform multimodal psychophysical testing.

In behavioral sciences, there is growing concern about the inflation of false-positive rates due to the amount of under-powered studies that have been shared in the past years. While problematic, having the possibility to recruit (lots of) participants (for a lot of time) is realistically not achievable for many research facilities. Factors that hinder the reaching of optimal sample sizes are, to name but a few, research costs, participants' availability and commitment, and logistics. We challenge these issues by introducing PsySuite, an Android app designed to foster a remote approach to multimodal behavioral testing. To validate PsySuite, we first evaluated its ability to generate stimuli appropriate to rigorous psychophysical testing, measuring both the app's accuracy (i.e., stimuli's onset, offset, and multimodal simultaneity) and precision (i.e., the stability of a given pattern across trials), using two different smartphone models. We then evaluated PsySuite's ability to replicate perceptual performances obtained using a classic psychophysical paradigm, comparing sample data collected with the app against those measured via a PC-based setup. Our results showed that PsySuite could accurately reproduce stimuli with a minimum duration of 7 ms, 17 ms, and 30 ms for the auditory, visual, and tactile modalities, respectively, and that perceptual performances obtained with PsySuite were consistent with the perceptual behavior observed using the classical setup. Combined with the high accessibility inherently supported by PsySuite, here we ought to share the app to further boost psychophysical research, aiming at setting it to a cheap, user-friendly, and portable level.

A deep learning based cognitive model to probe the relation between psychophysics and electrophysiology of flicker stimulus.

The flicker stimulus is a visual stimulus of intermittent illumination. A flicker stimulus can appear flickering or steady to a human subject, depending on the physical parameters associated with the stimulus. When the flickering light appears steady, flicker fusion is said to have occurred. This work aims to bridge the gap between the psychophysics of flicker fusion and the electrophysiology associated with flicker stimulus through a Deep Learning based computational model of flicker perception. Convolutional Recurrent Neural Networks (CRNNs) were trained with psychophysics data of flicker stimulus obtained from a human subject. We claim that many of the reported features of electrophysiology of the flicker stimulus, including the presence of fundamentals and harmonics of the stimulus, can be explained as the result of a temporal convolution operation on the flicker stimulus. We further show that the convolution layer output of a CRNN trained with psychophysics data is more responsive to specific frequencies as in human EEG response to flicker, and the convolution layer of a trained CRNN can give a nearly sinusoidal output for 10 hertz flicker stimulus as reported for some human subjects.

Low-cost, portable, easy-to-use kiosks to facilitate home-cage testing of non-human primates during vision-based behavioral tasks.

Non-human primates (NHPs), especially rhesus macaques, have significantly contributed to our understanding of the neural computations underlying human vision. Besides the established homologies in the visual brain areas between these species and our ability to probe detailed neural mechanisms in monkeys at multiple scales, NHPs' ability to perform human-like visual behavior makes them an extremely appealing animal model of human vision. Traditionally, such behavioral studies have been conducted in controlled laboratory settings, offering experimenters tight control over variables like luminance, eye movements, and auditory interference. However, in-lab experiments have several constraints, including limited experimental time, the need for dedicated human experimenters, additional lab space requirements, invasive surgeries for head-post implants, and extra time and training for chairing and head restraints. To overcome these limitations, we propose adopting home-cage behavioral training and testing of NHPs, enabling the administration of many vision-based behavioral tasks simultaneously across multiple monkeys with reduced human personnel requirements, no NHP head restraint, and monkeys' unrestricted access to experiments. In this article, we present a portable, low-cost, easy-to-use kiosk system developed to conduct home-cage vision-based behavioral tasks in NHPs. We provide details of its operation and build to enable more open-source development of this technology. Furthermore, we present validation results using behavioral measurements performed in the lab and in NHP home cages, demonstrating the system's reliability and potential to enhance the efficiency and flexibility of NHP behavioral research.

Texture perception at the foot sole: Comparison between walking, sitting, and to the hand.

