Dual structural color mechanisms in a scarab beetle

The cuticle of a Mycterophallus cetoniine scarab species displays both red iridescence due to a multilayer reflector mechanism and rainbow iridescence due to a superimposed diffraction grating mechanism. This is the first reported example of an animal possessing two independent classes of structural colors arising from interference at the wavelengths of visible light. In this work, the Mycterophallus cuticle is characterized by light microscopy, spectrophotometry, scanning electron microscopy, and transmission electron microscopy. We compare the cuticle of the Mycterophallus species to two closely related Lomaptera scarab species, one with only a multilayer reflector and the second with only a diffraction grating. We calculate the correspondence between the nanostructural parameters and the optical properties of the Mycterophallus cuticle to determine the relative optical contributions of the two color mechanisms and the interactions between them. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Ultrastructural variation tune wing coloration of a moth Asota caricae Fabricius, 1775

Butterflies and moths develop highly ordered coloration in their wing for signal transmission. We have investigated the ultrastructural arrangement of wing coloration of a moth Asota caricae, applying light, optical polarized, and scanning electron microscopy, and spectrophotometry. The forewing of the moth is brown in color with a white spot at the center. The hindwing is golden yellow in color with many black patches in it. The ventral part of the forewing and dorsal hindwing share the similar color pattern. The ventral part of the hindwing has dull coloration in comparison to the dorsal one although the pattern remains same. The spectrometry analysis reveals various patterns of absorbance and reflectance spectra for various colors. The peak observed for various colors remain same although the intensity of peak changes. Bright colors possess highly ordered structures whereas irregular structures are found in dull colored scales. The color variation observed due to dorsal and ventral part of the wing is due to the minute difference observed in terms of ultrastructural arrangement revealed by scanning electron microscope. The color pattern of A. caricae is due to variation of microstructures present within the scale.     

Species‐dependent microarchitectural traits of iridescent scales in the triad taxa of Ornithoptera birdwing butterflies

Ornithoptera birdwing butterflies have blue, green, or orange iridescent scales in different species or subspecies. To understand the species‐ or subspecies‐dependent scale color differences, we performed comparative morphometric analyses of iridescent scales from three closely related taxa: O. priamus priamus (green), O. priamus urvillianus (blue), and O. croesus (orange). The three types of Ornithoptera wings exhibited reversible color changes to longer wavelengths with different kinetics upon immersion in methanol, suggesting that their color differences are at least partly based on differences in the size of air cavities made by nanostructures. Cover scales of all three color types were visually semi‐transparent glass scales that exhibited color when placed on a dark background. The dorsoventral differences in coloration were observed in single scales, suggesting the optical importance of scale surfaces. Scanning electron microscopy of cover scales in cross section revealed that all color types exhibited finely sculpted tapered ridges and thick, irregular basal multilayers containing tandemly clustered granular objects and air cavities. Scale thickness, ridge height, and multilayer thickness were significantly different among the three color types, and granular object size was significantly different between orange scales and blue and green scales. We conclude that each of the three taxa of Ornithoptera butterflies possesses unique quantitative size values on tapered ridges and irregular multilayers with granular objects and air cavities to express unique structural color. These species‐ or subspecies‐dependent structural colors might have evolved via quantitative shifts in these microarchitectural traits rather than via changes in the basic developmental or architectural plan for color expression.     

多尺度植物色彩表征及其与人体响应的关系

色彩是人们对植物外观的第一印象,与植物视觉特征和人体响应关系密切,因此,对森林美景度提升和公众适宜性契合具有重要作用。结合色彩学形成的历史关键节点和1个世纪以来植物色彩的研究动态,将植物色彩研究的发展历程划分为萌芽、形成、发展和繁荣等4个阶段,并简介了各阶段的主要特点。同时,从植物器官、植物个体、植物群体(涵盖种群和群落)、整体景观等4个视觉尺度以及审美、生理与心理等3个影响层面,阐述与评价了植物色彩传达机制与人体响应关系的相关研究。在此基础上,围绕植物色彩特征提取中存在的色彩要素定量、特征因子选取、色彩量化方法等方面的不足之处,对未来研究方向进行展望,提出植物色彩尺度与公众响应关系剖析、较大尺度色彩评价因子的深入挖掘、色彩量化方法的多学科融入、公众对植物色彩响应关系综合比对、森林景观美景度提升实践等5个方向的发展趋势,以期为全面地了解植物色彩的传达机制与影响、科学地开展不同精度和尺度下的植物色彩研究提供参考。     

