Electrical neuromodulation biases brightness perception of flickering light

Human brightness estimation often pronouncedly dissociates from objective viewing conditions. Yet, the physiological substrate underlying subjective perception is still poorly understood. Rather than physical illumination, the subjective experience of brightness has been shown to correlate with temporal dynamics in the amplitude of cortical neural responses. Here, we aimed to experimentally manipulate visual flicker-evoked steady-state responses and related perception via concurrent modulation of cortical excitability by transcranial alternating current stimulation. Participants performed a brightness discrimination task of two visual flicker stimuli, one of which was targeted by same-frequency electrical stimulation at varying phase shifts. Transcranial electrical stimulation was applied with an occipital and a periorbital active control montage, based on finite-element method simulations of electric fields. Experimental results reveal that flicker brightness perception is modulated dependent on the phase shift between sensory and electrical stimulation, solely under stable flicker entrainment and exclusively under occipital electrical stimulation. The degree of induced brightness modulation was positively correlated with the strength of neuronal phase locking to the flicker, recorded prior to electrical stimulation. This finding was corroborated by a neural network model, demonstrating a comparable dependency between flicker-evoked phase synchronization and amplitude modulations of entrained neural rhythms by phase shifted visual and electric inputs. Our data suggest a causal role of the amplitude of neural activity in visual cortex for brightness perception in humans. This finding provides an important step towards understanding the basis of visual perception and further confirms electrical stimulation as a tool for advancing controlled modulations of neural excitability and related behavior.

A systematic investigation of differences in perceptual decision-making as a function of the time and method of participant recruitment

We examine the extent to which perceptual decision-making processes differ as a function of the time in the academic term in which the participant enrolls in the experiment and whether the participant is an undergraduate who completes the experiment for course credit, a paid participant who completes the experiment in the lab, or a paid participant recruited via Amazon Mechanical Turk who completes the experiment online. In Study 1, we conducted a survey to examine cognitive psychologists' expectations regarding the quality of data obtained from these different groups of participants. We find that cognitive psychologists expect performance and response caution to be lowest among undergraduate participants who enroll at the end of the academic term, and highest among paid in-lab participants. Studies 2 and 3 tested these expectations using two common perceptual decision-making paradigms. Overall, we found little evidence for systematic time-of-term effects among undergraduate participants. The different participant groups responded to standard stimulus quality and speed/accuracy emphasis manipulations in similar ways. Among participants recruited via Mechanical Turk, the effect of speed/accuracy emphasis on response caution was strongest. This group also showed poorer discrimination performance than the other groups in a motion discrimination task, but not in a brightness discrimination task. We conclude that online crowdsourcing platforms can provide high quality perceptual decision-making data, but give recommendations for how data quality can be maximized when using these platforms for recruitment.

Melanopsin- and L-cone–induced pupil constriction is inhibited by S- and M-cones in humans

The human retina contains five photoreceptor types: rods; short (S)-, mid (M)-, and long (L)-wavelength–sensitive cones; and melanopsin-expressing ganglion cells. Recently, it has been shown that selective increments in M-cone activation are paradoxically perceived as brightness decrements, as opposed to L-cone increments. Here we show that similar effects are also observed in the pupillary light response, whereby M-cone or S-cone increments lead to pupil dilation whereas L-cone or melanopic illuminance increments resulted in pupil constriction. Additionally, intermittent photoreceptor activation increased pupil constriction over a 30-min interval. Modulation of L-cone or melanopic illuminance within the 0.25–4-Hz frequency range resulted in more sustained pupillary constriction than light of constant intensity. Opposite results were found for S-cone and M-cone modulations (2 Hz), mirroring the dichotomy observed in the transient responses. The transient and sustained pupillary light responses therefore suggest that S- and M-cones provide inhibitory input to the pupillary control system when selectively activated, whereas L-cones and melanopsin response fulfill an excitatory role. These findings provide insight into functional networks in the human retina and the effect of color-coding in nonvisual responses to light, and imply that nonvisual and visual brightness discrimination may share a common pathway that starts in the retina.

Entrainment of theta, not alpha, oscillations is predictive of the brightness enhancement of a flickering stimulus

Frequency-dependent brightness enhancement, where a flickering light can appear twice as bright as an equiluminant constant light, has been reported to exist within the alpha (8 – 12 Hz) band. Could oscillatory neural activity be driving this perceptual effect? Here, in two experiments, human subjects reported which of two flickering stimuli were brighter. Strikingly, 4 Hz stimuli were reported as brighter more than 80% of the time when compared to all other tested frequencies, even though all stimuli were equiluminant and of equal temporal length. Electroencephalography recordings showed that inter-trial phase coherence (ITC) of theta (4 Hz) was: 1) Significantly greater than alpha, contralateral to the flickering stimulus; 2) Enhanced by the presence of a second ipsilateral 4 Hz flickering stimulus; and 3) Uniquely lateralized, unlike the alpha band. Importantly, on trials with two identical stimuli (i.e. 4 Hz vs 4 Hz), the brightness discrimination judgment could be predicted by the hemispheric balance in the amount of 4 Hz ITC. We speculate that the theta rhythm plays a distinct information transfer role, where its ability to share information between hemispheres via entrainment promotes a better processing of visual information to inform a discrimination decision.

Cholinergic Potentiation of Restoration of Visual Function after Optic Nerve Damage in Rats

Enhancing cortical plasticity and brain connectivity may improve residual vision following a visual impairment. Since acetylcholine plays an important role in attention and neuronal plasticity, we explored whether potentiation of the cholinergic transmission has an effect on the visual function restoration. To this end, we evaluated for 4 weeks the effect of the acetylcholinesterase inhibitor donepezil on brightness discrimination, visually evoked potentials, and visual cortex reactivity after a bilateral and partial optic nerve crush in adult rats. Donepezil administration enhanced brightness discrimination capacity after optic nerve crush compared to nontreated animals. The visually evoked activation of the primary visual cortex was not restored, as measured by evoked potentials, but the cortical neuronal activity measured by thallium autometallography was not significantly affected four weeks after the optic nerve crush. Altogether, the results suggest a role of the cholinergic system in postlesion cortical plasticity. This finding agrees with the view that restoration of visual function may involve mechanisms beyond the area of primary damage and opens a new perspective for improving visual rehabilitation in humans.