A linear chromatic mechanism drives the pupillary response.

Previous studies have shown that a chromatic mechanism can drive pupil responses. The aim of this research was to clarify whether a linear or nonlinear chromatic mechanism drives pupillary responses by using test stimuli of various colours that are defined in cone contrast space. The pupil and accommodation responses evoked by these test stimuli were continuously and simultaneously objectively measured by photorefraction. The results with isochromatic and isoluminant stimuli showed that the accommodative level remained approximately constant (< 0.25 D change in mean level) even when the concurrent pupillary response was large (ca. 0.30 mm). The pupillary response to an isoluminant grating was sustained, delayed (by ca. 60 ms) and larger in amplitude than that for a isochromatic uniform stimulus, which supports previous work suggesting that the chromatic mechanism contributes to the pupillary response. In a second experiment, selected chromatic test gratings were used and isoresponse contours in cone contrast space were obtained. The results showed that the isoresponse contour in cone contrast space is well described (r(2) = 0.99) by a straight line with a positive slope. The results indicate that a /L - M/ linear chromatic mechanism, whereby a signal from the long wavelength cone is subtracted from that of the middle wavelength cone and vice versa, drives pupillary responses.     

Visual pigment processes and prolonged pupillary responses in insect photoreceptor cells

The visual pigment in the peripheral retinular cells of the hoverfly Syrphus balteatus was investigated by absorbance difference measurements. Different visual pigments were found in the dorsal versus the ventral part of the eye in the male, but not in the female. In the male in the dorsal part of the eye the visual pigment has an isosbestic point at 513 nm; in the ventral part this value is 490 nm. The latter value is found in the female in both parts of the eye.Prolonged pupillary responses were studied in the male Syrphus and appeared to be most marked in the ventral part of the eye. In both hoverfly and blowfly prolonged pupillary responses are induced by short wavelength light only; i.e., by light which excessively can convert rhodopsin into metarhodopsin. By contrast, in butterflies red light (and a long dark adaptation time) is necessary to evoke a prolonged pupillary response. It was demonstrated in both hoverfly and blowfly that long wavelength light, which reconverts metarhodopsin into rhodopsin, inhibits a prolonged pupillary response; or, accelerates pupil opening.Based on material presented at the European Neurosciences Meeting, Florence, September 1978     

Spectral effects of the pupil in fly photoreceptors

Summary Photoreceptors of flies contain pigment granules which upon illumination of the receptors migrate towards the rhabdomere and act as a longitudinal pupil. Data in the literature concerning the effect of the pupil on the spectral sensitivity are contradictory. Therefore spectral sensitivity ofMusca photoreceptors upon light adaptation was reinvestigated.The change in spectral sensitivity of fly photoreceptors upon light adaptation as measured by Hardie (1979) was confirmed. Taking into account waveguide optics this change was explained from absorbance spectra of pupillary granules, measured by microspectrophotometry in squash preparations. Furthermore the pupil absorbance spectrum determined in vivo (Stavenga et al. 1973) was interpreted. The absence of a change in spectral sensitivity upon light adaptation measured by pupillary reflexion (Bernard and Stavenga 1979) is explained by a local-triggering of the pupil.     

Investigations of pigment granule transport systems in Gonodactylus oerstedii (Crustacea: Hoplocarida: Stomatopoda)

