Why do horseflies need polarization vision for host detection? Polarization helps tabanid flies to select sunlit dark host animals from the dark patches of the visual environment.

Horseflies (Tabanidae) are polarotactic, being attracted to linearly polarized light when searching for water or host animals. Although it is well known that horseflies prefer sunlit dark and strongly polarizing hosts, the reason for this preference is unknown. According to our hypothesis, horseflies use their polarization sensitivity to look for targets with higher degrees of polarization in their optical environment, which as a result facilitates detection of sunlit dark host animals. In this work, we tested this hypothesis. Using imaging polarimetry, we measured the reflection-polarization patterns of a dark host model and a living black cow under various illumination conditions and with different vegetation backgrounds. We focused on the intensity and degree of polarization of light originating from dark patches of vegetation and the dark model/cow. We compared the chances of successful host selection based on either intensity or degree of polarization of the target and the combination of these two parameters. We show that the use of polarization information considerably increases the effectiveness of visual detection of dark host animals even in front of sunny-shady-patchy vegetation. Differentiation between a weakly polarizing, shady (dark) vegetation region and a sunlit, highly polarizing dark host animal increases the efficiency of host search by horseflies.

Mayflies are least attracted to vertical polarization: A polarotactic reaction helping to avoid unsuitable habitats.

Like other aquatic insects, mayflies are positively polarotactic and locate water surfaces by means of the horizontal polarization of water-reflected light. However, may vertically polarized light also have implications for the swarming behaviour of mayflies? To answer this question, we studied in four field experiments the behavioural responses of Ephoron virgo and Caenis robusta mayflies to lamps emitting horizontally and vertically polarized and unpolarized light. In both species, unpolarized light induces positive phototaxis, horizontally polarized light elicits positive photo- and polarotaxis, horizontally polarized light is much more attractive than unpolarized light, and vertically polarized light is the least attractive if the stimulus intensities and spectra are the same. Vertically polarized light was the most attractive for C. robusta if its intensity was about two and five times higher than that of the unpolarized and horizontally polarized stimuli, respectively. We suggest that the mayfly behaviour observed in our experiments may facilitate the stability of swarming above water surfaces. Beside the open water surface reflecting horizontally polarized light, the shadow and mirror image of riparian vegetation at the edge of the water surface reflect weakly and non-horizontally (mainly vertically) polarized light. Due to their positive polarotaxis, flying mayflies remain continuously above the water surface, because they keep away from the unpolarized or non-horizontally polarizing edge regions (water surface and coast line) of water bodies. We also discuss how our findings can explain the regulation of mayfly colonization.

Testing avian compass calibration: comparative experiments with diurnal and nocturnal passerine migrants in South Sweden.

Cue-conflict experiments were performed to study the compass calibration of one predominantly diurnal migrant, the dunnock (Prunella modularis), and two species of nocturnal passerine migrants, the sedge warbler (Acrocephalus schoenobaenus), and the European robin (Erithacus rubecula) during autumn migration in South Sweden. The birds' orientation was recorded in circular cages under natural clear and simulated overcast skies in the local geomagnetic field, and thereafter the birds were exposed to a cue-conflict situation where the horizontal component of the magnetic field (mN) was shifted +90° or -90° at two occasions, one session starting shortly after sunrise and the other ca. 90 min before sunset and lasting for 60 min. The patterns of the degree and angle of skylight polarization were measured by full-sky imaging polarimetry during the cue-conflict exposures and orientation tests. All species showed orientation both under clear and overcast skies that correlated with the expected migratory orientation towards southwest to south. For the European robin the orientation under clear skies was significantly different from that recorded under overcast skies, showing a tendency that the orientation under clear skies was influenced by the position of the Sun at sunset resulting in more westerly orientation. This sun attraction was not observed for the sedge warbler and the dunnock, both orientating south. All species showed similar orientation after the cue-conflict as compared to the preferred orientation recorded before the cue-conflict, with the clearest results in the European robin and thus, the results did not support recalibration of the celestial nor the magnetic compasses as a result of the cue-conflict exposure.

Acquisition and visualization techniques for narrow spectral color imaging.