We frequently interact with textured surfaces with both our feet and hands. Like texture's importance for grasping, texture perception via the foot sole might provide important signals about the stability of a surface, aiding in maintaining balance. However, how textures are perceived by the foot, and especially under the high forces experienced during walking, is unknown. The current study builds on extensive research investigating texture perception at the hand by presenting everyday textures to the foot while stepping onto them, exploring them with the foot while sitting, and exploring them with the hand. Participants rated each texture along three perceptual dimensions: roughness, hardness, and stickiness. Participants also rated how stable their posture felt when standing upon each texture. Results show that perceptual ratings of each textural dimension were highly correlated across conditions. Hardness exhibited the greatest consistency and stickiness the weakest. Moreover, correlations between stepping and exploration with the foot were lower than those between exploration with the foot and exploration with the hand, suggesting that mode of interaction (high vs low force) impacts perception more than body region used (foot vs hand). On an individual level, correlations between conditions were higher than those between participants, suggesting that differences are greater between individuals than between mode of interaction or body region. When investigating the relationship to perceived stability, only hardness contributed significantly, with harder surfaces rated as more stable. Overall, tactile perception appears consistent across body regions and interaction modes, although differences in perception are greater during walking.

Expert CONsensus on Visual Evaluation in Retinal disease manaGEment: the CONVERGE study.

BACKGROUND/AIMS: Recent decades have seen significant advances in both structural and functional testing of retinal disease. However, the current clinical value of specific testing modalities, as well as future trends, need to be clearly identified in order to highlight areas for further development in routine care and clinical trials. METHODS: We designed a modified two-round Delphi study to obtain the opinion of a multidisciplinary group of 33 international experts involved in the field of retinal disease management/research to determine the level of agreement and consensus regarding the value and performance of specific structural and functional testing methods for retinal disease. On a Likert scale, a median of 1-2 indicated disagreement with the statement, and 5-6 indicated agreement with the statement. An IQR of ≤2 indicated consensus in the responses. Several questions also allowed comments on responses. RESULTS: There was overall agreement that structural testing currently predominates for detection and monitoring. There was moderate agreement that functional testing remains important and will continue to do so in the future because it provides complementary information. Certain respondents considered that properly designed and applied psychophysical tests are as reliable and repeatable as structural observations and that functional changes are the most important in the long run. Respondents considered future care and research to require a combination of structural and functional testing with strong consensus that the relative importance will depend on disease type and stage. CONCLUSION: The study obtained important insights from a group of international experts regarding current and future needs in the management of retinal disease using a mix of quantitative and qualitative approaches. Responses provide a rich range of opinions that will be of interest to researchers seeking to design tests for future patient care and clinical trials.

fMRI and gene therapy in adults with CNGB3 mutation.

Achromatopsia is an inherited retinal disease that affects 1 in 30,000-50,000 individuals and is characterised by an absence of functioning cone photoreceptors from birth. This results in severely reduced visual acuity, no colour vision, marked sensitivity to light and involuntary oscillations of the eyes (nystagmus). In most cases, a single gene mutation prevents normal development of cone photoreceptors, with mutations in the CNGB3 or CNGA3 gene being responsible for ∼80 % of all patients with achromatopsia. There are a growing number of studies investigating recovery of cone function after targeted gene therapy. These studies have provided some promise for patients with the CNGA3 mutation, but thus far have found limited or no recovery for patients with the CNGB3 mutation. Here, we developed colour-calibrated visual stimuli designed to isolate cone photoreceptor responses. We combined these with adapted fMRI techniques and pRF mapping to identify if cortical responses to cone-driven signals could be detected in 9 adult patients with the CNGB3 mutation after receiving gene therapy. We did not detect any change in brain activity after gene therapy when the 9 patients were analysed as a group. However, on an individual basis, one patient self-reported a change in colour perception, corroborated by improved performance on a psychophysical task designed to selectively identify cone function. This suggests a level of cone sensitivity that was lacking pre-treatment, further supported by a subtle but reliable change in cortical activity within their primary visual cortex.

Standard models of spatial vision mispredict edge sensitivity at low spatial frequencies.