Compound eyes of the stomatopod, Gonodactylus oerstedii, exhibit pupillary reflection responses which arise from migration of retinular cell pigment granules. In the light, reflectance from the eye increases as pigment granules accumulate around light-sensitive rhabdoms and scatter incoming light back out of the eye (pupillary closure). At dark onset, reflectance diminishes as pigment granules disperse centrifugally, enhancing photon capture by the rhabdom. We investigated the mechanisms of the pupillary response in intact animals by measuring reflectance from the eye under different temperature conditions. Lowering the temperature from 27° to 7 °C caused an increase in reflectance of infrared light in the absence of visible-light stimuli, indicating pupillary closure. When given light stimuli as temperature decreased, the eye continued to produce reflection increases which decreased in amplitude as the between stimulus reflectance level increased. All low-temperature effects were reversed when temperature was increased to normal. The rate of pupillary closure was insensitive to temperature, with a temperature quotient (Q₁₀) of 0.8 ± 0.1 s.e.m, while pupillary opening was extremely temperature sensitive (Q₁₀ of 5.4 ± 0.4). Different temperature sensitivities for pupillary opening and closing suggest that these processes involve different mechanisms.Abbreviations IOP intracelllular optical physiology - Q₁₀ temperature quotient     

Melanopsin Gene Polymorphism I394T Is Associated with Pupillary Light Responses in a Dose-Dependent Manner

Background

Melanopsin-containing intrinsically photosensitive retinal ganglion cells (ipRGCs) play an important role in non-image forming responses to light, such as circadian photoentrainment, light-induced melatonin suppression, and pupillary light response. Although it is known that there are some single nucleotide polymorphisms (SNPs) in the melanopsin (OPN4) gene in humans, the associations of the SNPs with non-image forming responses to light remains unclear. In the present study, we examined the associations of melanopsin gene polymorphisms with pupillary light response.

Methods

Japanese university students (mean age: 21.0±1.7 years) with the genotypes of TT (n = 38), TC (n = 28) and CC (n = 7) at rs1079610 (I394T) located in the coding region participated in the present study. They were matched by age and sex ratio. Dark-adapted pupil size (<1 lx) was first measured. Then steady-state pupil size was measured during exposure to five lighting conditions (10 lx, 100 lx, 1000 lx, 3000 lx, 6000 lx in the vertical direction at eye level).

Results

Significant interaction between the genotype of I394T (TT versus TC+CC) and luminance levels was found in pupil size. Under high illuminance levels (1000 lx, 3000 lx and 6000 lx), pupil sizes in subjects with the C allele were significantly smaller than those in subjects with the TT genotype. On the other hand, pupil size in subjects with the C allele under low illuminance (<1 lx) was significantly larger than that in subjects with the TT genotype. Percentages of pupil constriction under high illuminance levels were significantly greater in subjects with the C allele than in subjects with the TT genotype.

Conclusions

Human melanopsin gene polymorphism I394T interacted with irradiance in association with pupil size. This is the first evidence suggesting a functional connection between melanopsin gene polymorphism and pupillary light response as an index of non-image forming response to light.     

瞳孔控制系统非线性的频率特性与固有频率

本文用实验方法从时域和频域上揭示了瞳孔对光反应的非线性特性.在亮度以正弦变化的光刺激下,瞳孔反应波形呈同步倍频现象;描述函数的频率-振幅特性曲线呈多峰的锯齿形;具有1.2Hz左右的系统固有频率和极限环现象.     

The Light Growth Response of Phycomyces

With the help of an automated tracking system we have studied the characteristics of the transient light growth response of Phycomyces. The response shows a sharply defined latency. The Q₁₀ of the reciprocal latency is 2.4. Response patterns at different peaks of the action spectrum are the same. The gradual variation of response magnitude over a wide range of adapted intensifies parallels that of phototropism. The responses to saturating stimuli exhibit a strong oscillation with a constant period of 1.6 min and variable damping. The growth responses to sinusoidally varying light intensities show a system bandwidth of 2.5 x 10^-3 Hz. The linear dependence of phase shift on frequency is largely attributable to the latency observed with pulse stimuli. In the high intensity range a previously suspected increase of the steady-state growth rate with intensity has been confirmed. The light growth responses of mutants selected for diminished phototropism have been investigated. Many of these mutants have sizable but grossly distorted growth responses.     