The original colours of fossil beetles

Structural colours, the most intense, reflective and pure colours in nature, are generated when light is scattered by complex nanostructures. Metallic structural colours are widespread among modern insects and can be preserved in their fossil counterparts, but it is unclear whether the colours have been altered during fossilization, and whether the absence of colours is always real. To resolve these issues, we investigated fossil beetles from five Cenozoic biotas. Metallic colours in these specimens are generated by an epicuticular multi-layer reflector; the fidelity of its preservation correlates with that of other key cuticular ultrastructures. Where these other ultrastructures are well preserved in non-metallic fossil specimens, we can infer that the original cuticle lacked a multi-layer reflector; its absence in the fossil is not a preservational artefact. Reconstructions of the original colours of the fossils based on the structure of the multi-layer reflector show that the preserved colours are offset systematically to longer wavelengths; this probably reflects alteration of the refractive index of the epicuticle during fossilization. These findings will allow the former presence, and original hue, of metallic structural colours to be identified in diverse fossil insects, thus providing critical evidence of the evolution of structural colour in this group.     

Wing colors based on arrangement of the multilayer structure of wing scales in lycaenid butterflies (Insecta: Lepidoptera)

Male wing colors and wing scale morphology were examined for three species of lycaenid butterflies: Chrysozephyrus ataxus, Favonius cognatus and F. jezoensis. Measurement of spectral reflectance on the wing surface with a spectrophotometer revealed species‐specific reflection spectra, with one or two peaks in the ultraviolet and/or green ranges. Observations of wing scales using an optical microscope revealed that light was reflected from the inter‐ridge regions, where transmission electron microscopy revealed a multilayer structure. Based on the multilayer dimensions obtained, three models were devised and compared to explain the measured reflectance spectrum. The results showed that the best fit is a model in which thicknesses of thin films of the multilayer system are not constant and air spaces between cuticle layers are more or less packed with cuticle spacers. This suggests that the specific wing colors of the species examined are produced by the species‐specific arrangement of the multilayer structure of wing scales.     

Brightness contrast inhibits color induction: evidence for a new kind of color theory

A gray region can be made to look colored by a colored surround. This phenomenon, chromatic induction, depends on color differences around the boundary of the region. We performed experiments on chromatic induction with small, initially achromatic, targets on nine different colored surrounds ranging in color from blue to red. Using scaling of saturation as our measure of perceived color strength, we found that chromatic induction is at its maximum when the brightness contrast at the boundary between target and surroundings is minimal. This implies that the neural mechanism in the cerebral cortex that mediates the appearance of brightness at a boundary inhibits the activity of chromatic mechanisms at that same boundary. Observers matched the apparent brightness and luminance of each of the colored surrounds. For surround colors where brightness and luminance matches differ, brightness contrast, not luminance contrast, controls chromatic induction. These new findings, taken together with other evidence, require a new theory of color appearance that includes mutually inhibitory interactions between color and brightness mechanisms that are sensing color and brightness contrast at visual boundaries.     

Correlations between scale structure and pigmentation in butterfly wings

SUMMARY We examined the correlation between color and structure of wing scales in the nymphalid butterflies Bicyclus anynana and Heliconius melpomene . All scales in B. anynana are rather similar in comparison to the clear structural differences of differently pigmented scales in H. melpomene . Where scale structural differences in H. melpomene are qualitative, they seem to be quantitative in B. anynana . There is a "gradient" in the density of some structural elements, the cross ribs, in the scales of B. anynana : black, gold, and brown scales show progressively lower cross rib density within an individual. There is, however, high individual variation in the absolute cross rib densities (i.e., scales with a particular color and cross rib density in one individual may have a different color but similar density in another individual). By ectopically inducing color pattern during early pupal development, we examined whether a scale's color and its microstructure could be uncoupled. The effect of these manipulations appears to be different in B. anynana and H. melpomene . In Bicyclus , "black" scales induced by wing damage at an ectopic location normally containing brown scales acquire both an intermediate structure and color between that of brown and normal black scales. In Heliconius , however, intermediate colors or scale structure were never observed, and scales with an altered color (due to damage) always have the same structure as normal scales with that color. The results are discussed on the basis of gene expression patterns, variability in rates of scale development and pigment, and scale sclerotization pathways.     

The cause of 50 million-year-old colour

Multilayer reflectors cause structural, 'metallic' colours in a diversity of animals today, yet are unknown in extinct species. We identify a multilayer reflector, causing structural colour, in a 50-million-year-old beetle from Messel, Germany. It is proposed that the original material of this reflector has been preserved, although this is not a precondition for determining original colours from ancient multilayer reflectors. Therefore, the potential exists to reveal the original colours of other (particularly arthropod) extinct species.     

Hormonal control of GTP cyclohydrolase I gene expression and enzyme activity during color pattern development in wings of Precis coenia

Color patterns of butterfly wings are composed of single color points represented by each scale. In the case of Precis coenia, at the end of pupal development, different types of pigments are synthesized sequentially in the differently colored scales beginning with white (pterins) followed by red (ommatins) and then black (melanin). In order to explain how formation of these different colors is regulated, we examined the expression of an mRNA-encoding guanosine triphosphate-cyclohydrolase I (GTP-CH I; EC 3.5.4.16), the first key enzyme in the biosynthesis of pteridines, during pigment formation in the wings of P. coenia. The strongest positive signal was recognized around pigment formation one day before butterfly emergence. This GTP-CH I gene expression is paralleled by GTP-CH I enzyme activity measured in wing extracts. We also investigated the effect of 20-hydroxyecdysone on the expression of GTP-CH I mRNA and the enzyme activity during color formation. The results strongly suggest that the onset and duration of the expression of a GTP-CH I mRNA is triggered by a declining ecdysteroid hormone titer during late pupal development.     

Exceptional three-dimensional preservation and coloration of an originally iridescent fossil feather from the Middle Eocene Messel Oil Shale

A feather from the Eocene Messel Formation, Germany, has been demonstrated to have been originally structurally colored by densely packed sheets of melanosomes similar to modern iridescent feathers exhibiting thin-film diffraction. The fossil itself currently exhibits a silvery sheen, but the mechanism for generating this optical effect was not fully understood. Here we use scanning electron microscopy, electron probe microanalysis, and dual-beam focused ion beam scanning electron microscopy to investigate the source of the silvery sheen that occurs in the apical feather barbules. Focused ion beam scanning electron microscopy provides a powerful tool for studying three-dimensionality of nanostructures in fossils. Use of the method reveals that the flattened apical barbules are preserved almost perfectly, including smooth structural melanosome sheets on the obverse surface of the fossil feather that are identical to those that cause iridescence in modern bird feathers. Most of each apical barbule is preserved beneath a thin layer of sediment. The silvery sheen is generated by incoherent light diffraction between this sediment layer and melanosomes and, although related to the original iridescence of the feather, is not a feature of the feather itself. The reddish and greenish hues frequently exhibited by fossil feathers from the Messel Formation appear to be due to precipitates on the surface of individual melanosomes.     

Neural locus of color afterimages

After fixating on a colored pattern, observers see a similar pattern in complementary colors when the stimulus is removed [1-6]. Afterimages were important in disproving the theory that visual rays emanate from the eye, in demonstrating interocular interactions, and in revealing the independence of binocular vision from eye movements. Afterimages also prove invaluable in exploring selective attention, filling in, and consciousness. Proposed physiological mechanisms for color afterimages range from bleaching of cone photopigments to cortical adaptation [4-9], but direct neural measurements have not been reported. We introduce a time-varying method for evoking afterimages, which provides precise measurements of adaptation and a direct link between visual percepts and neural responses [10]. We then use in vivo electrophysiological recordings to show that all three classes of primate retinal ganglion cells exhibit subtractive adaptation to prolonged stimuli, with much slower time constants than those expected of photoreceptors. At the cessation of the stimulus, ganglion cells generate rebound responses that can provide afterimage signals for later neurons. Our results indicate that afterimage signals are generated in the retina but may be modified like other retinal signals by cortical processes, so that evidence presented for cortical generation of color afterimages is explainable by spatiotemporal factors that modify all signals.     

