Excitation of subjective light patterns (phosphenes) at different altitudes

Subjective light patterns (phosphenes) can be produced by electrical stimulation of the temples with rectangular current pulses of about 0.1 to 1 mA RMS within the electroencephalographic frequency range (1 to 70 c/s). The phosphenes either consist of formless flickering of the visual field or of different abstract geometrical figures on a dark background. Electrical excitation of phosphenes in Munich (altitude 530 m) and in the mountains at 2962 m showed the tendency that more phosphenes can be elicited at high altitude. There was no difference between the types of phosphenes obtained at the two altitude levels.     

Psychological aspects of perception of magnetophosphenes and electrophosphenes]

Scientific research at the Helmholtz Institute for Biomedical Engineering at Aachen University indicates that ELF (extremely-low frequency) electric and magnetic fields may generate visual perceptions in the human retina which are similar to the pressure phosphenes. During our own experiments we found that 90% of subjects undergoing a blind experiment reported various visual sensations which were mostly colored and moving. Our findings indicate that the psychological component of the perception of electric and magnetic phosphenes must not be underestimated. It is possible that there is a connection between retinal noise in the dark (due to metabolic processes [5, 8]) and magnetic or electric phosphenes.     

Spatially localized distortions of event time

A fundamental question about the perception of time is whether the neural mechanisms underlying temporal judgements are universal and centralized in the brain or modality specific and distributed. Time perception has traditionally been thought to be entirely dissociated from spatial vision. Here we show that the apparent duration of a dynamic stimulus can be manipulated in a local region of visual space by adapting to oscillatory motion or flicker. This implicates spatially localized temporal mechanisms in duration perception. We do not see concomitant changes in the time of onset or offset of the test patterns, demonstrating a direct local effect on duration perception rather than an indirect effect on the time course of neural processing. The effects of adaptation on duration perception can also be dissociated from motion or flicker perception per se. Although 20 Hz adaptation reduces both the apparent temporal frequency and duration of a 10 Hz test stimulus, 5 Hz adaptation increases apparent temporal frequency but has little effect on duration perception. We conclude that there is a peripheral, spatially localized, essentially visual component involved in sensing the duration of visual events.     

Temporal properties of the lens eyes of the box jellyfish Tripedalia cystophora

Box jellyfish (Cubomedusae) are visually orientating animals which posses a total of 24 eyes of 4 morphological types; 2 pigment cup eyes (pit eye and slit eye) and 2 lens eyes [upper lens-eye (ule) and lower lens-eye (lle)]. In this study, we use electroretinograms (ERGs) to explore temporal properties of the two lens eyes. We find that the ERG of both lens eyes are complex and using sinusoidal flicker stimuli we find that both lens eyes have slow temporal resolution. The average flicker fusion frequency (FFF) was found to be approximately 10 Hz for the ule and 8 Hz for the lle. Differences in the FFF and response patterns between the two lens eyes suggest that the ule and lle filter information differently in the temporal domain and thus are tuned to perform different visual tasks. The data collected in this study support the idea that the visual system of box jellyfish is a collection of special purpose eyes.     

THRESHOLD INTENSITY OF ILLUMINATION AND FLICKER FREQUENCY FOR THE EYE OF THE SUN-FISH

The sun-fish Lepomis responds to a moving system of stripes by a motion of its body. By changing the velocity of motion of the stripe system different flicker frequencies can be produced and thus the relation of flicker frequency to critical intensity of illumination can be studied. Threshold illumination varies with flicker frequency in such a way that with increasing flicker frequency the intensity of illumination must be increased to produce a threshold response in the fish. The curve of critical illumination as a function of frequency is made up of two distinct parts. For an intensity range below 0.04 millilambert and flicker frequencies below 10 per second, the rods are in function. For higher intensities and flicker frequencies above 10, the cones come into play. The maximum frequency of flicker which can be perceived by the fish''s eye is slightly above 50 per second. The flicker curve for the eye of Lepomis can easily be compared with that for the human eye. The extent of the curve for the fish is greater at low illuminations, the fish being capable of distinguishing flicker at illuminations lower than can the human eye. The transition of rod vision to cone vision occurs for the fish and for the human eye at the same intensity and flicker frequency. The maximum frequency of flicker which can be perceived is for both about the same.     

Flicker alternating potentials in the ERGs of insects in response to two different stimuli

Summary A special pattern of the flicker is studied in insects belonging to four Orders, i. e. the differential electrical synchronised response of the eye periodically stimulated by two slightly different alternating illuminations.After having checked that the flicker in response to a regular periodical stimulation at every frequency is made up of successive equal potentials, we use two slightly different alternate flickering flashes. It is established that the alternation of two periodical stimulations of a different duration, as well as the alternation of two periodical stimulations of a different intensity, results, over a certain frequency which depends on the insect studied, in the appearance of a flicker marked with the alternation of two potentials whose difference increases at the same time as the frequency of stimulation.The dependence of this phenomenon on modulation of the light flux is described. At a given frequency of stimulation, the alternation of a high and low potential is more obvious when the modulation is lower.A particular experiment allows us to admit that the differential threshold of electrical response to two different stimulations is under 0.25%, at the frequency 100 Hz, inCalliphora erythrocephala.All the phenomena observed can be explained by a mathematical theory which considers the characteristics of the amplitude of the response to sinusoidal stimulations of various frequencies, i. e. the characteristics of the transfer function of the frequencies.     

CRITICAL FREQUENCY OF FLICKER AS A FUNCTION OF INTENSITY OF ILLUMINATION FOR THE EYE OF THE BEE

The bee''s characteristic response to a movement of its visual field is used for the study of the relation between critical frequency of flicker and illumination. The critical flicker frequency varies with illumination in such a way that with increasing flicker frequency the intensity of illumination must be increased to produce a threshold response in the bee. The illuminations required to give a response in a bee at different flicker frequencies closely correspond to the intensities for threshold response in visual acuity tests. This is due to the different thresholds of excitability of the elements of the ommatidial mosaic. An analysis of the variation of the values for threshold intensities at the several flicker frequencies shows that the variation depends upon flicker frequency and upon the number of elements functioning at different intensities.     

fMRI of Retina-Originated Phosphenes Experienced by Patients with Leber Congenital Amaurosis

A phenomenon characterized by the experience of seeing light without any light actually entering the eye is called phosphenes or photopsias. Phosphenes can occur spontaneously or via induction by external stimuli. Previous reports regarding phosphenes have primarily focused on externally induced phosphenes such as by applying alternating or direct current to the cortex. A few of these reports used functional magnetic resonance (fMRI) to study activations induced by cortical phosphenes. However, there are no fMRI reports on spontaneous phosphenes originating from the retina and the resulting pattern of cortical activations. We performed fMRI during a reversing checkerboard paradigm in three LCA patients who underwent unilateral gene therapy and reported experiencing frequent phosphene on a daily basis. We observed bilateral cortical activation covering the entire visual cortices when patients reported experiencing phosphenes. In contrast, in the absence of phosphenes, activation was regulated by patient''s visual ability and demonstrated improved cortical activation due to gene therapy. These fMRI results illustrate the potential impact of phosphene perception on visual function and they may explain some of the variability that clinicians find in visual function testing in retinal degeneration. Although we did not perform correlations between visual function and phosphenes, we hope data presented here raises awareness of this phenomenon and its potential effect on visual function and the implications for clinical testing. We recommend a thorough history for phosphene experiences be taken in patients with retinal disease who are candidates for gene or molecular therapy. Lastly, these data illustrate the potential power of fMRI as an outcome measure of gene therapy and the negative impact phosphenes may have on vision testing. fMRI has proven to be a sensitive, non-invasive, and reproducible test paradigm for these purposes and can complement standard visual function testing.     

Frequency‐dependent and montage‐based differences in phosphene perception thresholds via transcranial alternating current stimulation

It is well known that applying transcranial alternating current stimulation (tACS) to the scalp can generate artefactual visual perceptions of flashing or shimmering light known as phosphenes. The thresholds for generating these phosphenes have been used by international standards bodies to provide conservative estimates of the field strength required to interfere with human neural functioning and set safety limits accordingly. However, the precise relationship between electric currents and phosphene perception thresholds remains uncertain. The present study used tACS to systematically investigate the effects of the location and the frequency of stimulation on phosphene perception thresholds. These thresholds were obtained from 24 participants using a within‐subject design as a function of scalp stimulation sites (FPz‐Cz versus Oz‐Cz) and stimulation frequency (2–30 Hz in steps of 2 Hz). Phosphene perception thresholds were consistently lower for FPz‐Cz stimulation, and regardless of tACS location were lowest for 16 Hz stimulation. Threshold variation between participants was very small, which is meaningful when setting standards based on phosphenes. Bioelectromagnetics. 2019;40:365–374. © 2019 Bioelectromagnetics Society.     

ON CRITICAL FREQUENCY AND CRITICAL ILLUMINATION FOR RESPONSE TO FLICKERED LIGHT

The curve of mean critical flicker frequency as a function of illumination has been determined for the reaction of the sunfish Lepomis to flicker. It exhibits expected quantitative disagreements with the curve of mean critical illumination as a function of flicker frequency in the same organism. The form of the dependence of the variation of critical frequency of flicker upon illumination can be predicted from a knowledge of the way in which variation of critical illumination depends upon flicker frequency. It is pointed out that these findings have an important bearing upon the interpretation of the data of intensity discrimination.     

Visual capacities of the owl monkey (Aotus trivirgatus): temporal contrast sensitivity.

Aotus monkeys were tested in a forced-choice discrimination task to determine their ability to discriminate sinusoidally flickering lights varying in temporal frequency and luminance contrast. Under conditions of moderate light adaptation this primate is maximally sensitive to lights flickering at 10 Hz while the highest frequency they can discriminate is about 42 Hz. At very low light levels (10(-5) ft L) maximum sensitivity is for 2.2--5 Hz flicker. The highest flicker rate that could be discriminated under these conditons was about 29 Hz. In comparison to humans tested in the same situation. Aotus monkeys show relatively lower sensitivity to temporal flicker under conditions of light adaptation but relatively higher sensitivity at very low light levels.     

The Frequency-Response Electroretinogram Distinguishes Cone and Abnormal Rod Function in rd12 Mice

Early studies on Rpe65 knockout mice reported that remaining visual function was attributable to cone function. However, this finding has been challenged more and more as time has passed. Electroretinograms (ERGs) showed that rd12 mice, a spontaneous animal model of RPE65 Leber’s congenital amaurosis, had sizeable photopic responses. Unfortunately, the recorded ERG waveform was difficult to interpret because of a remarkably delayed peak-time, which resembles a rod response more than a cone response. Here, we compare flicker ERGs in animals with normal rod and cone function (C57BL/6J mice), pure rod function (cpfl5 mice), and pure cone function (Rho^-/- mice) under different adaptation levels and stimulus intensities. These responses were then compared with those obtained from rd12 mice. Our results showed that normal rods respond to low frequency flicker (5 and 15 Hz) and that normal cones respond to both low and high frequency flicker (5–35 Hz). As was seen in cpfl5 mice, rd12 mice had recordable responses to low frequency flicker (5 and 15Hz), but not to high frequency flicker (25 and 35 Hz). We hypothesize that abnormal rods may be the source of residual vision in rd12 mice, which is proved correct here with double mutant rd12mice. In this study, we show, for the first time, that frequency-response ERGs can effectively distinguish cone- and rod-driven responses in the rd12 mouse. It is another simple and valid method for evaluating the respective contributions of retinal rods and cones.     

Conscious awareness of flicker in humans involves frontal and parietal cortex

Even when confined to the same spatial location, flickering and steady light evoke very different conscious experiences because of their distinct temporal patterns. The neural basis of such differences in subjective experience remains uncertain . Here, we used functional MRI in humans to examine the neural structures involved in awareness of flicker. Participants viewed a single point source of light that flickered at the critical flicker fusion (CFF) threshold, where the same stimulus is sometimes perceived as flickering and sometimes as steady (fused) . We were thus able to compare brain activity for conscious percepts that differed qualitatively (flickering or fused) but were evoked by identical physical stimuli. Greater brain activation was observed on flicker (versus fused) trials in regions of frontal and parietal cortex previously associated with visual awareness in tasks that did not require detection of temporal patterns . In contrast, greater activation was observed on fused (versus flicker) trials in occipital extrastriate cortex. Our findings indicate that activity of higher-level cortical areas is important for awareness of temporally distinct visual events in the context of a nonspatial task, and they thus suggest that frontal and parietal regions may play a general role in visual awareness.     

Transfer of stimulus control from a TFT to CRT screen

The use of television and computer screens for presenting stimuli to animals is increasing as it is non-invasive and can provide precise control over stimuli. Past studies have used cathode ray tube (CRT) screens; however, there is some evidence that these give different results to non-flickering thin film transistor (TFT) screens. Hens’ critical flicker fusion frequency ranges between 80 and 90 Hz - above standard CRT screens. Thus, stimuli presented on CRT screens may appear distorted to hens. This study aimed to investigate whether changing the flicker rate of CRT screens altered hens’ discrimination. Hens were trained (in a conditional discrimination) to discriminate between two stimuli on a TFT (flickerless) screen, and tested with the stimuli on a CRT screen at four flicker rates (60, 75, 85, and 100 Hz). The hens’ accuracy generally decreased as the refresh rate of the CRT screen decreased. These results imply that the change in flicker rate changed the appearance of the stimuli enough to affect the hens’ discrimination and stimulus control is disrupted when the stimuli appear to flicker.     

Temporal resolution in mesopelagic crustaceans

Mesopelagic crustaceans occupy a dim-light environment that is similar to that of nocturnal insects. In a light-limited environment, the requirement for greater sensitivity may result in slower photoreceptor transduction and increased summation time. This should be reflected by a lower temporal resolution, as indicated by a lower critical flicker fusion frequency (CFF). Therefore, one would predict that the CFFs of mesopelagic organisms would be relatively low compared with those of their shallow-water relatives, just as nocturnal insects tend to have lower CFFs than diurnal insects. Using an electrophysiological apparatus that was adapted for shipboard use, the dark-adapted CFFs of a variety of species of mesopelagic crustaceans were determined using the electroretinogram. The parameter examined was the maximum CFF--the point at which further increases in irradiance no longer result in a faster flicker fusion frequency. The results summarized here indicate that there is a trend towards lower CFFs with increasing habitat depth, with some interesting exceptions.     

REACTIONS OF LIMULUS TO ILLUMINATED FIELDS OF DIFFERENT AREA AND FLICKER FREQUENCY

In phototropic tests with young Limulus, the phototropic reactions to flickering fields were studied. If the two fields are equal in area and brightness but different in flicker frequency, the number of animals going to the two fields is proportional to their flicker frequencies. Equal stimulating effects of two fields differing in flicker frequency are obtained by reduction of the area of the faster flickering field. The areas for equal effect must be inversely proportional to their flicker frequencies. It seems that equal effects are dependent upon equality of the number of active excitation elements per unit of time.     

Perception of heterochromatic flicker by honeybees: a behavioural study

Freely flying honeybees were trained to discriminate a stimulus consisting of two alternating chromatic lights (heterochromatic flicker) from a steady mixture of the same two lights, using 3 different pairs of lights: blue-UV, UV-green, and green-UV. With each light pair, training to the heterochromatic flicker was conducted at several flicker frequencies, using experimentally naive bees in each training. In subsequent tests, the trained bees were given a choice between the two lights that constituted the flicker, presented steady, as well as between either of them and the steady mixture. We find that bees trained to particular frequencies of heterochromatic flicker prefer one of the component lights over the other as well as over the steady mixture, suggesting that the colour they perceive in the heterochromatic flicker to which they have been trained is shifted in the direction of one of the lights contained in the flicker. The colour shift occurs at flicker frequencies that depend on the pair of lights used. We propose that the shift is generated by an effect similar to the Brücke-Bartley phenomenon known from human vision. This effect is based on the enhancement of the photoreceptors' response upon onset of stimulation, causing an intermittent light to appear brighter than a steady light of identical physical intensity. We propose that the degree of enhancement might differ among the 3 spectral classes of photoreceptor, causing the colour perceived in a heterochromatic flicker to differ from that perceived in a steady mixture of its two light components.     

Shortening of subjective visual intervals followed by repetitive stimulation

Our previous research demonstrated that repetitive tone stimulation shortened the perceived duration of the preceding auditory time interval. In this study, we examined whether repetitive visual stimulation influences the perception of preceding visual time intervals. Results showed that a time interval followed by a high-frequency visual flicker was perceived as shorter than that followed by a low-frequency visual flicker. The perceived duration decreased as the frequency of the visual flicker increased. The visual flicker presented in one hemifield shortened the apparent time interval in the other hemifield. A final experiment showed that repetitive tone stimulation also shortened the perceived duration of preceding visual time intervals. We concluded that visual flicker shortened the perceived duration of preceding visual time intervals in the same way as repetitive auditory stimulation shortened the subjective duration of preceding tones.     

用稳态视觉诱发电位研究注意的选择机制

众所周知,注意是一种心理活动,是对外界刺激的一种选择,对人的其它认知活动有一定的调节作用。关于注意的选择机制,有两种不同的理论,一种是早期选择机制,另一种是晚期选择机制。稳态视觉诱发电位是视觉通路对外界刺激的一种物理反应,它的产生过程处于认知的早期阶段,如果它能受到注意的调节,这就支持了注意的早期选择机制。在现有的利用闪光稳态诱发电位对注意的研究中,注意的对象是认知任务,而不是闪光。该实验用两种不同频率的闪光分别做诱发源,对在注意与不注意闪光两种情况下(都没有认知任务)的稳态视觉诱发电位进行研究,发现其明显地受注意调节,而且对不同频率光刺激的调节大小不一样。因此,该实验结果支持了注意的早期选择机制。     

Frequency dependent flicker response enhancement in the lamina ganglionaris ofDrosophila

Summary At low light intensity and within the narrow frequency range of 55 to 66 s^–1, the eye ofDrosophila will follow a flashing light source by enhancing it's flicker response to every other flash. By contrast, at lower and higher frequencies the eye will follow every cycle of a respective flash frequency upto a fusion point around 200 s^–1.While the receptor cells involved are retinula cells R1–6, the flicker response enhancement is established to originate postsynaptically in the Large Monopolar Cells of the lamina with which the peripheral retinula cells synapse, and which respond with the cornealpositive on-transient component of the ERG. Not only is a prescribed frequency required for the enhancement, but also continuity of cue — since brief periods of light flashes within the required frequency range are resolved at every cycle.The flicker response behaviour provides further credence to the existence of fine tuning mechanisms together with amplification within the lamina neuropile.We are grateful to the late Dr. Richard Wright for his comments, and to Professor Aubrey Manning for the hospitality his Department gave to N.L.