A Model for the Origin and Properties of Flicker-Induced Geometric Phosphenes

We present a model for flicker phosphenes, the spontaneous appearance of geometric patterns in the visual field when a subject is exposed to diffuse flickering light. We suggest that the phenomenon results from interaction of cortical lateral inhibition with resonant periodic stimuli. We find that the best temporal frequency for eliciting phosphenes is a multiple of intrinsic (damped) oscillatory rhythms in the cortex. We show how both the quantitative and qualitative aspects of the patterns change with frequency of stimulation and provide an explanation for these differences. We use Floquet theory combined with the theory of pattern formation to derive the parameter regimes where the phosphenes occur. We use symmetric bifurcation theory to show why low frequency flicker should produce hexagonal patterns while high frequency produces pinwheels, targets, and spirals

Human Sensation of Transcranial Electric Stimulation.

Noninvasive transcranial electric stimulation is increasingly being used as an advantageous therapy alternative that may activate deep tissues while avoiding drug side-effects. However, not only is there limited evidence for activation of deep tissues by transcranial electric stimulation, its evoked human sensation is understudied and often dismissed as a placebo or secondary effect. By systematically characterizing the human sensation evoked by transcranial alternating-current stimulation, we observed not only stimulus frequency and electrode position dependencies specific for auditory and visual sensation but also a broader presence of somatic sensation ranging from touch and vibration to pain and pressure. We found generally monotonic input-output functions at suprathreshold levels, and often multiple types of sensation occurring simultaneously in response to the same electric stimulation. We further used a recording circuit embedded in a cochlear implant to directly and objectively measure the amount of transcranial electric stimulation reaching the auditory nerve, a deep intercranial target located in the densest bone of the skull. We found an optimal configuration using an ear canal electrode and low-frequency (<300 Hz) sinusoids that delivered maximally ~1% of the transcranial current to the auditory nerve, which was sufficient to produce sound sensation even in deafened ears. Our results suggest that frequency resonance due to neuronal intrinsic electric properties need to be explored for targeted deep brain stimulation and novel brain-computer interfaces

Effect of frequency and rhythmicity on flicker light-induced hallucinatory phenomena

Flicker light stimulation (FLS) uses stroboscopic light on closed eyes to induce transient visual hallucinatory phenomena, such as the perception of geometric patterns, motion, and colours. It remains an open question where the neural correlates of these hallucinatory experiences emerge along the visual pathway. To allow future testing of suggested underlying mechanisms (e.g., changes in functional connectivity, neural entrainment), we sought to systematically characterise the effects of frequency (3 Hz, 8 Hz, 10 Hz and 18 Hz) and rhythmicity (rhythmic and arrhythmic conditions) on flicker-induced subjective experiences. Using a novel questionnaire, we found that flicker frequency and rhythmicity significantly influenced the degree to which participants experienced simple visual hallucinations, particularly the perception of Klüver forms and dynamics (e.g., motion). Participants reported their experience of geometric patterns and dynamics was at highest intensity during 10 Hz rhythmic stimulation. Further, we found that frequency-matched arrhythmic FLS strongly reduced these subjective effects compared to equivalent rhythmic stimulation. Together, these results provide evidence that flicker rhythmicity critically contributes to the effects of FLS beyond the effects of frequency alone, indicating that neural entrainment may drive the induced phenomenal experience

SETTING THE STAGE FOR THE SUN: Interrogations of performance of flicker

Flicker is a coordinated beat which can be superimposed on the brain’s alpha rhythm, through a flickering light source, and is associated with hallucinatory or visionary states of mind. In this experimental piece, I weave ideas and performance into a poetic image and text piece proposing some hypotheses and questions surrounding the relationship of moving bodies, the sun, and eureka moments of visions and dreams of new worlds

Subjective visual experiences of colour and form induced by temporally modulated light

