Circular polarization vision in a stomatopod crustacean

We describe the addition of a fourth visual modality in the animal kingdom, the perception of circular polarized light. Animals are sensitive to various characteristics of light, such as intensity, color, and linear polarization [1, 2]. This latter capability can be used for object identification, contrast enhancement, navigation, and communication through polarizing reflections [2-4]. Circularly polarized reflections from a few animal species have also been known for some time [5, 6]. Although optically interesting [7, 8], their signal function or use (if any) was obscure because no visual system was known to detect circularly polarized light. Here, in stomatopod crustaceans, we describe for the first time a visual system capable of detecting and analyzing circularly polarized light. Four lines of evidence-behavior, electrophysiology, optical anatomy, and details of signal design-are presented to describe this new visual function. We suggest that this remarkable ability mediates sexual signaling and mate choice, although other potential functions of circular polarization vision, such as enhanced contrast in turbid environments, are also possible [7, 8]. The ability to differentiate the handedness of circularly polarized light, a visual feat never expected in the animal kingdom, is demonstrated behaviorally here for the first time.     

Color and polarization vision in foraging Papilio

This paper gives an overview of behavioral studies on the color and polarization vision of the Japanese yellow swallowtail butterfly, Papilio xuthus. We focus on indoor experiments on foraging individuals. Butterflies trained to visit a disk of certain color correctly select that color among various other colors and/or shades of gray. Correct selection persists under colored illumination, but is systematically shifted by background colors, indicating color constancy and simultaneous color contrast. While their eyes contain six classes of spectral receptors, their wavelength discrimination performance indicates that their color vision is tetrachromatic. P. xuthus innately prefers brighter targets, but can be trained to select dimmer ones under certain conditions. Butterflies trained to a dark red stimulus select an orange disk presented on a bright gray background over one on dark gray. The former probably appears darker to them, indicating brightness contrast. P. xuthus has a strong innate preference for vertically polarized light, but the selection of polarized light changes depending on the intensity of simultaneously presented unpolarized light. Discrimination of polarization also depends on background intensity. Similarities between brightness and polarization vision suggest that P. xuthus perceive polarization angle as brightness, such that vertical polarization appears brighter than horizontal polarization.     

Behavioural relevance of polarization sensitivity as a target detection mechanism in cephalopods and fishes

Aquatic habitats are rich in polarized patterns that could provide valuable information about the environment to an animal with a visual system sensitive to polarization of light. Both cephalopods and fishes have been shown to behaviourally respond to polarized light cues, suggesting that polarization sensitivity (PS) may play a role in improving target detection and/or navigation/orientation. However, while there is general agreement concerning the presence of PS in cephalopods and some fish species, its functional significance remains uncertain. Testing the role of PS in predator or prey detection seems an excellent paradigm with which to study the contribution of PS to the sensory assets of both groups, because such behaviours are critical to survival. We developed a novel experimental set-up to deliver computer-generated, controllable, polarized stimuli to free-swimming cephalopods and fishes with which we tested the behavioural relevance of PS using stimuli that evoke innate responses (such as an escape response from a looming stimulus and a pursuing behaviour of a small prey-like stimulus). We report consistent responses of cephalopods to looming stimuli presented in polarization and luminance contrast; however, none of the fishes tested responded to either the looming or the prey-like stimuli when presented in polarization contrast.     

Optical polarization properties of the diffraction spectra from single fibers of skeletal muscle.

The diffraction spectra of laser light from single fibers of skeletal muscle exhibit a large degree of optical depolarization. When the linearly polarized incident laser source is oriented at polarization angles between 0 less than theta less than pi/2 rad with respect to the fiber axis, the diffracted light is elliptically polarized. These results show that the phase angle of the ellipse rotates by as much as 20 degrees when the fiber is stretched from 2.4 to 3.8 microns. To further ascertain that the observed phenomenon is diffraction related, an experiment monitoring the spectra of scattered light in between diffraction orders showed this signal to be significantly more linearly polarized. These results suggest that the degree of elliptical polarization of the diffraction spectra is a sensitive probe of A-band dynamics, including changes of the anisotropic S-2 elements.     

