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Photos of all butterflies examined in this study. They are arranged on a map of the world in order to indicate their natural habitat.
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The bright colors found on the wings of some butterflies have been widely examined during recent decades because they are frequently caused by nano-structures and not by pigments or dyes. Sometimes it is puzzling to discover the physical origin of these structural colors because the color-causing nano-structures are integrated into a complex struct...
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... we analysed 10 different buttery species from 7 sub-families and 3 families of Lepidoptera. Fig. 1 displays photos of the dorsal sides of all examined butteries roughly positioned on a world map in accordance with their natural habitats. They originate from all ve continents and differ in colour appearance and habitat. Table 1 summarizes the examined species together with their respective subfamilies and families. All butteries ...
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... scanning electron microscopy (SEM). Some scales were also examined by transmission electron microscopy (TEM). Fig. 2 displays examples of such images for the Eurasian buttery Aglais urticae commonly known as 'small tortoiseshell'. Its dorsal wings are mostly covered with orange/brownish and black areas but also contain blue and white spots (see Fig. 1). Imaging the wing by scanning electron microscopy reveals that the scales lie on one another like roof tiles all over the wing (Fig. 2a). Zooming into single scales shows that the scales of A. urticae exhibit a hollow structure (Fig. 2b). Ridges, ribs, and trabeculae can be easily distinguished. 14 The ridges and ribs belong to the ...
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... The dazzling iridescent hues, characterized by their remarkable reflectivity, in the Morpho butterfly's wings originate from a complex nanostructure. This nanostructure emerges through the intricate interplay of multilayer interference, the harmonious arrangement of multiscale wing components, the presence of periodic geometric shapes and particle's density changes, and a diverse array of sizes and shapes-ranging from lattice frames to hexagonal tiles and lamella-all facilitated by dynamic movement patterns [46,47]. According to Louvers' features and functions, the correspondent criteria can be abstracted from the aforementioned biomimetic principles as morphology and mechanism. ...
The number of desk workers who frequently conduct their jobs at home has increased dramatically during Covid-19. Work-from-home flexibility makes it attractive for workers and companies, resulting in a “Work-Style Reform” after the Covid-19 pandemic. However, the quick conversion of home spaces into workplaces cannot always sufficiently respond to users’ visual comfort and daylight performance needs which are primary contributors to occupant well-being and productivity. Therefore, this study adopts a mixed-methodology method that integrates parametric thinking, biomimetic, conceptual design, kinetic strategy and the DIVA approach to develop a real-time parametric-generative circular design for multi-objective adaptability that optimizes visual comfort and electric lighting energy efficiency for multiple occupants simultaneously. Parametric simulations of 1458 different options (five different runs per case: a total of 7290) were conducted to assess how the louvers perform regarding daylight, glare, and electric energy usage. Implementing an interactive kinetic louver greatly improved daylight performance in all orientations while simultaneously avoiding visual discomfort for multiple occupants. Furthermore, the use of this façade modification resulted in a substantial decrease in electrical lighting energy consumption, reducing the values from 14.22 to 0.2 kWh/m2/year, 8.1 to 0.18 kWh/m2/year, and 12.88 to 0.18 kWh/m2/year for South, East, and West orientations, respectively. Integrating users' lighting level preferences and the dynamic transitory sensitive area on the façade considerably reduces electric lighting consumption by around 99% compared to the ASHRAE 90.1 standard's lighting profile.
... Their evolutionary history is entirely painted by colours, reflecting multiple adaptations making butterflies excellent study models for elucidating factors and processes associated with the distribution, evolution and functionality of colours in nature (Mallet and Joron 1999;Kemp and Rutowski 2011;Adams et al. 2014;Kunte et al. 2021;Schirmer et al. 2023). Adults, in particular, have a rich "palette" of colours which give rise from simple and homogeneous to more complex and heterogeneous patterns, composing distinct strategies to maximise their fitness in different ecosystems (Hiyama et al. 2012;Köchling et al. 2020;Spaniol et al. 2020). Brightness (i.e. the amount of light reflected by the wing surface) plays a crucial role in thermoregulation, being closely associated with abiotic environmental characteristics, and different behaviours related to body temperature control (Clench 1966;Zeuss et al. 2014;Bladon et al. 2020). ...
Organismal colours have long captivated and inspired naturalists and scientists. Since colours depend on the life history of a species, it is expected that they respond to environmental changes, especially in an increasingly anthropized world. Aiming to evaluate how this trait responds to different anthropogenic disturbances, we assess wing colour aspects of fruit-feeding butterflies sampled in Atlantic Forest remnants. These remnants, with well-defined understory and canopy, are crossed by artificial edges and surround exotic pine silviculture areas of different ages, representing landscapes commonly found in the subtropical Atlantic Forest. We obtained colour measurements of brightness, saturation, contrast, colour diversity and heterogeneity, and the presence of eyespots and iridescence for dorsal and ventral wing surfaces of 47 fruit-feeding butterfly species. We evaluated colour trait distribution and abundance in the distinct native (understory and canopy) and anthropized (edge, old and young Pinus) habitats, and hypothesised that butterfly assemblage colouration will differ in each habitat due to biotic and abiotic differences. In addition, butterfly assemblages in anthropized environments should present less diverse colour traits due to the pressures generated by anthropogenic actions, like microclimate changes and higher exposure to predation. As expected, the natural environments have butterflies with diverse colours and unique contrast colour traits. These patterns are not found in anthropized ones despite artificial edges presenting brighter and even more diverse colours. However, pine silviculture areas present butterflies with less diverse colours, leading the most colourful species to disappear. We demonstrate that different anthropogenic disturbances can impact butterfly colouration. Our results reinforce the close relationship between butterfly colouration and the environment, highlighting that colours can be used as bioindicators for conservation purposes, representing a useful form of functional biodiversity.
... While this complexity makes the self-assembly of these structures fascinating, imaging and capturing this gamut of complexity is extremely challenging. The available imaging methods can either capture the overall field of view at a lower resolution (scanning electron microscopy (SEM)) or can capture high-resolution details of the structure at a specific region (resin section transmission electron microscopy) [1]. Further, three-dimensional (3D) characterization techniques either can only image the topology of a single surface (atomic force microscopy) [1] or are limited to fewer synchrotron facilities (ptychographic x-ray tomography (PXCT)) [2], rendering them impractical to promptly test a large number of scales for gene expression studies. ...
... The available imaging methods can either capture the overall field of view at a lower resolution (scanning electron microscopy (SEM)) or can capture high-resolution details of the structure at a specific region (resin section transmission electron microscopy) [1]. Further, three-dimensional (3D) characterization techniques either can only image the topology of a single surface (atomic force microscopy) [1] or are limited to fewer synchrotron facilities (ptychographic x-ray tomography (PXCT)) [2], rendering them impractical to promptly test a large number of scales for gene expression studies. ...
The complexity of self-assembled photonic structures like butterfly wing scales spans 3–4 orders of magnitude in length scales. These monolithic structures produced by single cells on a butterfly wing typically measure a few hundred micrometres across, with intricate features going well below the nanometre scale. While this complexity makes the self-assembly of these structures fascinating, imaging and capturing this gamut of complexity is extremely challenging. The available imaging methods can either capture the overall field of view at a lower resolution (scanning electron microscopy (SEM)) or can capture high-resolution details of the structure at a specific region (resin section transmission electron microscopy). Further, three-dimensional (3D) characterization techniques either can only image the topology of a single surface (atomic force microscopy) or are limited to fewer synchrotron facilities (ptychographic x-ray tomography (PXCT)) [2], rendering them impractical to promptly test a large number of scales for gene expression studies.
Automation is key to imaging large complex structures like the butterfly scale; the advent of cryo-electron microscopy (cryo-EM) aided the development of automation tools for transmission electron microscopy (TEM). Here we propose an imaging framework which utilizes the new possibilities of TEM provided by such automation tools, energy filters and direct electron detectors to rapidly image an extended field of view of photonic crystals in butterfly wing scales at nanoscale resolution. In addition, we demonstrate that the application of pop-out 3D metrology in this framework provides 3D morphological insights.
... However, this rather obvious assumption may be erroneous. Butterfly scales are a multipurpose convolution of features resulting from constraints and needs that may or may not be related to the many factors involved in the generation of colour [2]. Therefore, identifying the particular contributions of each feature of the scales to determine their functionality is central to understanding the physiological and evolutionary aspects of butterflies. ...
... Therefore, the independent contribution of ridge heights to the overall colour of butterfly wing scales should be carefully integrated into future studies. Our findings support the idea that the diversity and complexity of the colour-producing structures in butterfly wings-and by extension, the cuticles of other arthropods-cannot be solely explained as an efficient strategy for producing different colours [2], as this feat can be achieved by simpler and more efficient design variations. At the same time, other functionalities of the cuticle, such as further optical properties (iridescence, angle independency, light polarisation, etc.) [21], interactions with water [22], aerodynamics [23], structural requirements or even the winding evolutionary path to these structures [24], need to be considered when studying and replicating biological structures that produce colour. ...
The colourful wings of butterflies result from the interaction between light and the intricate chitinous nanostructures on butterflies' scales. This study demonstrates that just by reproducing the chitinous ridges present in butterfly scales (i.e., without any other secondary structure), the entire colour palette is achieved. This result was achieved using a new methodology based on the controlled reproduction of parts of the biological structure of complex chitinous systems using their native chemistry, enabling the isolation of different features' contributions. Here we isolate the contribution of the ridges and their variations as producing and modulating colour hue. The results suggest that complicated butterfly scales may be non-ideal solutions for producing colour when multifunctionality is not considered.
... On a micro or macro scale, plants and animals adapt to dynamic environmental stimuli through several motions and transformations [11,12]. Many species, such as Morpho butterfly wings iridescence, demonstrate a broad range of geometrical and symmetrical spatial patterns in nature [13]. Morpho butterflies are members of the genus Morpho and are divided into numerous dozen species, including rhetenor and didius. ...
... The bright blue hue of the Morpho butterfly's wings has grabbed scientists' curiosity as an example of iridescent coloring with high reflectivity and minimal angle dependency [14][15][16]. Interference between multilayers and multiscale wing parts, geometric shapes, and movement behavior can explain the blue color's high reflectivity [13,[17][18][19]. These species use structural color, interesting optical properties, including iridescence, high reflectance, and substantial polarization effects [14,20]. ...
... The structures have varied sizes and shapes, such as lattice frames and hexagonal tiles [20,21]. Moreover, differing movement behaviors significantly impact the iridescence phenomenon by applying periodic organization and geometrical variations in the butterfly wings (Fig. 1) [13,17]. ...
Daylight contains dynamic features such as color and intensity that may vary over time. The ideas of Orosi windows and biomimetic kinetic lessons of butterfly wings are viewed as essential tactics for regulating dynamic daylight. As a result, this research looks at a methodological technique for providing an appropriate combination of kinetic behavior and movement with colorful glass compositions to boost occupants' daylight performance. The research is being carried out using a multidisciplinary method that includes general morphological analysis (GMA) and a kinetic design strategy. The methodology uses GMA for parametric exploration between interconnected principles of Orosi windows and biomimetic lessons of Morpho butterfly wings to make a matrix of relationships between parameters. The kinetic façade changes the depth and scale of the hexagonal modular elements across a south direction in Iran. Moreover, the most optimal depth and scale of the hexagonal grid regarding the daylight performance metrics have been used to change the composition of colored glasses by periodic movements due to dynamic sun-timing and occupant's positions. The annual daylight metrics analysis of all kinetic alternatives represents high daylight performance by UDI, EUDI, sDA of 93.5, 6.29, and 78.37, respectively. The facade increases the amount of UDI up to 5.61 times regarding the base case while decreasing the amount of EUDI in the room by an average of 91.8%. Concerning predicting the risk of glare, as a point in time metric, the façade prevents visual discomfort by keeping most scenarios in the imperceptible range.
... Universal parameters of the grid-like units on the surface, like the widths and distances of ridges and crossribs, are various when comparing previous reports of butterfly wing scales (Yoshida & Emoto, 2010;Dhungel & Otaki, 2014;Davis et al., 2020;Kochling et al., 2020). These variations might lead to a change in cooling efficiency. ...
The radiative cooling of butterfly wing scales hierarchy has great value in understanding how poikilotherms adapt to the environment and developing bionic materials. However, it remains unclear what the cooling system is like and how the variation of hierarchy affects the cooling efficiency. Therefore, the correlation between the variations of the structure and emissivity of scale hierarchy is thoroughly investigated in Tirumala limniace (Cramer, 1775), whose thermal properties are highly heterogeneous among different wings and regions but similar between males and females. Patterns were deduced from the biological and model simulation experiments. The scale hierarchy varies at the micro- to nanolevel on both surface and section, corresponding to the variating emissivity. Scales on wing veins and margins have large nanostructured units with small lumens and are distinctly thickened, which bring extraordinarily high emissivity. The variations of light and dark scales, respectively, lead to the high emissivity of the middle region of wings and the front wings. Generally, the elevation of the inner surface area and the thickness of the chitin is the key to enhancing the cooling efficiency. For the first time, the effects of the variation of hierarchy toward emissivity of the mid-infrared spectrum are systematically clarified. It is demonstrated that wing scales integrally differentiate in coping with the heterogeneous cooling needs, which may benefit in balancing multifunctions and the development toward the adaptation to the abiotic environment. The study provides insights into the comprehensive thermoregulation system of butterflies and the further development of radiative cooling materials.
... This point assists in dry self-cleaning and improves aerodynamics. The lowest value (0.15) is found in Morpho didius [25]. These ratios influence the adherence of ZnO/nicotine molecules. ...
... The butterfly wing scales exposed with ZnO/nicotine clearly show a partial bleaching influence (Figure 2), that is, partial extraction of the short-wavelength absorbing pigment so that the absorbance of the partially bleached wing scales provides a much more refined picture. The absorbance is still high because the excitation light is absorbed by the ommochrome pigment in addition to melanin [25]. In Figure 4, the redshift of absorbance is detectable after the growth of ZnO/nicotine molecules over the butterfly wing scales. ...
This study aimed to elucidate the optical functions of naturally butterfly wing scales via precise control of morphology as an effective photonic sensor and confirm the content of metal oxide nanoparticles in surrounding nicotine. Metal oxide nanoparticles mixed with nicotine were deposited on the wing scales through the spin-coating method and hence investigated using optical microscopy and spectroscopy. Experimental results demonstrated that absorption intensities of ZnO and TiO2 mixed with nicotine on Danaus genutia were remarkably enhanced. Due to the relatively high concentration of zinc found in e-cigarette aerosol, the intensity of ZnO/nicotine modelled as aerosol adsorption on Danaus genutia, further held a certain linear relationship with the concentration of ZnO. The limit of detection of ZnO was as low as 1 nM. The working mechanism of our sensor was explained through the molecular adsorption after H-bond formation of ZnO/nicotine molecules as high-index materials on the wing scales of Danaus genutia without aggregation. This photonic sensor is an alternative to the present-day methods for the rapid test of ZnO content, which is very simple without complicated instrumentation. Furthermore, our method might become a starting point for the advancement of portable instruments for onsite ZnO detection.
Organismal colours have long captivated and inspired naturalists and scientists. Since colours depend on species' life history, it is expected that they respond to environmental changes, especially in an increasingly anthropized world. