The Differential Contribution of Macular Pigments and Foveal Anatomy to the Perception of Maxwell's Spot and Haidinger's Brushes.

The relationship of macular pigments and foveal anatomy to the perception of Maxwell's spot (MS) and Haidinger's brushes (HB) entoptic phenomena were investigated. Dual-wavelength-autofluorescence and OCT were used to define macular pigment density and foveal anatomy in 52 eyes. MS was generated by alternating unpolarized red/blue and red/green uniform field illumination. HB was generated by alternating the linear polarization axis of a uniform blue field. In Experiment 1, horizontal widths of MS and HB were measured using a micrometer system and compared with macular pigment densities and OCT-defined morphometry. MS radius (mean 1.4°) was significantly less than HB radius (mean 1.6°), with the spatial extent of both phenomena falling between the boundaries of the foveola and foveal pit. Multiple regression showed MS and HB radii to be significantly associated with the macular pigment spatial profile radius. HB radius, but not MS radius, was also significantly associated with foveolar morphometry. Experiment 2 compared perceptual profiles of MS with macular pigment distribution patterns and demonstrated close agreement. The size and appearance of MS is a direct indicator of macular pigment density and distribution. Measures of HB radii are less specific, with dependence on both macular pigment density and foveal structure.

The Effect of Age-Related Macular Degeneration on Polarization Pattern Perception.

Purpose: The purpose of this study was to determine if a battery of polarization-modulated stimuli, quantified as a single metric, is effective in identifying macular disease in the presence/absence of cataract or pseudophakia. Methods: Using a modified liquid crystal display, polarization pattern perception (PPP) for a formulated battery of geometric and logMAR stimuli was evaluated in participants that had either no eye pathology (healthy participants) or were grouped according to the presence of cataract, pseudophakia, and/or age-related macular degeneration (AMD). PPP was quantified as response frequencies to individual stimuli, and as a novel monocular polarization sensitivity score (Ps) based on perception of the stimulus battery set. Results: Stimulus response frequencies were pattern-dependent and, compared with healthy participants, reduced for cataract and AMD groups but not for subjects with pseudophakia. Compared with healthy eyes (n = 47, median Ps = 17), Ps was significantly reduced by AMD (n = 59, median Ps = 1, P < 0.001) and, to a lesser extent, by cataracts (n = 80, median Ps = 6, P < 0.001). There was no significant difference between Ps for healthy and pseudophakic eyes (n = 47, median Ps = 13, P = 0.323). There was no significant correlation between Ps and logMAR visual acuity. Conclusions: In the absence of significant cataract, or in pseudophakia, a set of polarization-modulated visual stimuli, quantified as the Ps score, distinguishes AMD from healthy maculae. Translational Relevance: Perception of polarization-modulated stimuli, previously shown to be macula-dependent in a laboratory setting, is effective as a test of macular function in health and disease in a clinic setting.

Seeing polarization of light with the naked eye.

Many readers may know that scores of animal species sense the polarization of light for purposes including navigation, predation, and communication1. It is commonly thought that humans lack any sensitivity to polarization of light (e.g., Morehouse2). We hope to convince you otherwise by describing three examples where humans can detect polarization of light with the naked eye, by showing you how to see it yourself, and by describing its uses.

The Clinical Application of Polarization Pattern Perception.

Purpose: Determine the repeatability of and optimum stimulus parameters for testing polarization pattern perception in a real-world clinical population, and assess the ability of polarization perception to distinguish normal from abnormal eyes. Methods: Polarization perception was evaluated in staff and patients attending ophthalmology clinics at Warwick Hospital, UK. A series of visual stimuli were presented in pseudorandom order using a liquid-crystal-display-based polarization pattern generator. Stimuli included geometric patterns, gratings, checkerboards, and optotypes. Participants had one or both eyes diagnosed as normal or abnormal following ophthalmic examination, optical coherence tomography, and measures of visual acuity. Measurement scores were assigned to the eye(s) of each participant depending on the total number of stimuli perceived or identified. Results: Stimuli covered the range of spatial scales resolvable within polarization perception by normal and abnormal eyes. Different stimuli had different saliencies. For each stimulus type, polarization perception in the abnormal group was significantly reduced compared with normal eyes (P < 0.001). Relative stimulus salience was broadly similar for normal-eye and abnormal-eye viewing groups, being greatest for radially symmetric patterns and least for optotypes. Checkerboard pattern salience had an inverse logarithmic relationship with check fundamental spatial frequency. A devised metric covering the dynamic range of polarization perception was repeatable, and the score derived from the metric was reduced in the abnormal group compared with the normal group (P < 0.001). Conclusions: Clinically useful metrics of polarization perception distinguish between normal and abnormal eyes. Translational Relevance: Perception of spatial patterns formed of non-uniform polarization fields has potential as a quantitative clinical diagnostic measurement.

The electrophysiological response to polarization-modulated patterned visual stimuli.

Recent reports indicate that the subjective ability of humans to discriminate between polarization E-vector orientations approaches that of many invertebrates. Here, we show that polarization-modulated patterned stimuli generate an objectively recordable electrophysiological response in humans with normal vision. We investigated visual evoked potential (VEP) and electroretinographic (ERG) responses to checkerboard patterns defined solely by their polarization E-vector orientation alternating between ± 45°. Correcting for multiple comparisons, paired-samples t-tests were conducted to assess the significance of post-stimulus deflections from baseline measures of noise. Using standard check pattern sizes for clinical electrophysiology, and a pattern-reversal protocol, participants showed a VEP response to polarization-modulated patterns (PolVEP) with a prominent and consistent positive component near 150 ms (p < 0.01), followed by more variable negative components near 200 ms and 300 ms. The effect was unrecordable with visible wavelengths >550 nm. Further, pseudo-depolarization negated the responses, while control studies provided confirmatory evidence that the PolVEP response was not the product of luminance artefacts. Polarization-modulated patterns did not elicit a recordable ERG response. The possible origins of the PolVEP signals, and the absence of recordable ERG signals, are discussed. We conclude that evoked cortical responses to polarization-modulated patterns provide an objective measure of foveal function, suitable for both humans and non-human primates with equivalent macular anatomy.

Polarization perception in humans: on the origin of and relationship between Maxwell's spot and Haidinger's brushes.

Under specific conditions of illumination and polarization, differential absorption of light by macular pigments is perceived as the entoptic phenomena of Maxwell's spot (MS) or Haidinger's brushes (HB). To simulate MS and HB, an existing computational model of polarization-dependent properties of the human macula was extended by incorporating neuronal adaptation to stabilized retinal images. The model predicted that polarized light modifies the appearance of MS leading to the perception of a novel phenomenon. The model also predicted a correlation between the observed diameters of MS and HB. Predictions were tested psychophysically in human observers, whose measured differences in the diameters of each entoptic phenomenon generated with depolarized and linearly polarized light were consistent with the model simulations. These findings support a common origin of each phenomenon, and are relevant to the clinical use of polarization stimuli in detecting and monitoring human eye disorders, including macular degeneration. We conclude: (i) MS and HB both result from differential light absorption through a radial diattenuator, compatible with the arrangement of macular pigments in Henle fibres; (ii) the morphology of MS is dependent on the degree of linear polarization; (iii) perceptual differences between MS and HB result from different states of neural adaptation.

Computational simulation of human perception of spatially dependent patterns modulated by degree and angle of linear polarization.

Recent studies on polarization perception have shown that humans are sensitive to patterned stimuli modulated by either angle of linear polarization (AoP) or degree of polarization (DoP). Here, we present a model of human polarization sensitivity that incorporates both AoP and DoP as spatially dependent input variables. Applying the model to both sinusoidal- and square-wave-modulated DoP and AoP inputs, we demonstrate the theoretical similarities and differences generated by such inputs. Our model indicates the following: (i) edge boundaries between two adjacent areas of different linear polarization are preserved for both AoP- and DoP-modulated stimuli; and (ii) compared with DoP stimuli, AoP stimuli generate greater luminance changes at the photoreceptor level, suggesting that AoP-modulated patterns are potentially more salient than DoP patterns. The computational model is supported experimentally with an optical test of the model comprising a radial diattenuating polarizing filter and modified liquid crystal displays generating DoP- and AoP-modulated outputs. Psychophysical measures of human sensitivity confirm the increased salience of AoP- relative to DoP-modulated stimuli. These findings have practical application to the selection of DoP- and AoP-modulated stimuli for the investigation of macular function and macular pigment density in healthy and diseased eyes.

Haidinger's brushes elicited at varying degrees of polarization rapidly and easily assesses total macular pigmentation.

Macular pigments (MPs), by absorbing potentially toxic short-wavelength (400-500 nm) visible light, provide protection against photo-chemical damage thought to be relevant in the pathogenesis of age-related macular degeneration (AMD). A method of screening for low levels of MPs could be part of a prevention strategy for helping people to delay the onset of AMD. We introduce a new method for assessing MP density that takes advantage of the polarization-dependent absorption of blue light by MPs, which results in the entoptic phenomenon called Haidinger's brushes (HB). Subjects were asked to identify the direction of rotation of HB when presented with a circular stimulus illuminated with an even intensity of polarized white light in which the electric field vector was rotating either clockwise or anti-clockwise. By reducing the degree of polarization of the stimulus light, a threshold for perceiving HB (degree of polarization threshold) was determined and correlated (r2=0.66) to macular pigment optical density assessed using dual-wavelength fundus autofluoresence. The speed and ease of measurement of degree of polarization threshold makes it well suited for large-scale screening of macular pigmentation.

Computational simulation of Haidinger's brushes.

Haidinger's brushes (HB) are entoptic phenomena resulting from differential absorption of linear polarized light by the human macula. Computational models have assisted in understanding the behavior of these subjective phenomena but have been limited in their application. This study presents a revised computational model that incorporates known determinants of the form and behavior of HB. The model generates both static and animated simulations of HB that can be quantified by their density, contrast, and radial/circumferential extent. Measured physiological parameters are used to demonstrate the dependency of HB on macular pigment (MP) density, MP distribution, and ocular retardation. Physiological variations in these parameters explain the reported variations in the perception of HB.

The spectral, spatial and contrast sensitivity of human polarization pattern perception.

It is generally believed that humans perceive linear polarized light following its conversion into a luminance signal by diattenuating macular structures. Measures of polarization sensitivity may therefore allow a targeted assessment of macular function. Our aim here was to quantify psychophysical characteristics of human polarization perception using grating and optotype stimuli defined solely by their state of linear polarization. We show: (i) sensitivity to polarization patterns follows the spectral sensitivity of macular pigment; (ii) the change in sensitivity across the central field follows macular pigment density; (iii) polarization patterns are identifiable across a range of contrasts and scales, and can be resolved with an acuity of 15.4 cycles/degree (0.29 logMAR); and (iv) the human eye can discriminate between areas of linear polarization differing in electric field vector orientation by as little as 4.4°. These findings, which support the macular diattenuator model of polarization sensitivity, are unique for vertebrates and approach those of some invertebrates with a well-developed polarization sense. We conclude that this sensory modality extends beyond Haidinger's brushes to the recognition of quantifiable spatial polarization-modulated patterns. Furthermore, the macular origin and sensitivity of human polarization pattern perception makes it potentially suitable for the detection and quantification of macular dysfunction.

Human perception of visual stimuli modulated by direction of linear polarization.

This study explores both theoretically and experimentally the human perception of polarized light beyond that currently established. The radial analyser theory of Haidinger's phenomenon (HP) is used to predict the effect of observing visual stimuli comprising patterned zones characterized by orthogonal planes of linear polarization (linear polarization direction fields, LPD-fields). Any pattern can be represented as an LPD-field including optotypes and geometric forms. Simulated percepts differ from the original patterns although edges are mostly preserved. In edge-rich images a cross of attenuating contrast spanning the field of view is predicted. The mathematical model is verified experimentally using a liquid crystal display (LCD)-based polarization modulator imaged through a tangential (azimuthal) analyser with properties complementary to a radial analyser. The LCD device is then used in vivo to elicit perceptual responses in human subjects. Normal humans are found to readily detect spatially and temporally modulated isoluminant spatially-isochromatic, highly polarized LPD stimuli. Most subjects match the stimuli to corresponding images of theoretically predicted percepts. In particular edge perception and the presence of the contrast cross was confirmed. Unlike HP, static patterned LPD stimuli are perceived without difficulty. The simplest manifestation of human polarization perception is HP which is the fundamental element of an open set of stimulus-dependent percepts. This study demonstrates that humans have the ability to perceive and identify visual pattern stimuli defined solely by polarization state modulation.

The theory and implications of the biaxial model of corneal birefringence.

PURPOSE: A theoretical model of biaxial optical anisotropy is derived and its structural, biomechanical and developmental implications are discussed with reference to known corneal anatomy. METHODS A concise review of the theory of optical crystallography is followed by the derivation of a theoretical model of the optical anisotropic properties of a dome of biaxial birefringent crystalline material. The model is then applied to parameters relevant to the biaxial model of human corneal birefringence. RESULTS: Theoretical distributions of refractive indices and vibration directions for transmitted monochromatic light are derived for the central and paracentral zones of a model human cornea. Contours of equal refractive index (equirefringence curves) are found to have orthogonal confocal spheroconic geometry. CONCLUSIONS: A novel model of corneal structure is proposed in which discrete uniaxial positive birefringent fibre-like elements conform to the derived spheroconic geometry. Biomechanical implications and the relationship of the birefringent elements to known corneal anatomy are discussed. The crystallographic conventions are proposed as a standard for further investigations of corneal birefringence.

Circular polarization biomicroscopy: a method for determining human corneal stromal lamellar organization in vivo.

The theory of polarization biomicroscopy is explored using Stokes vectors and Mueller matrices. It is established that circular polarization can be used to simultaneously detect birefringent elements at any orientation unlike orientation-sensitive techniques using linear polarized light alone. A method of biomicroscopy using circular polarized light is described and tested in a physical model. The method is then used to investigate the lamellar structure of human corneas in vivo in pairs of eyes of 38 subjects. An approximate confocal elliptic/hyperbolic distribution of stromal fibrils, presumed to be collagen, is clearly identified within central and intermediate areas of the cornea. All subjects tested demonstrate approximate mirror symmetry between pairs of eyes typically with a preferred orientation of central fibrils at approximately 15 degrees to the horizontal in a superotemporal-inferonasal direction.

A Mueller matrix model of Haidinger's brushes.

Stokes vectors and Mueller matrices are used to model the polarisation properties (birefringence, dichroism and depolarisation) of any optical system, in particular the human eye. An explanation of the form and behaviour of the entoptic phenomenon of Haidinger's brushes is derived that complements and expands upon a previous study. The relationship between the appearance of Haidinger's brushes and intrinsic ocular retardation is quantified and the model allows prediction of the effect of any retarder of any orientation placed between a source of polarised light and the eye. The simple relationship of minimum contrast of Haidinger's brushes to the cosine of total retardation is derived.