This paper introduces a new approach in narrow-band imaging (NBI). Existing NBI techniques generate images by selecting discrete bands over the full visible spectrum or an even wider spectral range. In contrast, here we perform the sampling with filters covering a tight spectral window. This image acquisition method, named narrow spectral imaging, can be particularly useful when optical information is only available within a narrow spectral window, such as in the case of deep-water transmittance, which constitutes the principal motivation of this work. In this study we demonstrate the potential of the proposed photographic technique on nonunderwater scenes recorded under controlled conditions. To this end three multilayer narrow bandpass filters were employed, which transmit at 440, 456, and 470 nm bluish wavelengths, respectively. Since the differences among the images captured in such a narrow spectral window can be extremely small, both image acquisition and visualization require a novel approach. First, high-bit-depth images were acquired with multilayer narrow-band filters either placed in front of the illumination or mounted on the camera lens. Second, a color-mapping method is proposed, using which the input data can be transformed onto the entire display color gamut with a continuous and perceptually nearly uniform mapping, while ensuring optimally high information content for human perception.

Spottier targets are less attractive to tabanid flies: on the tabanid-repellency of spotty fur patterns.

During blood-sucking, female members of the family Tabanidae transmit pathogens of serious diseases and annoy their host animals so strongly that they cannot graze, thus the health of the hosts is drastically reduced. Consequently, a tabanid-resistant coat with appropriate brightness, colour and pattern is advantageous for the host. Spotty coats are widespread among mammals, especially in cattle (Bos primigenius). In field experiments we studied the influence of the size and number of spots on the attractiveness of test surfaces to tabanids that are attracted to linearly polarized light. We measured the reflection-polarization characteristics of living cattle, spotty cattle coats and the used test surfaces. We show here that the smaller and the more numerous the spots, the less attractive the target (host) is to tabanids. We demonstrate that the attractiveness of spotty patterns to tabanids is also reduced if the target exhibits spottiness only in the angle of polarization pattern, while being homogeneous grey with a constant high degree of polarization. Tabanid flies respond strongly to linearly polarized light, and we show that bright and dark parts of cattle coats reflect light with different degrees and angles of polarization that in combination with dark spots on a bright coat surface disrupt the attractiveness to tabanids. This could be one of the possible evolutionary benefits that explains why spotty coat patterns are so widespread in mammals, especially in ungulates, many species of which are tabanid hosts.

No evidence for behavioral responses to circularly polarized light in four scarab beetle species with circularly polarizing exocuticle.

The strongest known circular polarization of biotic origin is the left-circularly polarized (LCP) light reflected from the metallic shiny exocuticle of certain beetles of the family Scarabaeidae. This phenomenon has been discovered by Michelson in 1911. Although since 1955 it has been known that the human eye perceives a visual illusion when stimulated by circularly polarized (CP) light, it was discovered only recently that a stomatopod shrimp is able to perceive circular polarization. It is pertinent to suppose that scarab beetles reflecting LCP light in an optical environment (vegetation) being deficient in CP signals may also perceive circular polarization and use it to find each other (mate/conspecifics) as until now it has been believed. We tested this hypothesis in six choice experiments with several hundred individuals of four scarab species: Anomala dubia, Anomala vitis (Coleoptera, Scarabaeidae, Rutelinae), and Cetonia aurata, Potosia cuprea (Coleoptera, Scarabaeidae, Cetoniinae), all possessing left-circularly polarizing exocuticle. From the results of our experiments we conclude that the studied four scarab species are not attracted to CP light when feeding or looking for mate or conspecifics. We demonstrated that the light reflected by host plants of the investigated scarabs is circularly unpolarized. Our results finally solve a puzzle raised over one hundred years ago, when Michaelson discovered that scarab beetles reflect circularly polarized light.

Imaging polarimetry of the fogbow: polarization characteristics of white rainbows measured in the high Arctic.

The knowledge on the optics of fogbows is scarce, and their polarization characteristics have never been measured to our knowledge. To fill this gap we measured the polarization features of 16 fogbows during the Beringia 2005 Arctic polar research expedition by imaging polarimetry in the red, green and blue spectral ranges. We present here the first polarization patterns of the fogbow. In the patterns of the degree of linear polarization p, fogbows and their supernumerary bows are best visible in the red spectral range due to the least dilution of fogbow light by light scattered in air. In the patterns of the angle of polarization α fogbows are practically not discernible because their α-pattern is the same as that of the sky: the direction of polarization is perpendicular to the plane of scattering and is parallel to the arc of the bow, independently of the wavelength. Fogbows and their supernumeraries were best seen in the patterns of the polarized radiance. In these patterns the angular distance δ between the peaks of the primary and the first supernumerary and the angular width σ of the primary bow were determined along different radii from the center of the bow. δ ranged between 6.08° and 13.41°, while σ changed from 5.25° to 19.47°. Certain fogbows were relatively homogeneous, meaning small variations of δ and σ along their bows. Other fogbows were heterogeneous, possessing quite variable δ- and σ-values along their bows. This variability could be a consequence of the characteristics of the high Arctic with open waters within the ice shield resulting in the spatiotemporal change of the droplet size within the fog.

On the trail of Vikings with polarized skylight: experimental study of the atmospheric optical prerequisites allowing polarimetric navigation by Viking seafarers.

Between AD 900 and AD 1200 Vikings, being able to navigate skillfully across the open sea, were the dominant seafarers of the North Atlantic. When the Sun was shining, geographical north could be determined with a special sundial. However, how the Vikings could have navigated in cloudy or foggy situations, when the Sun's disc was unusable, is still not fully known. A hypothesis was formulated in 1967, which suggested that under foggy or cloudy conditions, Vikings might have been able to determine the azimuth direction of the Sun with the help of skylight polarization, just like some insects. This hypothesis has been widely accepted and is regularly cited by researchers, even though an experimental basis, so far, has not been forthcoming. According to this theory, the Vikings could have determined the direction of the skylight polarization with the help of an enigmatic birefringent crystal, functioning as a linearly polarizing filter. Such a crystal is referred to as 'sunstone' in one of the Viking's sagas, but its exact nature is unknown. Although accepted by many, the hypothesis of polarimetric navigation by Vikings also has numerous sceptics. In this paper, we summarize the results of our own celestial polarization measurements and psychophysical laboratory experiments, in which we studied the atmospheric optical prerequisites of possible sky-polarimetric navigation in Tunisia, Finland, Hungary and the high Arctic.

An unexpected advantage of whiteness in horses: the most horsefly-proof horse has a depolarizing white coat.

White horses frequently suffer from malign skin cancer and visual deficiencies owing to their high sensitivity to the ultraviolet solar radiation. Furthermore, in the wild, white horses suffer a larger predation risk than dark individuals because they can more easily be detected. In spite of their greater vulnerability, white horses have been highly appreciated for centuries owing to their natural rarity. Here, we show that blood-sucking tabanid flies, known to transmit disease agents to mammals, are less attracted to white than dark horses. We also demonstrate that tabanids use reflected polarized light from the coat as a signal to find a host. The attraction of tabanids to mainly black and brown fur coats is explained by positive polarotaxis. As the host's colour determines its attractiveness to tabanids, this parameter has a strong influence on the parasite load of the host. Although we have studied only the tabanid-horse interaction, our results can probably be extrapolated to other host animals of polarotactic tabanids, as the reflection-polarization characteristics of the host's body surface are physically the same, and thus not species-dependent.

Spatiotemporal change of sky polarization during the total solar eclipse on 29 March 2006 in Turkey: polarization patterns of the eclipsed sky observed by full-sky imaging polarimetry.

Using 180 degrees field-of-view (full-sky) imaging polarimetry, we measured the spatiotemporal change of the polarization of skylight during the total solar eclipse on 29 March 2006 in Turkey. We present our observations here on the temporal variation of the celestial patterns of the degree p and angle alpha of linear polarization of the eclipsed sky measured in the red (650 nm), green (550 nm), and blue (450 nm) parts of the spectrum. We also report on the temporal and spectral change of the positions of neutral (unpolarized, p = 0) points, and points with local minima or maxima of p of the eclipsed sky. Our results are compared with the observations performed by the same polarimetric technique during the total solar eclipse on 11 August 1999 in Hungary. Practically the same characteristics of celestial polarization were encountered during both eclipses. This shows that the observed polarization phenomena of the eclipsed sky may be general.

Imaging polarimetry of glass buildings: why do vertical glass surfaces attract polarotactic insects?