Binocular alternating pulse stimuli: Experimental and modeling studies of the pupil reflex to light

In our experiment, alternating pulse stimuli of both low and high intensity are used to study the pupil reflex to light. When applied monocularly, high intensity stimulation normally results in a sustained contraction; when alternated between the two eyes, it is found to produce small transient responses similar to those obtained with low intensity monocular stimulation. In order to study the mechanisms regulating these binocular responses, a model of the pupillary light reflex is constructed. It includes parallel AC and DC pathways for processing the light stimulus to produce motor signals to the iris muscles, nonlinear parameter control of pathway gains dependent upon internal operating level, binocular summation of DC pathway signals to produce that operating level, equal motor responses of both pupils, and iris neuromuscular delays and lags. The model is found to simulate the experimental data. It shows the binocular transient responses to be due to the canceling by summation of the symmetric DC pathway responses to alternating stimuli, thus allowing the AC pathway signals to become manifest. Therefore the dilatory portion of the transient responses is shown to be due to the lead-lag operator in the AC pathway and not to the off-dilatation elicited by removal of the light stimulus from the eye. Finally the results of our study are used to discuss the Marcus Gunn pupillary sign, a clinical test utilizing this binocular alternating pulse stimulation for detecting unilateral afferent defects.     

The Dependence of the Photopupil Response on Flash Duration and Intensity

The changes in pupil size were recorded by infrared pupillographic methods in response to light flashes of different durations and intensities for a 13 degree 34 minute centrally fixated circular field. For such stimuli, the threshold intensities for (rod) vision and for the pupil response were found to be about the same. The response amplitudes were related to the logarithm of the flash energy, the reciprocity law remaining valid up to about one-half second. The curve relating flash energy and pupil response was clearly divisible into two parts commensurate with the duplex character of the human retina. A similar dichotomy appears in curves relating response amplitude to response latency. Since the pupil response is determined by total flash energy, intense long flashes produce larger pupil responses than shorter (and perceptually brighter) ones of the same intensity.     

在双脉冲光刺激下瞳孔系统的采样控制特性

本文用实验揭示了瞳孔对光动态反应具有采样控制特性。实验中采用各种不同时间间隔的双脉冲光,以开环的方式(Maxwellian View)刺激瞳孔,当双脉冲之间间隔较长时,瞳孔反应相当于对双脉冲光的两次脉冲分别产生瞬态收缩;当双脉冲时间间隔短于0.6s 时,其反应就成了一次瞬态收缩,与单个光脉冲所引起的瞳孔反应一样。同—受试者的多次实验结果相同,不同受试者所得结果也基本一致。故瞳孔对脉冲刺激光引起反应后,必须至少约隔0.6s 才能对另一次脉冲光产生反应,这就说明了瞳孔动态反应具有离散的采样控制特性。实验还进一步证明,瞳孔系统的控制机制是双重模式的控制:不同的刺激条件下,瞳孔反应可呈现为瞬态反应(AC)或持续反应(DC),瞬态反应的 AC 通道为离散的采样控制,持续反应的 DC 通道为连续控制。     

Visual motion,binocular correspondence and binocular rivalry

The human pupillary control system has been the subject of interest to biologists and engineers as an example of a sensorimotor reflex which can be embedded in a control system paradigm. We present a nonlinear feedback model whose compact structure allows us to hypothesize possible physiological mechanisms which generate the proper behavior of the pupil system. The important pupil responses, including pupil size effect, asymmetry, and response to high-frequency stimuli, are defined. This model was simulated on a digital computer and comparisons to the paradigm experimental responses were performed, demonstrating a fit to each of the observed conditions. Improvements on previous models are discussed.     