Skylight polarization as perceived by desert ants and measured by video polarimetry

North African desert ants belonging to different genera and inhabiting different areas (sand dunes, salt pans, inundation flats and gravel plains) exhibit different ways of skylight navigation: some rely especially on the polarized light in the sky, others depend more effectively on the position of the sun. Are these differences due to species- or genus-specific idiosyncrasies of the ant's skylight compass, or are they caused by differences in the overall degree of polarization prevailing in the celestial hemisphere that vaults the different kinds of habitat? Theoretically, such differences are to be expected, as various parameters known to influence the degree of polarization in the Earth's atmosphere – such as the albedo of the ground and the content of water vapour, dust and haze in the airlayers above the ground – do vary between the different types of habitat mentioned above. The first wide-field, video-polarimetric study of skylight polarization presented here clearly shows that at any particular locality the temporal (day-to-day) variations of the degree of skylight polarization are much more pronounced than the differences recorded at the same local time at different localities. In contrast, the angle of polarization is unaffected by atmospheric disturbances and accords well with the predictions of Rayleigh scattering. Consequently, differences in behavioural performances of navigating North African desert ants are due to interspecific and intergeneric differences in the ants' navigational systems rather than to general differences in the skylight stimuli experienced by the ants during navigation.     

The polarization of fluorescence of DNA stains depends on the incorporation density of the dye molecules.

BACKGROUND: The fluorescence induced by polarized light sources, such as the lasers that are used in flow cytometry, is often polarized and anisotropic. In addition, most optical detector systems are sensitive to the direction of polarization. These two factors influence the accuracy of fluorescence intensity measurements. The intensity of two light sources can be compared only if all details of the direction and degree of polarization are known. In a previous study, we observed that fluorescence polarization might be modified by dye-dye interactions. This report further investigates the role of dye density in fluorescence polarization anisotropy. METHODS: We measured the polarization distribution of samples stained with commonly used DNA dyes. To determine the role of fluorophore proximity, we compared the monomeric and a dimeric form of the DNA dyes ethidium bromide (EB), thiazole orange (TO), and oxazole yellow (YO). RESULTS: In all dyes sampled, fluorescence polarization is less at high dye concentrations than at low concentrations. The monomeric dyes exhibit a higher degree of polarization than the dimeric dyes of the same species. CONCLUSIONS: The polarization of fluorescence from DNA dyes is related to the density of incorporation into the DNA helix. Energy transfer between molecules that are in close proximity loosens the linkage between the excitation and emission dipoles, thereby reducing the degree of polarization of the emission.     

Modulated polarization microscopy: a promising new approach to visualizing cytoskeletal dynamics in living cells

In an effort to visualize cytoskeletal filaments in living cells, we have developed modulated polarization microscopy. Modulated polarization microscopy visualizes cytoskeletal filaments based on their birefringence but differs from the standard polarization microscopy by exploiting the angle dependence of birefringence. A prototype instrument has been developed using two Faraday rotators under computer control to change the angle of plane polarized light at a known rate. By placing one Faraday rotator before and one after the specimen, rotation produced by the first Faraday rotator is cancelled by the second. This allows the use of fixed polarizer and analyzer in a crossed configuration and continuous imaging of the specimen between crossed polarizers. The variation in polarization angle of light illuminating the specimen causes birefringent elements to oscillate in brightness. Images acquired as polarization angle is varied are then processed by a Fourier filter image-processing algorithm. The Fourier filtering algorithm isolates those signals that vary at the proper rate, whereas static or random signals are removed. Here we show that the modulated polarization microscope can reveal cytoskeletal elements including stress fibers and microtubules in living cells.     

Frequency-Dependent Polarization Properties of Local Electric Field in Gold–Dielectric Multi-Nanoshells

The polarization properties of the local electric field in the gold–dielectric–gold multilayer nanoshells are investigated by theoretical calculation based on the quasi-static approximation. The calculation results show that the complete polarized incident light does not only stimulate the same directional polarized local electric field. The polarized angle of the local field may changes from 0° to 90° as the wavelength and location are changed. The distributions of local field polarization are different in dielectric layer or gold shell and display different features in different plasmonic hybridization mode. As the incident wavelength is increased, the hot spot of polarizing angle moves monotonously in the middle dielectric shell, whereas moves nonmonotonously in the gold shell and surrounding environment. In the gold shell, the gap between hot spots of polarizing angle may occur at the resonance frequency. However, the hot spots of polarizing angle always occur at the resonance frequencies in the surrounding environment. These interesting results show that the single-molecule detection based on metal nanostructure induced surface-enhanced Raman scattering and surface enhanced fluorescence could be optimized by adjusting the incident light polarization and frequency.     

