Browsing by Author "Chiou, Tsyr-Huei"
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Item Biological polarized light reflectors in stomatopod crustaceans(SPIE, 2005-09-10) Chiou, Tsyr-Huei; Cronin, Thomas W.; Caldwell, Roy L.; Marshall, JustinBody parts that can reflect highly polarized light have been found in several species of stomatopod crustaceans (mantis shrimps). These polarized light reflectors can be grossly divided into two major types. The first type, usually red or pink in color to the human visual system, is located within an animal’s cuticle. Reflectors of the second type, showing iridescent blue, are located beneath the exoskeleton and thus are unaffected by the molt cycle. We used reflection spectropolarimetry and transmission electron microscopy (TEM) to study the reflective properties and the structures that reflect highly polarized light in stomatopods. For the first type of reflector, the degree of polarization usually changes dramatically, from less than 20% to over 70%, with a change in viewing angle. TEM examination indicates that the polarization reflection is generated by multilayer thin-film interference. The second type of reflector, the blue colored ones, reflects highly polarized light to all viewing angles. However, these reflectors show a slight chromatic change with different viewing angles. TEM sections have revealed that streams of oval-shaped vesicles might be responsible for the production of the polarized light reflection. In all the reflectors we have examined so far, the reflected light is always maximally polarized at around 500 nm, which is close to the wavelength best transmitted by sea water. This suggests that the polarized light reflectors found in stomatopods are well adapted to the underwater environment. We also found that most reflectors produce polarized light with a horizontal e-vector. How these polarized light reflectors are used in stomatopod signaling remains unknown.Item Circular Polarization Vision in a Stomatopod Crustacean(Elsevier B.V, 2008-03-25) Chiou, Tsyr-Huei; Kleinlogel, Sonja; Cronin, Tom; Caldwell, Roy; Loeffler, Birte; Siddiqi, Afsheen; Goldizen, Alan; Marshall, JustinWe 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 . This latter capability can be used for object identification, contrast enhancement, navigation, and communication through polarizing reflections . Circularly polarized reflections from a few animal species have also been known for some time . Although optically interesting , 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. 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.Item Fine structure and optical properties of biological polarizers in crustaceans and cephalopods(SPIE, 2008-03-24) Chiou, Tsyr-Huei; Caldwell, Roy L.; Hanlon, Roger T.; Cronin, Thomas W.The lighting of the underwater environment is constantly changing due to attenuation by water, scattering by suspended particles, as well as the refraction and reflection caused by the surface waves. These factors pose a great challenge for marine animals which communicate through visual signals, especially those based on color. To escape this problem, certain cephalopod mollusks and stomatopod crustaceans utilize the polarization properties of light. While the mechanisms behind the polarization vision of these two animal groups are similar, several distinctive types of polarizers (i.e. the structure producing the signal) have been found in these animals. To gain a better knowledge of how these polarizers function, we studied the relationships between fine structures and optical properties of four types of polarizers found in cephalopods and stomatopods. Although all the polarizers share a somewhat similar spectral range, around 450- 550 nm, the reflectance properties of the signals and the mechanisms used to produce them have dramatic differences. In cephalopods, stack-plates polarizers produce the polarization patterns found on the arms and around their eyes. In stomatopods, we have found one type of beam-splitting polarizer based on photonic structures and two absorptive polarizer types based on dichroic molecules. These stomatopod polarizers may be found on various appendages, and on the cuticle covering dorsal or lateral sides of the animal. Since the efficiencies of all these polarizer types are somewhat sensitive to the change of illumination and viewing angle, how these animals compensate with different behaviors or fine structural features of the polarizer also varies.Item Light habitats and the role of polarized iridescence in the sensory ecology of neotropical nymphalid butterflies (Lepidoptera: Nymphalidae)(The Company of Biologists Ltd, 2007-02-12) Douglas, Jonathan M.; Cronin, Thomas W.; Chiou, Tsyr-Huei; Dominy, Nathaniel J.The exploitation of polarized light may increase perceived visual contrast independent of spectrum and intensity and thus have adaptive value in forest habitats, where illumination varies greatly in brightness and spectral properties. Here we investigate the extent to which Costa Rican butterflies of the family Nymphalidae exhibit polarized wing reflectance and evaluate the types of habitats in which the trait is commonly found. We also examine the degree of polarized reflectance of wing patterns in