One well-established characteristic of early visual processing is the contrast sensitivity function (CSF) which describes how sensitivity varies with the spatial frequency (SF) content of the visual input. The CSF prompted the development of a now standard model of spatial vision. It represents the visual input by activity in orientation- and SF selective channels which are nonlinearly recombined to predict a perceptual decision. The standard spatial vision model has been extensively tested with sinusoidal gratings at low contrast because their narrow SF spectra isolate the underlying SF selective mechanisms. It is less studied how well these mechanisms account for sensitivity to more behaviourally relevant stimuli such as sharp edges at high contrast (i.e. object boundaries) which abound in the natural environment and have broader SF spectra. Here, we probe sensitivity to edges (2-AFC, edge localization) in the presence of broadband and narrowband noises. We use Cornsweet luminance profiles with peak frequencies at 0.5, 3 and 9 cpd as edge stimuli. To test how well mechanisms underlying sinusoidal contrast sensitivity can account for edge sensitivity, we implement a single- and a multi-scale model building upon standard spatial vision model components. Both models account for most of the data but also systematically deviate in their predictions, particularly in the presence of pink noise and for the lowest SF edge. These deviations might indicate a transition from contrast- to luminance-based detection at low SFs. Alternatively, they might point to a missing component in current spatial vision models.

One mechanism of sudden sensorineural hearing loss after sildenafil and sexual activity.

OBJECTIVES: A case of sudden sensorineural hearing loss following use of sildenafil was examined in detail over a period of three days from first report to recovery. DESIGN: Case study. The subject presented with sudden sensorineural hearing loss and diplacusis a day after onset. Testing involved detailed interview, standard audiometry, detailed inter-octave audiometry, and measurement of detailed psychophysical frequency tuning curves during a two day recovery period. STUDY SAMPLE: One male aged in his thirties with otherwise normal hearing. RESULTS: Although standard audiometry was within normal limits, detailed inter-octave audiometry and psychophysical frequency tuning curves were consistent with a punctate unilateral intra-cochlear lesion that resolved over a period of three days. CONCLUSIONS: This is the first report of such a frequency-specific audiometric shift and diplacusis after sildenafil, and is not consistent with previous reports of direct ototoxic pharmacological effects. We propose that the lesion was most likely caused by a cochlear bleed, and may have been due to physical exertion rather than a direct pharmaceutical effect. The study highlights the important role of additional diagnostic testing that can be easily achieved in a clinical setting with minimal equipment.

The Relation Between Subjective and Objective Measures of Visual Awareness: Current Evidence, Attempt of a Synthesis and Future Research Directions.

Within the realm of consciousness research, different methods of measuring the content of visual awareness are used: On the one hand, subjective measures require a report of sensory experiences related to a stimulus. On the other hand, objective measures rely on the observer's performance to accurately detect or discriminate the stimulus. The most appropriate measure of awareness is currently debated. To contribute to this debate, we review findings on the relation between subjective and objective measures of awareness. Although subjective measures sometimes lag behind objective measures, a substantial number of studies demonstrates a convergence of measures. Based on the reviewed studies, we identify five aspects relevant for achieving a convergence of measures. Future research could then identify and empirically test the boundary conditions, under which a convergence or divergence of subjective and measures of awareness is observed.

Developmentally stable representations of naturalistic image structure in macaque visual cortex.

To determine whether post-natal improvements in form vision result from changes in mid-level visual cortex, we studied neuronal and behavioral responses to texture stimuli that were matched in local spectral content but varied in "naturalistic" structure. We made longitudinal measurements of visual behavior from 16 to 95 weeks of age, and of neural responses from 20 to 56 weeks. We also measured behavioral and neural responses in near-adult animals more than 3 years old. Behavioral sensitivity reached half-maximum around 25 weeks of age, but neural sensitivities remained stable through all ages tested. Neural sensitivity to naturalistic structure was highest in V4, lower in V2 and inferotemporal cortex (IT), and barely discernible in V1. Our results show a dissociation between stable neural performance and improving behavioral performance, which may reflect improved processing capacity in circuits downstream of visual cortex.

Haptic based fundamentals of laparoscopic surgery simulation for training with objective assessments.