Recently it was observed that the Hydropsyche pellucidula caddis flies swarm near sunset at the vertical glass surfaces of buildings standing on the bank of the Danube river in Budapest, Hungary. These aquatic insects emerge from the Danube and are lured to dark vertical panes of glass, where they swarm, land, copulate, and remain for hours. It was also shown that ovipositing H. pellucidula caddis flies are attracted to highly and horizontally polarized light stimulating their ventral eye region and thus have positive polarotaxis. The attraction of these aquatic insects to vertical reflectors is surprising, because after their aerial swarming, they must return to the horizontal surface of water bodies from which they emerge and at which they lay their eggs. Our aim is to answer the questions: Why are flying polarotactic caddis flies attracted to vertical glass surfaces? And why do these aquatic insects remain on vertical panes of glass after landing? We propose that both questions can be partly explained by the reflection-polarization characteristics of vertical glass surfaces and the positive polarotaxis of caddis flies. We measured the reflection-polarization patterns of shady and sunlit, black and white vertical glass surfaces from different directions of view under clear and overcast skies by imaging polarimetry in the red, green, and blue parts of the spectrum. Using these polarization patterns we determined which areas of the investigated glass surfaces are sensed as water by a hypothetical polarotactic insect facing and flying toward or landed on a vertical pane of glass. Our results strongly support the mentioned proposition. The main optical characteristics of "green," that is, environmentally friendly, buildings, considering the protection of polarotactic aquatic insects, are also discussed. Such "green" buildings possess features that attract only a small number of polarotactic aquatic insects when standing in the vicinity of fresh waters. Since vertical glass panes of buildings are abundant in the man-made optical environment, and polarotactic aquatic insects are spread worldwide, our results are of general interest in the visual and behavioral ecology of aquatic insects.

Imaging polarimetry of forest canopies: how the azimuth direction of the sun, occluded by vegetation, can be assessed from the polarization pattern of the sunlit foliage.

Radiance, color, and polarization of the light in forests combine to create complex optical patterns. Earlier sporadic polarimetric studies in forests were limited by the narrow fields of view of the polarimeters used in such studies. Since polarization patterns in the entire upper hemisphere of the visual environment of forests could be important for forest-inhabiting animals that make use of linearly polarized light for orientation, we measured 180 degrees field-of-view polarization distributions in Finnish forests. From a hot air balloon we also measured the polarization patterns of Hungarian grasslands lit by the rising sun. We found that the pattern of the angle of polarization alpha of sunlit grasslands and sunlit tree canopies was qualitatively the same as that of the sky. We show here that contrary to an earlier assumption, the alpha-pattern characteristic of the sky always remains visible underneath overhead vegetation, independently of the solar elevation and the sky conditions (clear or partly cloudy with visible sun's disc), provided the foliage is sunlit and not only when large patches of the clear sky are visible through the vegetation. Since the mirror symmetry axis of the alpha-pattern of the sunlit foliage is the solar-antisolar meridian, the azimuth direction of the sun, occluded by vegetation, can be assessed in forests from this polarization pattern. Possible consequences of this robust polarization feature of the optical environment in forests are briefly discussed with regard to polarization-based animal navigation.

Polarization patterns of thick clouds: overcast skies have distribution of the angle of polarization similar to that of clear skies.

The distribution of polarization in the overcast sky has been practically unknown. Earlier the polarization of light from heavily overcast skies (when the Sun's disc was invisible) has been measured only sporadically in some celestial points by point-source polarimetry. What kind of patterns of the degree p and angle alpha of linear polarization of light could develop after transmission through a thick layer of ice or water clouds? To answer this question, we measured the p and alpha patterns of numerous totally overcast skies on the Arctic Ocean and in Hungary by full-sky imaging polarimetry. We present here our finding that depending on the optical thickness of the cloud layer, the pattern of alpha of light transmitted through the ice or water clouds of totally overcast skies is qualitatively the same as the alpha pattern of the clear sky. Under overcast conditions the value of alpha is determined predominantly by scattering on cloud particles themselves. Nevertheless, the degrees of linear polarization of light from overcast skies were rather low (p

Anomalous celestial polarization caused by forest fire smoke: why do some insects become visually disoriented under smoky skies?