Our understanding of human visual perception generally rests on the assumption that conscious visual states represent the interaction of spatial structures in the environment and our nervous system. This assumption is questioned by circumstances where conscious visual states can be triggered by external stimulation which is not primarily spatially defined. This work discusses psychophysical experiments investigating flicker induced subjective experiences of colour and form. Using the presentation of spatially uniform flicker with a precise temporal resolution it is shown that subjective experiences do not only depend on stimulation frequency, but also on stimulation phase. In addition, the occurrence of one subjective experience appears to be associated with the occurrence of others. The phenomenology of subjective colour and form is further explored and the physiological mechanisms underlying subjective colour are investigated using electroencephalography. While these data indicate that conscious visual experience may be evoked directly by particular variations in the flow of spatially unstructured light over time it may be assumed that the systems reponsible are essentially temporal in character and capable of representing a variety of visual forms and colours, coded in different frequencies or at different phases of the same processing rhythm

Quasicrystal patterns in a neural field model

Doubly periodic patterns in planar neural field models have been extensively studied since the 1970s for their role in explaining geometric visual hallucinations. The study of activity patterns that lack translation invariance has received little, if any, attention. Here we show that a scalar neural field model with a translationally invariant kernel can support quasicrystal solutions and that these can be understood using many of the theoretical tools developed previously for materials science. Our approach is constructive in that we consider constraints on the nonlocal kernel describing interactions in the neural field that lead to the simultaneous excitation of two periodic spatial patterns with incommensurate wavelengths. The resulting kernel has a shape that is a modulation of a Mexican-hat kernel. In the neighborhood of the degenerate bifurcation of a homogeneous steady state, we use a Fourier amplitude approach to determine the value of a Lyapunov functional for various periodic and quasicrystal states. For some values of the parameters defining a translationally invariant synaptic kernel of the model, we find that quasicrystal states have the lowest value of the Lyapunov functional. We observe patterns of 12-fold, 10-fold, and 6-fold rotational symmetry that are stable, but none with 8-fold symmetry. We describe some of the visual hallucination patterns that would be perceived from these quasicrystal cortical patterns, making use of the well known inverse retinocortical map from visual neuroscience

Neural Correlates of Conscious Perception. The Role of Primary Visual Cortex in Visual Awareness.

This study investigates which neural populations represent low-level dimensions of conscious perception. First, a general framework is presented that will allow the separation of different aspects of visual awareness. A set of six criteria is developed that allows one to assess whether a neural population could in principle represent a dimension of conscious perception. These criteria are then applied to previous studies on the neurophysiology and neuropsychology of conscious perception. In the following empirical section a study on the relationship between perceived contrast and activity in primary visual cortex is performed using a combination of EEG, MEG and psychophysics. Lateral masking was used to dissociate the physical and the perceived contrast of a target grating. Transient potentials and magnetic fields evoked by the flashed target gratings were recorded and compared to psychophysical judgements of perceived contrast. At all investigated contrast levels, the amplitudes of electrophysiological transients correlated better with perceived than with physical target contrast. This held especially for the late transient. Source localisation indicated that the transients in question are likely to originate in primary visual cortex. The study presented here is the first ever to study perceptual constancy by recording psychophysics and physiological responses synchronously. The results identify the activity of primary visual cortex as the most likely neural basis of perceived contrast

Migraine Visual Aura and Cortical Spreading Depression—Linking Mathematical Models to Empirical Evidence

This review describes the subjective experience of visual aura in migraine, outlines theoretical models of this phenomenon, and explores how these may be linked to neurochemical, electrophysiological, and psychophysical differences in sensory processing that have been reported in migraine with aura. Reaction–diffusion models have been used to model the hallucinations thought to arise from cortical spreading depolarisation and depression in migraine aura. One aim of this review is to make the underlying principles of these models accessible to a general readership. Cortical spreading depolarisation and depression in these models depends on the balance of the diffusion rate between excitation and inhibition and the occurrence of a large spike in activity to initiate spontaneous pattern formation. We review experimental evidence, including recordings of brain activity made during the aura and attack phase, self-reported triggers of migraine, and psychophysical studies of visual processing in migraine with aura, and how these might relate to mechanisms of excitability that make some people susceptible to aura. Increased cortical excitability, increased neural noise, and fluctuations in oscillatory activity across the migraine cycle are all factors that are likely to contribute to the occurrence of migraine aura. There remain many outstanding questions relating to the current limitations of both models and experimental evidence. Nevertheless, reaction–diffusion models, by providing an integrative theoretical framework, support the generation of testable experimental hypotheses to guide future research

Flicker light stimulation induces thalamocortical hyperconnectivity with LGN and higher-order thalamic nuclei

The thalamus is primarily known as a relay for sensory information; however, it also critically contributes to higher-order cortical processing and coordination. Thalamocortical hyperconnectivity is associated with hallucinatory phenomena that occur in various psychopathologies (e.g., psychosis, migraine aura) and altered states of consciousness (ASC; e.g., induced by psychedelic drugs). However, the exact functional contribution of thalamocortical hyperconnectivity in forming hallucinatory experiences is unclear. Flicker light stimulation (FLS) can be used as an experimental tool to induce transient visual hallucinatory phenomena in healthy participants. Here, we use FLS in combination with fMRI to test how FLS modulates thalamocortical connectivity between specific thalamic nuclei and visual areas. We show that FLS induces thalamocortical hyperconnectivity between lateral geniculate nucleus (LGN), early visual areas, and proximal upstream areas of the ventral visual stream (e.g., hV4, VO1). Further, an exploratory analysis indicates specific higher-order thalamic nuclei, such as anterior and mediodorsal nuclei, to be strongly affected by FLS. Here, the connectivity changes to upstream cortical visual areas directly reflect a frequency-dependent increase in experienced visual phenomena. Together, these findings contribute to the identification of specific thalamocortical interactions in the emergence of visual hallucinations

States and sequences of paired subspace ideals and their relationship to patterned brain function

It is found here that the state of a network of coupled ordinary differential equations is partially localizable through a pair of contractive ideal subspaces, chosen from dual complete lattices related to the synchrony and synchronization of cells within the network. The first lattice is comprised of polydiagonal subspaces, corresponding to synchronous activity patterns that arise from functional equivalences of cell receptive fields. This lattice is dual to a transdiagonal subspace lattice ordering subspaces transverse to these network-compatible synchronies. Combinatorial consideration of contracting polydiagonal and transdiagonal subspace pairs yields a rich array of dynamical possibilities for structured networks. After proving that contraction commutes with the lattice ordering, it is shown that subpopulations of cells are left at fixed potentials when pairs of contracting subspaces span the cells' local coordinates - a phenomenon named glyph formation here. Treatment of mappings between paired states then leads to a theory of network-compatible sequence generation. The theory's utility is illustrated with examples ranging from the construction of a minimal circuit for encoding a simple phoneme to a model of the primary visual cortex including high-dimensional environmental inputs, laminar speficicity, spiking discontinuities, and time delays. In this model, glyph formation and dissolution provide one account for an unexplained anomaly in electroencephalographic recordings under periodic flicker, where stimulus frequencies differing by as little as 1 Hz generate responses varying by an order of magnitude in alpha-band spectral power. Further links between coupled-cell systems and neural dynamics are drawn through a review of synchronization in the brain and its relationship to aggregate observables, focusing again on electroencephalography. Given previous theoretical work relating the geometry of visual hallucinations to symmetries in visual cortex, periodic perturbation of the visual system along a putative symmetry axis is hypothesized to lead to a greater concentration of harmonic spectral energy than asymmetric perturbations; preliminary experimental evidence affirms this hypothesis. To conclude, connections drawn between dynamics, sensation, and behavior are distilled to seven hypotheses, and the potential medical uses of the theory are illustrated with a lattice depiction of ketamine xylazine anaesthesia and a reinterpretation of hemifield neglect