Force is crucial for learning psychomotor skills in laparoscopic tissue manipulation. Fundamental laparoscopic surgery (FLS), on the other hand, only measures time and position accuracy. FLS is a commonly used training program for basic laparoscopic training through part tasks. The FLS is employed in most of the laparoscopic training systems, including box trainers and virtual reality (VR) simulators. However, many laparoscopic VR simulators lack force feedback and measure tissue damage solely through visual feedback based on virtual collisions. Few VR simulators that provide force feedback have subjective force metrics. To provide an objective force assessment for haptic skills training in the VR simulators, we extend the FLS part tasks to haptic-based FLS (HFLS), focusing on controlled force exertion. We interface the simulated HFLS part tasks with a customized bi-manual haptic simulator that offers five degrees of freedom (DOF) for force feedback. The proposed tasks are evaluated through face and content validity among laparoscopic surgeons of varying experience levels. The results show that trainees perform better in HFLS tasks. The average Likert score observed for face and content validity is greater than 4.6 ± 0.3 and 4 ± 0.5 for all the part tasks, which indicates the acceptance of the simulator among subjects for its appearance and functionality. Face and content validations show the need to improve haptic realism, which is also observed in existing simulators. To enhance the accuracy of force rendering, we incorporated a laparoscopic tool force model into the simulation. We study the effectiveness of the model through a psychophysical study that measures just noticeable difference (JND) for the laparoscopic gripping task. The study reveals an insignificant decrease in gripping-force JND. A simple linear model could be sufficient for gripper force feedback, and a non-linear LapTool force model does not affect the force perception for the force range of 0.5-2.5 N. Further study is required to understand the usability of the force model in laparoscopic training at a higher force range. Additionally, the construct validity of HFLS will confirm the applicability of the developed simulator to train surgeons with different levels of experience.

Multiplexing intensity and frequency sensations for artificial touch by modulating temporal features of electrical pulse trains.

In neural prostheses, intensity modulation of a single channel (i.e., through a single stimulating electrode) has been achieved by increasing the magnitude or width of each stimulation pulse, which risks eliciting pain or paraesthesia; and by changing the stimulation rate, which leads to concurrent changes in perceived frequency. In this study, we sought to render a perception of tactile intensity and frequency independently, by means of temporal pulse train patterns of fixed magnitude, delivered non-invasively. Our psychophysical study exploits a previously discovered frequency coding mechanism, where the perceived frequency of stimulus pulses grouped into periodic bursts depends on the duration of the inter-burst interval, rather than the mean pulse rate or periodicity. When electrical stimulus pulses were organised into bursts, perceived intensity was influenced by the number of pulses within a burst, while perceived frequency was determined by the time between the end of one burst envelope and the start of the next. The perceived amplitude was modulated by 1.6× while perceived frequency was varied independently by 2× within the tested range (20-40 Hz). Thus, the sensation of intensity might be controlled independently from frequency through a single stimulation channel without having to vary the injected electrical current. This can form the basis for improving strategies in delivering more complex and natural sensations for prosthetic hand users.

Epistemic circularity and measurement validity in quantitative psychology: insights from Fechner's psychophysics.

The validity of psychological measurement is crucially connected to a peculiar form of epistemic circularity. This circularity can be a threat when there are no independent ways to assess whether a certain procedure is actually measuring the intended target of measurement. This paper focuses on how Fechner addressed the measurement circularity that emerged in his psychophysical research. First, I show that Fechner's approach to the problem of circular measurement involved a core idealizing assumption of a shared human physiology. Second, I assess Fechner's approach to this issue against the backdrop of his own epistemology of measurement and the measurement context of his time. Third, I claim that, from a coherentist and historically-situated perspective, Fechner's quantification can be regarded as a first successful step of a longer-term quantification process. To conclude, I draw from these insights some general epistemological reflections that are relevant to current quantitative psychology.

Visuotactile integration in individuals with fibromyalgia.

Our brain constantly integrates afferent information, such as visual and tactile information, to perceive the world around us. According to the maximum-likelihood estimation (MLE) model, imprecise information will be weighted less than precise, making the multisensory percept as precise as possible. Individuals with fibromyalgia (FM), a chronic pain syndrome, show alterations in the integration of tactile information. This could lead to a decrease in their weight in a multisensory percept or a general disruption of multisensory integration, making it less beneficial. To assess multisensory integration, 15 participants with FM and 18 pain-free controls performed a temporal-order judgment task in which they received pairs of sequential visual, tactile (unisensory conditions), or visuotactile (multisensory condition) stimulations on the index and the thumb of the non-dominant hand and had to determine which finger was stimulated first. The task enabled us to measure the precision and accuracy of the percept in each condition. Results indicate an increase in precision in the visuotactile condition compared to the unimodal conditions in controls only, although we found no intergroup differences. The observed visuotactile precision was correlated to the precision predicted by the MLE model in both groups, suggesting an optimal integration. Finally, the weights of the sensory information were not different between the groups; however, in the group with FM, higher pain intensity was associated with smaller tactile weight. This study shows no alterations of the visuotactile integration in individuals with FM, though pain may influence tactile weight in these participants.