Quantitative Beziehungen zwischen Lichtreiz und Kontraktion des Musculus sphincter pupillae vom Scheibenzüngler (Discoglossus pictus)

Summary The Musculus sphincter pupillae of the toad Discoglossus pictus contracts when exposed to light, even if the iris with the muscle is cut out of the eye. Normally the size of the pupil is controlled by this photosensitivity and in addition by a nervous control mechanism. —Experiments are discribed in which the M. sphincter pupillae of the isolated iris was stimulated by various light programs. The contractile force was measured isometrically with a compensation system. It is shown that the response of the sphincter-muscle to changing light stimuli depends on the mean light intensity; i.e. the sphincter-muscle is an adapting system. The amplitude and phaseshift of the reactions to sinusoidal light stimuli were measured. When stimulus and response are both considered as functions of time, the transformation of stimulus in response by the muscle is linear, provided that the frequency of the sinusoidal light is high and its intensity and modulation degree are low. From this result the hypothesis is derived, that the transformation of stimulus into response is linear only when the level of adaptation does not change during contraction. From the response to sinusoidal light program it was possible to predict the reaction to square wave light programs. The time-course of the reaction to step up light stimuli are similar to the time-course of receptor potentials of phasic-tonic sense cells, showing a maximum, a minimum and a plateau. Within certain limits the amplitude of the reaction depends linearly on the tension of the muscle. Since in isometric experiments the reaction consists of an increase of muscle tension, the muscle becomes more sensitive during reaction and the contractile force increases for a long time, even when light intensity is held constant.     

Sinusoidal and delta function responses of visual cells of the Limulus eye

Dynamic responses of visual cells of the Limulus eye to stimuli of sinusoids and narrow pulses of light superimposed on a nonzero mean level have been obtained. Amplitudes and phase angles of averaged sinusoidal generator potential are plotted with respect to frequency of intensity modulation for different mean levels of light adaptation. At frequencies above 10 CPS, generator potential amplitudes decrease sharply and phase lag angle increases. At frequencies below 1 CPS, amplitude decreases. A maximum of amplitude in the region of 1 to 2 CPS is apparent with increased mean intensity. The generator potential responses are compared with those of differential equation models. Variation of gain with mean intensity for incremental stimuli is consistent with logarithmic sensitivity of the photoreceptor. Frequency response of the photoreceptor derived from narrow pulses of light predicts the frequency response obtained with sinusoidal stimuli, and the photoreceptor is linear for small signals in the light-adapted state.     

Pupil Dilation Dynamics Track Attention to High-Level Information

It has long been thought that the eyes index the inner workings of the mind. Consistent with this intuition, empirical research has demonstrated that pupils dilate as a consequence of attentional effort. Recently, Smallwood et al. (2011) demonstrated that pupil dilations not only provide an index of overall attentional effort, but are time-locked to stimulus changes during attention (but not during mind-wandering). This finding suggests that pupil dilations afford a dynamic readout of conscious information processing. However, because stimulus onsets in their study involved shifts in luminance as well as information, they could not determine whether this coupling of stimulus and pupillary dynamics reflected attention to low-level (luminance) or high-level (information) changes. Here, we replicated the methodology and findings of Smallwood et al. (2011) while controlling for luminance changes. When presented with isoluminant digit sequences, participants'' pupillary dilations were synchronized with stimulus onsets when attending, but not when mind-wandering. This replicates Smallwood et al. (2011) and clarifies their finding by demonstrating that stimulus-pupil coupling reflects online cognitive processing beyond sensory gain.     

Pupillenreaktionen auf sinusförmige Leuchtdichteänderungen

Summary Nonlinear properties of the human pupillary light reflex have been studied using binocular light stimuli of sinusoidally varying intensity. As reported earlier, the pupil responds with a net contraction to the modulation of the light intensity within a wide range of modulation frequencies. Results of the present experiments show that the net contraction does not depend upon the relative phase difference of the binocular light stimuli. However in the lower frequency range, where the in-phase stimuli result in a periodic light reflex, the out-of-phase stimuli fail to do so. These results allow certain conclusions to be drawn regarding the localisation within the photopupil reflex arc of the nonlinearities involved.     

Light adaptation in turtle cones. Testing and analysis of a model for phototransduction.