Signal evolution and morphological complexity in hummingbirds (Aves: Trochilidae)

Understanding how animal signals are produced is critical for understanding their evolution because complexity and modularity in the underlying morphology can affect evolutionary patterns. Hummingbird feathers show some of the brightest and most iridescent colors in nature. These are produced by optically complex stacks of hollow, platelet-shaped organelles called melanosomes. Neither how these morphologies produce colors nor their evolution has been systematically studied. We first used nanoscale morphological measurements and optical modeling to identify the physical basis of color production in 34 hummingbird species. We found that, in general, the melanosome stacks function as multilayer reflectors, with platelet thickness and air space size explaining variation in hue (color) and saturation (color purity). Additionally, light rays reflected from the outer keratin surface interact with those reflected by small, superficial melanosomes to cause secondary reflectance peaks, primarily in short (blue) wavelengths. We then compared variation of both the morphological components and the colors they produce. The outer keratin cortex evolves independently and is more variable than other morphological traits, possibly due to functional constraints on melanosome packing. Intriguingly, shorter wavelength colors evolve faster than longer wavelength colors, perhaps due to developmental processes that enables greater lability of the shapes of small melanosomes. Together, these data indicate that increased structural complexity of feather tissues is associated with greater variation in morphology and iridescent coloration.     

Precise colocalization of interacting structural and pigmentary elements generates extensive color pattern variation in <Emphasis Type="Italic">Phelsuma</Emphasis>lizards

Background

Color traits in animals play crucial roles in thermoregulation, photoprotection, camouflage, and visual communication, and are amenable to objective quantification and modeling. However, the extensive variation in non-melanic pigments and structural colors in squamate reptiles has been largely disregarded. Here, we used an integrated approach to investigate the morphological basis and physical mechanisms generating variation in color traits in tropical day geckos of the genus Phelsuma.

Results

Combining histology, optics, mass spectrometry, and UV and Raman spectroscopy, we found that the extensive variation in color patterns within and among Phelsuma species is generated by complex interactions between, on the one hand, chromatophores containing yellow/red pteridine pigments and, on the other hand, iridophores producing structural color by constructive interference of light with guanine nanocrystals. More specifically, we show that 1) the hue of the vivid dorsolateral skin is modulated both by variation in geometry of structural, highly ordered narrowband reflectors, and by the presence of yellow pigments, and 2) that the reflectivity of the white belly and of dorsolateral pigmentary red marks, is increased by underlying structural disorganized broadband reflectors. Most importantly, these interactions require precise colocalization of yellow and red chromatophores with different types of iridophores, characterized by ordered and disordered nanocrystals, respectively. We validated these results through numerical simulations combining pigmentary components with a multilayer interferential optical model. Finally, we show that melanophores form dark lateral patterns but do not significantly contribute to variation in blue/green or red coloration, and that changes in the pH or redox state of pigments provide yet another source of color variation in squamates.

Conclusions

Precisely colocalized interacting pigmentary and structural elements generate extensive variation in lizard color patterns. Our results indicate the need to identify the developmental mechanisms responsible for the control of the size, shape, and orientation of nanocrystals, and the superposition of specific chromatophore types. This study opens up new perspectives on Phelsuma lizards as models in evolutionary developmental biology.

     

Behavioral research in pigeons with ARENA: An Automated Remote Environmental Navigation Apparatus

Three experiments established the effectiveness of an Automated Remote Environmental Navigation Apparatus (ARENA) developed in our lab to study behavioral processes in pigeons. The technology utilizes one or more wireless modules, each capable of presenting colored lights as visual stimuli to signal reward and of detecting subject peck responses. In Experiment 1, subjects were instrumentally shaped to peck at a single ARENA module following an unsuccessful autoshaping procedure. In Experiment 2, pigeons were trained with a simultaneous discrimination procedure during which two modules were illuminated different colors; pecks to one color (S+) were reinforced while pecks to the other color (S−) were not. Pigeons learned to preferentially peck the module displaying the S+. In Experiment 3, two modules were lit the same color concurrently from a set of six colors in a conditional discrimination task. For three of the colors pecks to the module in one location (e.g., upper quadrant) were reinforced while for the remaining colors pecks at the other module (e.g., lower quadrant) were reinforced. After learning this discrimination, the color-reinforced location assignments were reversed. Pigeons successfully acquired the reversal. ARENA is an automated system for open-field studies and a more ecologically valid alternative to the touchscreen.     

Perceived speed of colored stimuli

The influence of contrast and color on perceived motion was measured using a speed-matching task. Observers adjusted the speed of an L cone contrast pattern to match that of a variety of colored test patterns. The dependence of speed on test contrast was the same for all test colors measured, differing only by a sensitivity factor. This result suggests that the reduced apparent speed of low contrast targets and certain colored targets is caused by a common cortical mechanism. The cone contrast levels that equate perceived speed differ substantially from those that equate visibility. This result suggests that the neural mechanisms governing speed perception and visibility differ. Perceived speed differences caused by variations in color can be explained by color responses that are characteristic of motion-selective cortex.     

Aggressive Behavior and Courtship Rejection in Brightly and Plainly Colored Female Keeled Earless Lizards (Holbrookia propinqua)

Relationships between bright secondary sexual coloration and behavior were studied in female Holbrookia propinqua, which develop striking orange and yellow colors during the breeding season. In tethered introduction studies, brightly colored females performed aggressive courtship rejection behaviors toward conspecific males; plainly colored females were not aggressive toward males, but attempted to avoid them. Responses of females of the two color patterns toward conspecific females of both color phases were not detectably different. Experimental introductions of lizards with coloration modified by paint showed that females of both color patterns recognize any other lizard bearing the bright female colors as female, regardless of actual sex. Both the orange and yellow components were shown to contribute to sex recognition. The yellow component alone allowed accurate sex identification, but only half the females responded to males painted with only the orange female component as if they were females. Because females did not behave differently toward other females on the basis of coloration, the hypothesis that bright coloration evolved as an adaptive signal between females is rejected. The dark ventrolateral stripes of male and plainly colored females did not appear to affect intraspecific social responses by females.     

彩叶植物叶片呈色分子机制研究进展

彩叶植物叶片呈现不同的颜色主要是受遗传因素和外部环境的共同作用,揭示彩叶植物叶片呈色机制对选育彩叶植物新品种和彩叶植物的应用推广具有重要理论和实践意义。目前对彩叶植物呈色机制的研究主要集中于叶片中色素变化、光合特性、叶片结构和环境条件等方面。该文主要対近年来有关彩叶植物叶片中叶绿素代谢途径、类胡萝卜素代谢途径、次生代谢途径、光合作用和叶绿体发育相关结构基因和转录因子调控机制的研究进展进行综述,并对以后的研究方向进行了展望,为培育彩叶植物新品种提供了理论基础,也为人工调控叶色以及叶色的定向遗传改良提供了参考。     

Human visual system integrates color signals along a motion trajectory

Whether fundamental visual attributes, such as color, motion, and shape, are analyzed separately in specialized pathways has been one of the central questions of visual neuroscience. Although recent studies have revealed various forms of cross-attribute interactions, including significant contributions of color signals to motion processing, it is still widely believed that color perception is relatively independent of motion processing. Here, we report a new color illusion, motion-induced color mixing, in which moving bars, the color of each of which alternates between two colors (e.g., red and green), are perceived as the mixed color (e.g., yellow) even though the two colors are never superimposed on the retina. The magnitude of color mixture is significantly stronger than that expected from direction-insensitive spatial integration of color signals. This illusion cannot be ascribed to optical image blurs, including those induced by chromatic aberration, or to involuntary eye movements of the observer. Our findings indicate that color signals are integrated not only at the same retinal location, but also along a motion trajectory. It is possible that this neural mechanism helps us to see veridical colors for moving objects by reducing motion blur, as in the case of luminance-based pattern perception.     

Fechner-Benham subjective colors do not induce McCollough after-effects

Fechner-Benham subjective color is widely believed to be governed by local interactions in early (probably retinal) mechanisms. Here we report three lines of phenomenological evidence that suggest otherwise: subjective colors seen in spatially extended stimuli (a) are dependent on global aspects of the stimuli; (b) can become multistable in position; and (c) even after being stabilized do not support the creation of McCollough's colored after-effects--a cortically based phenomenon generally thought to be more central than Fechner-Benham color. These phenomena suggest a central locus that controls perception of subjective color, characterized by pattern dependent interactions among cortical mechanisms that draw their inputs from peripheral units.