representative species belonging to the nymphalid subfamilies Charaxinae, Heliconiinae, Morphinae and Nymphalinae. Polarized reflectance was evaluated using museum specimens illuminated with a light source that simulated the spectrum of ambient sunlight and viewed through a polarized filter. Of the 144 species examined, 75 species exhibited polarized reflectance patterns. These species were significantly more likely to occupy forest habitats than open habitats. A concentrated changes test performed on a phylogeny of the Nymphalidae, with the Papilionidae as an outgroup, provides further support for the correlated evolution of polarized iridescence and life in a forest light environment. These results are consistent with the hypothesis that the production and detection of polarized light may have adaptive communicative value in those species inhabiting forest habitats with complex light conditions. The potential utility of polarized iridescence and iridescent wing coloration within differing ambient spectral environments is discussed to provide a basis for future investigation of the polarized light ecology of butterflies.Item A novel function for a carotenoid: astaxanthin used as a polarizer for visual signalling in a mantis shrimp(The Company of Biologists Ltd, 2011-11-17) Chiou, Tsyr-Huei; Place, Allen R.; Caldwell, Roy L.; Marshall, N. Justin; Cronin, Thomas W.Biological signals based on color patterns are well known, but some animals communicate by producing patterns of polarized light. Known biological polarizers are all based on physical interactions with light such as birefringence, differential reflection or scattering. We describe a novel biological polarizer in a marine crustacean based on linear dichroism of a carotenoid molecule. The red-colored, dichroic ketocarotenoid pigment astaxanthin is deposited in the antennal scale of a stomatopod crustacean, Odontodactylus scyllarus. Positive correlation between partial polarization and the presence of astaxanthin indicates that the antennal scale polarizes light with astaxanthin. Both the optical properties and the fine structure of the polarizationally active cuticle suggest that the dipole axes of the astaxanthin molecules are oriented nearly normal to the surface of the antennal scale. While dichroic retinoids are used as visual pigment chromophores to absorb and detect polarized light, this is the first demonstration of the use of a carotenoid to produce a polarizing signal. By using the intrinsic dichroism of the carotenoid molecule and orienting the molecule in tissue, nature has engineered a previously undescribed form of biological polarizer.Item Polarization signals in mantis shrimps(SPIE, 2009-08-11) Cronin, Thomas W.; Chiou, Tsyr-Huei; Caldwell, Roy L.; Roberts, Nicholas; Marshall, JustinWhile color signals are well known as a form of animal communication, a number of animals communicate using signals based on patterns of polarized light reflected from specialized body parts or structures. Mantis shrimps, a group of marine crustaceans, have evolved a great diversity of such signals, several of which are based on photonic structures. These include resonant scattering devices, structures based on layered dichroic molecules, and structures that use birefringent layers to produce circular polarization. Such biological polarizers operate in different spectral regions ranging from the near-UV to medium wavelengths of visible light. In addition to the structures that are specialized for signal production, the eyes of many species of mantis shrimp are adapted to detect linearly polarized light in the ultraviolet and in the green, using specialized sets of photoreceptors with oriented, dichroic visual pigments. Finally, a few mantis shrimp species produce biophotonic retarders within their photoreceptors that permit the detection of circularly polarized light and are thus the only animals known to sense this form of polarization. Mantis shrimps use polarized light in species-specific signals related to mating and territorial defense, and their means of manipulating light’s polarization can inspire designs for artificial polarizers and achromatic retarders.Item Polarization signals in the marine environment(SPIE, 2003-12-12) Cronin, Thomas W.; Shashar, Nadav; Caldwell, Roy L.; Marshall, Justin; Cheroske, Alexander G.; Chiou, Tsyr-HueiAlthough natural light sources produce depolarized light, partially linearly polarized light is naturally abundant in the scenes animals view, being produced by scattering in air or water or by reflection from shiny surfaces. Many species of animals are sensitive to light's polarization, and use this sensitivity to orient themselves using polarization patterns in the atmosphere or underwater. A few animal species have been shown to take this polarization sensitivity to another level of sophistication, seeing the world as a polarization image, analogous to the color images humans and other animals view. This sensory capacity has been incorporated into biological signals by a smaller assortment of species, who use patterns of polarization on their bodies to communicate with conspecific animals. In other words, they use polarization patterns for tasks similar to those for which other animals use biologically