Light adaptation in cones was characterized by measuring the changes in temporal frequency responses to sinusoidal modulation of light around various mean levels spanning a range of four log units. We have shown previously that some aspects of cone adaptation behavior can be accounted for by a biochemical kinetic model for phototransduction in which adaptation is mediated largely by a sigmoidal dependence of guanylate cyclase activity on the concentration of free cytoplasmic Ca2+, ([Ca2+]i) (Sneyd and Tranchina, 1989). Here we extend the model by incorporating electrogenic Na+/K+ exchange, and the model is put to further tests by simulating experiments in the literature. It accounts for (a) speeding up of the impulse response, transition from monophasic to biphasic waveform, and improvement in contrast sensitivity with increasing background light level, I0; (b) linearity of the response to moderate modulations around I0; (c) shift of the intensity-response function (linear vs. log coordinates) with change in I0 (Normann and Perlman, 1979); the dark-adapted curve adheres closely to the Naka-Rushton equation; (d) steepening of the sensitivity vs. I0 function with [Ca2+]i fixed at its dark level, [Ca2+]i dark; (Matthews et al., 1988, 1990); (e) steepening of the steady-state intensity-response function when [Ca2+]i is held fixed at its dark level (Matthews et al., 1988; 1990); (f) shifting of a steep template saturation curve for normalized photocurrent vs. light-step intensity when the response is measured at fixed times and [Ca2+]i is held fixed at [Ca2+]i dark (Nakatani and Yau, 1988). Furthermore, the predicted dependence of guanylate cyclase activity on [Ca2+] closely matches a cooperative inhibition equation suggested by the experimental results of Koch and Stryer (1988) on cyclase activity in bovine rods. Finally, the model predicts that some changes in response kinetics with background light will still be present, even when [Ca2+]i is held fixed at [Ca2]i dark.     

Long rise times of sound pulses in grasshopper songs improve the directionality cues received by the CNS from the auditory receptors

Auditory receptors of the locust (Locusta migratoria) were investigated with respect to the directionality cues which are present in their spiking responses, with special emphasis on how directional cues are influenced by the rise time of sound signals. Intensity differences between the ears influence two possible cues in the receptor responses, spike count and response latency. Variation in rise time of sound pulses had little effect on the overall spike count; however, it had a substantial effect on the temporal distribution of the receptor's spiking response, especially on the latencies of first spikes. In particular, with ramplike stimuli the slope of the latency vs. intensity curves was steeper as compared to stimuli with steep onsets (Fig. 3). Stimuli with flat ramplike onsets lead to an increase of the latency differences of discharges between left and right tympanic receptors. This type of ramplike stimulus could thus facilitate directional hearing. This hypothesis was corroborated by a Monte Carlo simulation in which the probability of incorrect directional decisions was determined on the basis of the receptor latencies and spike counts. Slowly rising ramps significantly improved the decisions based on response latency, as compared to stimuli with sudden onsets (Fig. 4). These results are compared to behavioural results obtained with the grasshopper Ch. biguttulus. The stridulation signals of the females of this species consist of ramplike pulses, which could be an adaptation to facilitate directional hearing of phonotactically approaching males.Abbreviations HFR high frequency receptor - ILD interaural level difference - LFR low frequency receptor - SPL sound pressure level - WN white noise     

The oscillatory responses of skate electroreceptors to small voltage stimuli

Tonic nerve activity in skate electroreceptors is thought to result from spontaneous activity of the lumenal membranes of the receptor cells which is modulated by applied stimuli. When physiological conditions are simulated in vitro, the receptor epithelium produces a current which flows inward across the lumenal surface. This epithelial current exhibits small spontaneous sinusoidal fluctuations about the mean that are associated with corresponding but delayed fluctuations in postsynaptic response. Small voltage stimuli produce damped oscillations in the epithelial current similar in time-course to the spontaneous fluctuations. For lumen-negative, excitatory stimuli, these responses are predominantly an increase over the mean inward current. For inhibitory stimuli they are predominantly a decrease. Increased inward current across the lumenal membranes of the receptor cells increases depolarization of the presynaptic membranes in the basal faces leading to increased release of transmitter and an excitatory postsynaptic response. Decreased inward current decreases depolarization of the presynaptic membranes leading to a reduction in transmitter release and an inhibitory postsynaptic response. Clear changes in postsynaptic response are detectable during stimuli as small as 5 microV with saturation occurring at +/- 400 microV. The evoked oscillations in epithelial current are damped and the postsynaptic responses decline during maintained stimuli with large off-responses occurring at stimulus termination. The initial peak of the off-response is similar to the response produced by onset of an oppositely directed stimulus. These observations substantiate the role of receptor cell excitability in the detection of small voltage changes.     