The precision of signals encoding active self-movement.

Everyday actions like moving the head, walking around, and grasping objects are typically self-controlled. This presents a problem when studying the signals encoding such actions because active self-movement is difficult to control experimentally. Available techniques demand repeatable trials, but each action is unique, making it difficult to measure fundamental properties like psychophysical thresholds. We present a novel paradigm that recovers both precision and bias of self-movement signals with minimal constraint on the participant. The paradigm relies on linking image motion to previous self-movement, and two experimental phases to extract the signal encoding the latter. The paradigm takes care of a hidden source of external noise not previously accounted for in techniques that link display motion to self-movement in real time (e.g., virtual reality). We use head rotations as an example of self-movement, and show that the precision of the signals encoding head movement depends on whether they are being used to judge visual motion or auditory motion. We find that perceived motion is slowed during head movement in both cases. The "nonimage" signals encoding active head rotation (motor commands, proprioception, and vestibular cues) are therefore biased toward lower speeds and/or displacements. In a second experiment, we trained participants to rotate their heads at different rates and found that the imprecision of the head rotation signal rises proportionally with head speed (Weber's law). We discuss the findings in terms of the different motion cues used by vision and hearing, and the implications they have for Bayesian models of motion perception.NEW & NOTEWORTHY We present a psychophysical technique for measuring the precision of signals encoding active self-movements. Using head movements, we show that 1) precision is greater when active head rotation is performed using visual comparison stimuli versus auditory; 2) precision decreases with head speed (Weber's law); 3) perceived speed is lower during head rotation. The findings may reflect the steps needed to convert different cues into common units, and challenge standard Bayesian models of motion perception.

The association between olfactory subdomains and frailty: A prospective case‒control study investigation.

BACKGROUND: Amidst the rise of frailty among a globally aging population, olfactory decline has emerged as a harbinger of frailty and mortality in population-level studies. However, the relationships between frailty and the olfactory subdomains of identification (OI), discrimination (OD), and threshold (OT) remain unexplored. This study prospectively examined the association between olfactory subdomains and the physical frailty phenotype (PFP) to investigate olfactory evaluation as a means of frailty screening. METHODS: A case‒control study of 45 frail and 45 non-frail individuals matched by age and sex. OT, OD, OI (range 0‒16), and composite sum (threshold, discrimination, and identification scores [TDI], range 0‒48) were measured with Sniffin' Sticks. PFP was defined by presence of three or more criteria: physical inactivity, self-reported exhaustion, muscle weakness, slow gait, and unintentional weight loss. Conditional logistic regression evaluated associations between olfactory subdomains and frailty. RESULTS: Ninety individuals with mean age of 83.1 ± 4.9 years, 60% female (n = 54), and 87.8% white (n = 79) were included. Olfactory scores were significantly lower in the frail group for OI (9.2 vs. 12.1, p < 0.001), OD (8.1 vs. 11.6, p < 0.001), OT (4.4 vs. 8.5, p < 0.001), and TDI (21.7 vs. 32.2, p < 0.001) than in the non-frail group. A single-point decrease in olfactory score was associated with increased odds of frailty in OT (odds ratio [OR]: 2.21, 95% confidence interval: [1.22, 3.98]), OD (OR: 2.19, 95% CI: [1.32, 3.65]), OI (OR: 2.29, 95% CI: [1.19, 4.39]), and TDI (OR: 1.54, 95% CI: [1.14, 2.08]). CONCLUSION: The robust association between olfactory subdomain scores and frailty suggests that olfaction may be an accessible signifier of frailty. Future studies should investigate this relationship longitudinally to assess predictive relationships.

The Processing of Short Time Intervals: Some Critical Issues.

Humans have the capability to make judgments about the relative duration of time intervals with accuracy (correct perceived duration) and precision (low variability). However, this capability has limitations, some of which are discussed in the present chapter. These limitations, either in terms of accuracy or precision, are obvious when there are changes in the physical characteristics of the stimuli used to mark the intervals to be judged. The characteristics are the structure (filled vs. empty) of the intervals and the sensory origin of the stimuli used to mark them. The variability of time estimates also depends on the use of single intervals by opposition to the use of sequences of intervals, and on the duration range under investigation. In addition to the effect caused by the physical characteristics of the stimuli, the perceived duration also relies on the way of presenting successive stimuli and on whether the intervals are marked by a single source or by different sources with distance (spatial effect) between them.

PSYCHOACOUSTICS-WEB: A free online tool for the estimation of auditory thresholds.

PSYCHOACOUSTICS-WEB is an online tool written in JavaScript and PHP that enables the estimation of auditory sensory thresholds via adaptive threshold tracking. The toolbox implements the transformed up-down methods proposed by Levitt (Journal of the Acoustical Society of America, 49, 467-477, (1971) for a set of classic psychoacoustical tasks: frequency, intensity, and duration discrimination of pure tones; duration discrimination and gap detection of noise; and amplitude modulation detection with noise carriers. The toolbox can be used through a common web browser; it works with both fixed and mobile devices, and requires no programming skills. PSYCHOACOUSTICS-WEB is suitable for laboratory, classroom, and online testing and is designed for two main types of users: an occasional user and, above all, an experimenter using the toolbox for their own research. This latter user can create a personal account, customise existing experiments, and share them in the form of direct links to further users (e.g., the participants of a hypothetical experiment). Finally, because data storage is centralised, the toolbox offers the potential for creating a database of auditory skills.

Predicting lever press in a vibrotactile yes/no detection task from S1 cortex of freely behaving rats by µECoG arrays.

AIM OF THE STUDY: Brain-computer interfaces (BCIs) may help patients with severe neurological deficits communicate with the external world. Based on microelectrocorticography (µECoG) data recorded from the primary somatosensory cortex (S1) of unrestrained behaving rats, this study attempts to decode lever presses in a psychophysical detection task by using machine learning algorithms. MATERIALS AND METHODS: 16-channel Pt-Ir microelectrode arrays were implanted on the S1 of two rats, and µECoG was recorded during a vibrotactile yes/no detection task. For this task, the rats were trained to press the right lever when they detected the vibrotactile stimulus and the left lever when they did not. The multichannel µECoG data was analysed offline by time-frequency methods and its features were used for binary classification of the lever press at each trial. Several machine learning algorithms were tested as such. RESULTS: The psychophysical sensitivities (A') were similar and low for both rats (0.58). Rat 2 (B'': -0.11) had higher bias for the right lever than Rat 1 (B'': - 0.01). The lever presses could be predicted with accuracies over 66% with all the tested algorithms, and the highest average accuracy (78%) was with the support vector machine. CONCLUSION: According to the recent studies, sensory feedback increases the benefit of the BCIs. The current proof-of-concept study shows that lever presses can be decoded from the S1; therefore, this area may be utilised for a bidirectional BCI in the future.

Transcranial Alternating Current Stimulation over Frontal Eye Fields Mimics Attentional Modulation of Visual Processing.

Attentional control over sensory processing has been linked to neural alpha oscillations and related inhibition of cerebral cortex. Despite the wide consensus on the functional relevance of alpha oscillations for attention, precise neural mechanisms of how alpha oscillations shape perception and how this top-down modulation is implemented in cortical networks remain unclear. Here, we tested the hypothesis that alpha oscillations in frontal eye fields (FEFs) are causally involved in the top-down regulation of visual processing in humans (male and female). We applied sham-controlled, intermittent transcranial alternating current stimulation (tACS) over bilateral FEF at either 10 Hz (alpha) or 40 Hz (gamma) to manipulate attentional preparation in a visual discrimination task. Under each stimulation condition, we measured psychometric functions for contrast perception and introduced a novel linear mixed modeling approach for statistical control of neurosensory side effects of the electric stimulation. tACS at alpha frequency reduced the slope of the psychometric function, resulting in improved subthreshold and impaired superthreshold contrast perception. Side effects on the psychometric functions were complex and showed large interindividual variability. Controlling for the impact of side effects on the psychometric parameters by using covariates in the linear mixed model analysis reduced this variability and strengthened the perceptual effect. We propose that alpha tACS over FEF mimicked a state of endogenous attention by strengthening a fronto-occipitoparietal network in the alpha band. We speculate that this network modulation enhanced phasic gating in occipitoparietal cortex leading to increased variability of single-trial psychometric thresholds, measurable as a reduction of psychometric slope.