Retinal ultrastructure may mediate polarization sensitivity in larvae of the Sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae)

A number of invertebrates are known to be sensitive to the polarization of light and use this trait in orientation, communication, or prey detection. In these animals polarization sensitivity tends to originate in rhabdomeric photoreceptors that are more or less uniformly straight and parallel. Typically, polarization sensitivity is based on paired sets of photoreceptors with orthogonal orientation of their rhabdomeres. Sunburst diving beetle larvae are active swimmers and highly visual hunters which could potentially profit from polarization sensitivity. These larvae, like those of most Dytiscids, have a cluster of six lens eyes or stemmata (designated E1 through E6) on each side of the head capsule. We examined the ultrastructure of the photoreceptor cells of the principal eyes (E1 and E2) of first instar larvae to determine whether their rhabdomeric organization could support polarization sensitivity. A detailed electron microscopical study shows that the proximal retinas of E1 and E2 are in fact composed of photoreceptors with predominantly parallel microvilli and that neighboring rhabdomeres are oriented approximately perpendicularly to one another. A similar organization is observed in the medial retina of E1, but not in the distal retinas of E1&2. Our findings suggest that T. marmoratus larvae might be able to analyze polarized light. If so, this could be used by freshly hatched larvae to find water or within the water to break the camouflage of common prey items such as mosquito larvae. Physiological and behavioral tests are planned to determine whether larvae of T. marmoratus can actually detect and exploit polarization signals.     

Behavioural evidence for polarization vision in crickets

ABSTRACT. Tethered field crickets, Gryllus campestris L., walking on an air-suspended bail exhibit a spontaneous response to the e-vector of polarized light presented from above: E-vector orientation controls strength and direction of turning tendency. Experiments in which different eye regions are covered with paint suggest that this response is mediated by the anatomically and physiologically specialized dorsal rim area of the compound eye. We conclude that crickets have polarization vision and that the dorsal rim area of the eye plays a key role in this sensory capacity.     

Behavioral analysis of polarization vision in tethered flying locusts

For spatial navigation many insects rely on compass information derived from the polarization pattern of the sky. We demonstrate that tethered flying desert locusts (Schistocerca gregaria) show e-vector-dependent yaw-torque responses to polarized light presented from above. A slowly rotating polarizer (5.3° s^–1) induced periodic changes in yaw torque corresponding to the 180° periodicity of the stimulus. Control experiments with a rotating diffuser, a weak intensity pattern, and a stationary polarizer showed that the response is not induced by intensity gradients in the stimulus. Polarotaxis was abolished after painting the dorsal rim areas of the compound eyes black, but remained unchanged after painting the eyes except the dorsal rim areas. During rotation of the polarizer, two e-vectors (preferred and avoided e-vector) induced no turning responses: they were broadly distributed from 0 to 180° but, for a given animal, were perpendicular to each other. The data demonstrate polarization vision in the desert locust, as shown previously for bees, flies, crickets, and ants. Polarized light is perceived through the dorsal rim area of the compound eye, suggesting that polarization vision plays a role in compass navigation of the locust.     

Why do dusk-active cockchafers detect polarization in the green? The polarization vision in Melolontha melolontha is tuned to the high polarized intensity of downwelling light under canopies during sunset

In the retina of dusk-active European cockchafers, Melolontha melolontha, the linear polarization of downwelling light (skylight or light from the tree canopy) is detected by photoreceptors in upward-pointing ommatidia with maximal sensitivity at 520 nm in the green portion of the spectrum. To date no attempt has been made to answer the question of why these beetles detect polarization in the green. Here we present an atmospheric optical and receptor-physiological explanation of why longer wavelengths are advantageous for the perception of polarization of downwelling light under canopies illuminated by the setting sun. Our explanation focuses on illumination situations during sunset in canopied optical environments, because cockchafers are active at sunset and fly predominantly under canopies during their swarming, feeding, and mating periods. Using three simple atmospheric optical models, we computed the degree of linear polarization, the linearly polarized intensity of downwelling light, the quantum catch, and quantum catch difference between polarization detectors with orthogonal microvilli under canopies illuminated by the setting sun as functions of wavelength and solar zenith angle. Based upon these computations, we show that the green sensitivity of polarization detectors in M. melolontha is tuned to the high polarized intensity of downwelling light in the green under canopies during sunset, an optimal compromise between simultaneous maximization of the quantum catch and the quantum catch difference. We also briefly discuss how green-sensitive polarization detectors can function efficiently enough during the pre-feeding and egg-laying flights of cockchafers, which always occur prior to sunset and under the sky.     

Circular Polarization of Transmitted Light by Sapphirinidae Copepods

Circularly polarized light, rare in the animal kingdom, has thus far been documented in only a handful of animals. Using a rotating circular polarization (CP) analyzer we detected CP in linearly polarized light transmitted through epipelagic free living Sapphirina metallina copepods. Both left and right handedness of CP was detected, generated from specific organs of the animal''s body, especially on the dorsal cephalosome and prosome. Such CP transmittance may be generated by phase retardance either in the muscle fibers or in the multilayer membrane structure found underneath the cuticle. Although the role, if any, played by circularly polarized light in Sapphirinidae has yet to be clarified, in other animals it was suggested to take part in mate choice, species recognition, and other forms of communication.

Highlights

Planktonic Sapphirinidae copepods were found to circularly polarize the light passing through them. Circular polarization may be created by unique, multilayered features of the membrane structure found under their cuticle or by organized muscle fibers.     

Polarization sensitivity as a contrast enhancer in pelagic predators: lessons from in situ polarization imaging of transparent zooplankton

Because light in the pelagic environment is partially polarized, it has been suggested that the polarization sensitivity found in certain pelagic species may serve to enhance the contrast of their transparent zooplankton prey. We examined its potential during cruises in the Gulf of Mexico and Atlantic Ocean and at a field station on the Great Barrier Reef. First, we collected various species of transparent zooplankton and micronekton and photographed them between crossed polarizers. Many groups, particularly the cephalopods, pelagic snails, salps and ctenophores, were found to have ciliary, muscular or connective tissues with striking birefringence. In situ polarization imagery of the same species showed that, while the degree of underwater polarization was fairly high (approx. 30% in horizontal lines of sight), tissue birefringence played little to no role in increasing visibility. This is most likely due to the low radiance of the horizontal background light when compared with the downwelling irradiance. In fact, the dominant radiance and polarization contrasts are due to unpolarized downwelling light that has been scattered from the animal viewed against the darker and polarized horizontal background light. We show that relatively simple algorithms can use this negative polarization contrast to increase visibility substantially.     

Perception of polarization plane,colour and movement in two dimensions by the crab,Carcinus

Summary The spectral sensitivity of movement perception in Carcinus has a peak near 508 m, irrespective of the direction of the movement or of the plane of polarization of the stimulating light, although the primary sensory cells are 10 times as sensitive to light in one plane of polarization as in the plane at right angles. This balance in the input to the movement perception system breaks down with a stimulating light at <439 and at >650 m. Only one set of connexions between the primary receptors in the movement perception system agrees with all the results of manipulating the various attributes of the visual input, and this model has unique properties which allow the subsequent discrimination of each attribute of the stimulus independent of the other attributes.I would like to thank Robert Santer for his assistance with the numerous experimental observations which form the basis of this paper.     

Evidence for true polarization vision based on a two-channel analyzer system in the eye of the water bug,Notonecta glauca

Summary Water bugs (Notonecta glauca) were set into flight in a room with a homogeneously illuminated ceiling and a light-emitting platform on the floor. In these conditions polarized UV light from the platform was more effective in causing the animals to fly down to the surface of the platform than was unpolarized UV light several times as intense. Experiments with an array of baffles that restricted the directions from which the polarization film on the platform could be seen showed that the polarized UV light is effective in eliciting descent only when the e-vector is perpendicular to the median sagittal plane of the animal (horizontal). It can be concluded that polarized UV light with horizontal e-vector is distinguished, as a special sensory quality, from unpolarized UV light.Notonecta thus provides an example of true polarization vision.The special orthogonal arrangement of the microvilli in the rhabdomeres of the UV visual cells in the ventral part of the eye (cf. Schwind 1983 b and Schwind et al., in press) is suggestive with regard to polarization vision. The microvilli of the two UV visual cells in the ommatidia looking forward and down are horizontal and vertical, respectively, and hence could serve as a two-channel analyzer system capable of distinguishing the polarized UV light reflected by a water surface from unpolarized UV light.     