Characterization of cytosolic calcium oscillations induced by phenylephrine and vasopressin in single fura-2-loaded hepatocytes

Changes of cytosolic free Ca2+ [( Ca2+]i) in response to receptor activation were studied at the single cell level by using digital imaging fluorescence microscopy of fura-2-loaded primary cultured hepatocytes. In response to phenylephrine and vasopressin, individual hepatocytes displayed dose-dependent oscillations of [Ca2+]i similar to those observed in aequorin-injected hepatocytes by Woods et al. (Woods, N. M., Cuthbertson, K. S. R., and Cobbold, P. H. (1986) Nature 329, 719-721). With increasing agonist concentration, the frequency of oscillations increased and the latent period decreased. For a given cell, peak [Ca2+]i was independent of applied agonist concentration. However, there was considerable variation from cell to cell in the absolute value of peak [Ca2+]i. There was also marked intercellular heterogeneity in the latency, frequency, and overall pattern of the Ca2+ responses. Such asynchronous responses can be explained in part by the apparent differential agonist sensitivity of individual cells for latency and frequency. At high doses, phenylephrine maintained an oscillatory pattern, whereas vasopressin produced a complex mixture of spiking and sustained [Ca2+]i responses. Vasopressin and phenylephrine also displayed differently shaped [Ca2+]i oscillations at submaximal doses, due primarily to a slower rate of decay with vasopressin. Despite the large cell-cell variation in the patterns of [Ca2+]i oscillations, successive readditions of the same agonist elicited identical cell-specific patterns of oscillation. In the absence of extracellular Ca2+ the frequency but not the magnitude of [Ca2+]i oscillations was decreased. Buffering of [Ca2+]i by increasing the fura-2 load of single hepatocytes also decreased the frequency of oscillations without affecting the peak Ca2+ level. These data provide further support for the importance of frequency modulation in agonist-induced Ca2+ responses and suggest that Ca2+ itself plays an important role in regulating the frequency of [Ca2+]i oscillations. Furthermore, the data demonstrate a broad heterogeneity in hepatocyte [Ca2+]i oscillations which may underlie the nonoscillatory responses of cell populations.     

Pupil Size Changes as an Active Information Channel for Biofeedback Applications

Pupil size is usually regarded as a passive information channel that provides insight into cognitive and affective states but defies any further control. However, in a recent study (Ehlers et al. 2015) we demonstrate that sympathetic activity indexed by pupil dynamics allows strategic interference by means of simple cognitive techniques. Utilizing positive/negative imaginings, subjects were able to expand pupil diameter beyond baseline variations; albeit with varying degrees of success and only over brief periods. The current study provides a comprehensive replication on the basis of considerable changes to the experimental set-up. Results show that stricter methodological conditions (controlled baseline settings and specified user instructions) strengthen the reported effect, whereas overall performance increases by one standard deviation. Effects are thereby not restricted to pupillary level. Parallel recordings of skin conductance changes prove a general enhancement of induced autonomic arousal. Considering the stability of the results across studies, we conclude that pupil size information exceeds affective monitoring and may constitute an active input channel in human–computer interaction. Furthermore, since variations in pupil diameter reliably display self-induced changes in sympathetic arousal, the relevance of this parameter is strongly indicated for future approaches in clinical biofeedback.