produced color patterns. Polarization signals are particularly useful in marine environments, where the spectrum of incident light is variable and unpredictable. Here, cephalopod mollusks (octopuses, squids, and cuttlefish) and stomatopod crustaceans (mantis shrimps) have developed striking patterns of polarization used in communication.Item Polarization Vision and Its Role in Biological Signaling(Oxford University Press, 2003-08-01) Cronin, Thomas W.; Shashar, Nadav; Caldwell, Roy L.; Marshall, Justin; Cheroske, Alexander G.; Chiou, Tsyr-HueiVisual pigments, the molecules in photoreceptors that initiate the process of vision, are inherently dichroic, differentially absorbing light according to its axis of polarization. Many animals have taken advantage of this property to build receptor systems capable of analyzing the polarization of incoming light, as polarized light is abundant in natural scenes (commonly being produced by scattering or reflection). Such polarization sensitivity has long been associated with behavioral tasks like orientation or navigation. However, only recently have we become aware that it can be incorporated into a high-level visual perception akin to color vision, permitting segmentation of a viewed scene into regions that differ in their polarization. By analogy to color vision, we call this capacity polarization vision. It is apparently used for tasks like those that color vision specializes in: contrast enhancement, camouflage breaking, object recognition, and signal detection and discrimination. While color is very useful in terrestrial or shallow-water environments, it is an unreliable cue deeper in water due to the spectral modification of light as it travels through water of various depths or of varying optical quality. Here, polarization vision has special utility and consequently has evolved in numerous marine species, as well as at least one terrestrial animal. In this review, we consider recent findings concerning polarization vision and its significance in biological signaling.Item A shape-anisotropic reflective polarizer in a stomatopod crustacean(Springer Nature Publishing AG, 2016-02-17) Jordan, Thomas M.; Wilby, David; Chiou, Tsyr-Huei; Feller, Kathryn D.; Caldwell, Roy L.; Cronin, Thomas W.; Roberts, Nicholas W.Many biophotonic structures have their spectral properties of reflection ‘tuned’ using the (zeroth-order) Bragg criteria for phase constructive interference. This is associated with a periodicity, or distribution of periodicities, parallel to the direction of illumination. The polarization properties of these reflections are, however, typically constrained by the dimensional symmetry and intrinsic dielectric properties of the biological materials. Here we report a linearly polarizing reflector in a stomatopod crustacean that consists of 6–8 layers of hollow, ovoid vesicles with principal axes of ~550 nm, ~250 nm and ~150 nm. The reflection of unpolarized normally incident light is blue/green in colour with maximum reflectance wavelength of 520 nm and a degree of polarization greater than 0.6 over most of the visible spectrum. We demonstrate that the polarizing reflection can be explained by a resonant coupling with the first-order, in-plane, Bragg harmonics. These harmonics are associated with a distribution of periodicities perpendicular to the direction of illumination, and, due to the shape-anisotropy of the vesicles, are different for each linear polarization mode. This control and tuning of the polarization of the reflection using shape-anisotropic hollow scatterers is unlike any optical structure previously described and could provide a new design pathway for polarization-tunability in man-made photonic devices.Item Spectral and spatial properties of polarized light reflections from the arms of squid (Loligo pealeii) and cuttlefish (Sepia officinalis L.)(The Company of Biologists Ltd, 2007-07-16) Chiou, Tsyr-Huei; Mäthger, Lydia M.; Hanlon, Roger T.; Cronin, Thomas W.On every arm of cuttlefish and squid there is a stripe of high-reflectance iridophores that reflects highly polarized light. Since cephalopods possess polarization vision, it has been hypothesized that these polarized stripes could serve an intraspecific communication function. We determined how polarization changes when these boneless arms move. By measuring the spectral and polarizing properties of the reflected light from samples at various angles of tilt and rotation, we found that the actual posture of the arm has little or no effect on partial polarization or the e-vector angle of the reflected light. However, when the illumination angle changed, the partial polarization of the reflected light also changed. The spectral reflections of the signals were also affected by the angle of illumination but not by the orientation of the sample. Electron microscope samples showed that these stripes are composed of several groups of multilayer platelets within the iridophores. The surface normal to each group is oriented at a different angle, which produces essentially constant reflection of polarized light over a range of viewing angles. These results demonstrate that cuttlefish and squid could send out reliable polarization signals to a receiver regardless of arm orientation.