The effects of forest fire smoke on sky polarization and animal orientation are practically unknown. Using full-sky imaging polarimetry, we therefore measured the celestial polarization pattern under a smoky sky in Fairbanks, Alaska, during the forest fire season in August 2005. It is quantitatively documented here that the celestial polarization, a sky attribute that is necessary for orientation of many polarization-sensitive animal species, above Fairbanks on 17 August 2005 was in several aspects anomalous due to the forest fire smoke: (i) The pattern of the degree of linear polarization p of the reddish smoky sky differed considerably from that of the corresponding clear blue sky. (ii) Due to the smoke, p of skylight was drastically reduced (p(max)

Polarization of 'water-skies' above arctic open waters: how polynyas in the ice-cover can be visually detected from a distance.

The foggy sky above a white ice-cover and a dark water surface (permanent polynya or temporary lead) is white and dark gray, phenomena called the 'ice-sky' and the 'water-sky,' respectively. Captains of icebreaker ships used to search for not-directly-visible open waters remotely on the basis of the water sky. Animals depending on open waters in the Arctic region may also detect not-directly-visible waters from a distance by means of the water sky. Since the polarization of ice-skies and water-skies has not, to our knowledge, been studied before, we measured the polarization patterns of water-skies above polynyas in the arctic ice-cover during the Beringia 2005 Swedish polar research expedition to the North Pole region. We show that there are statistically significant differences in the angle of polarization between the water-sky and the ice-sky. This polarization phenomenon could help biological and man-made sensors to detect open waters not directly visible from a distance. However, the threshold of polarization-based detection would be rather low, because the degree of linear polarization of light radiated by water-skies and ice-skies is not higher than 10%.

Imaging polarimetry of the circularly polarizing cuticle of scarab beetles (Coleoptera: Rutelidae, Cetoniidae).

The light reflected from the metallic-shiny regions of the cuticle of certain beetles belonging to the Scarabaeoidea is known since 1911 to be left-handed circularly polarized. Only photographs of a few selected species of scarabs, taken through left- and right-circular polarizers, have earlier been published. Through a right-circular polarizer these beetles appear more or less dark. This demonstration is, however, inadequate to quantitatively investigate the spatial distribution and the wavelength dependency of the circular polarization of light reflected from the scarab cuticle. In order to overcome this problem, we have developed a portable, rotating analyzer, linear/circular, digital, and imaging polarimeter. We describe here our polarimetric technique and present for the first time the linear and circular polarization patterns of the scarab species Chrysophora chrysochlora, Plusiotis resplendens (Rutelidae), and Cetonischema jousselini (Cetoniidae) in the red (650 nm), green (550 nm), and blue (450 nm) parts of the spectrum. We found the wavelength- and species-dependent circular polarization patterns in scarabs to be of a rather complex nature. These patterns are worthy of further studies.

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.

Polarizational colours could help polarization-dependent colour vision systems to discriminate between shiny and matt surfaces, but cannot unambiguously code surface orientation.

It was hypothesized that egg-laying Papilio butterflies could use polarizational colours as a cue to detect leaf orientation and to discriminate between shiny and matt leaves. These hypotheses would be supported if the following general questions were answered positively: (1) Can surface orientation be unambiguously coded by the polarizational colours perceived by polarization-sensitive colour vision systems? (2) Are the changes in the polarizational colours due to retinal rotation significantly different between shiny and matt surfaces? Using video polarimetry, we measured the reflection-polarizational characteristics of a shiny green hemisphere in the red, green and blue spectral ranges for different solar elevations and directions of view with respect to the solar azimuth as well as for sunlit and shady circumstances under clear skies. The continuously curving hemisphere models numerous differently oriented surfaces. Using the polarization- and colour-sensitive retina model developed earlier, we computed the polarizational colours of the hemisphere, and investigated the correlation between colours and local surface orientation. We also calculated the maximal changes of the polarizational colours of shiny and matt hemispheres induced by rotation of the retina. We found that a surface with any orientation can possess almost any polarizational colour under any illumination condition. Consequently, polarizational colours cannot unambiguously code surface orientation. Polarization sensitivity is even disadvantageous for the detection of surface orientation by means of colours. On the other hand, the colour changes due to retinal rotation can be significantly larger for shiny surfaces than for matt ones. Thus, polarizational colours could help discrimination between shiny and matt surfaces. The physical and perceptional reasons for these findings are explained in detail. Our results and conclusions are of general importance for polarization-dependent colour vision systems.