Cone photoreceptor mechanisms and the detection of polarized light in fish

Summary Although numerous studies have demonstrated the detection of polarized light in vertebrates, little is known of the photoreceptor mechanisms involved. Recent evidence, however, indicates that cyprinid fishes possess both ultraviolet (UV) and polarization sensitivity suggesting that some vertebrates, like many invertebrates, may employ UV-sensitive cone receptors in polarization sensitivity. In this report, we describe experiments that determine which spectral types of receptors participate in the detection of polarized light. We used a heart-rate conditioning technique to measure increment thresholds of immobilized goldfish for plane-polarized, narrow-band (10 nm half max.) spectral stimuli (380 nm, 460 nm, 540 nm, 660 nm). A typical experiment involved isolating the activity of a cone photoreceptor mechanism by chromatic adaptation and measuring increment thresholds for spectral stimuli at e-vector orientations of the polarizer between 0° to 180° in 30° steps. The UV-, green- and red-sensitive cone receptor mechanisms showed clear evidence of polarization sensitivity while the blue-sensitive cone receptor mechanism was polarizationally insensitive. The average amplitude (base to peak height on Fig. 4) of the polarization sensitivity curves (UV-, green- and red-curves) was 0.67 log unit (standard deviation of 0.12 log unit), with the UV-sensitive cone receptor mechanism most sensitive to the vertical e-vector axis and the green- and red-sensitive cone receptor mechanisms most sensitive to the horizontal e-vector axis. The observation that different cone photoreceptor mechanisms have orthogonal polarization sensitivity in fish suggests that the perception of polarized light may enhance the capacity for visual discrimination in lower vertebrates.     

Mueller matrix decomposition for polarized light assessment of biological tissues

The Mueller matrix represents the transfer function of an optical system in its interactions with polarized light and its elements relate to specific biologically or clinically relevant properties. However, when many optical polarization effects occur simultaneously, the resulting matrix elements represent several “lumped” effects, thus hindering their unique interpretation. Currently, no methods exist to extract these individual properties in turbid media. Here, we present a novel application of a Mueller matrix decomposition methodology that achieves this objective. The methodology is validated theoretically via a novel polarized‐light propagation model, and experimentally in tissue simulating phantoms. The potential of the approach is explored for two specific biomedical applications: monitoring of changes in myocardial tissues following regenerative stem cell therapy, through birefringence‐induced retardation of the light's linear and circular polarizations, and non‐invasive blood glucose measurements through chirality‐induced rotation of the light's linear polarization. Results demonstrate potential for both applications. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Development of a polarization imaging method to detect paraffin-embedded pathology tissues before applying other techniques

The present article describes the development of a technique, applied to paraffin-embedded tissues, which uses three different wavelengths of monochromatic light (λ1 = 445 nm, λ2 = 540 nm and λ3 = 660 nm) for the measures of the degree of polarization, degree of linear polarization, degree of circular polarization and birefringence, all obtained from measurements of Stokes parameters by using polarized light. The goal of this study was to detect changes in developing embryonic mouse eye when pregnant mice fed diets without folic acid for variable periods compared with a healthy control group. We present a biomedical diagnostic technique based on polarized light detection applied to paraffin-embedded tissues to visualize the structural damage to aid us in the diagnosis before applying other techniques. Through this method, we can visualize and identify which parts of the tissue were altered with respect to the control group.     

Optical depolarization changes in single, skinned muscle fibers. Evidence for cross-bridge involvement.

Optical ellipsometry studies of single, skinned muscle fibers conducted on the diffraction orders have yielded spectra that are sensitive to the state of the fiber. The linearly polarized light field vector becomes elliptically polarized as it passes through the fiber and may be collected at the diffraction orders. Fibers that have been subjected to extraction of myosin (0.6 M KCl) retain a weak diffraction pattern and exhibit a substantially decreased depolarization of incident linearly polarized light. A significant decrease in polarization is seen in skinned fibers that are subject to an increase in pH from 7.0 to 8.0. This increase in pH results in a decrease of approximately 30% in the depolarization angle of single fibers. The major decrease in depolarization angle that we observe at pH 8.0 is consistent with the notion that as cross-bridges move out from the shaft of the thick filament, their ability to cause depolarization of the incident linearly polarized light decreases. This interpretation is also consistent with the work of Ueno and Harrington where the decrease in the ability to cross-link S-1 and S-2 to the thick filament at pH 8.2 suggests cross-bridge movement away from the thick filament. A large decrease in birefringence, seen after treatment of skinned fibers with alpha-chymotrypsin, appears to be related to the breakdown of myosin into rod, S-1, heavy meromyosin, and light meromyosin.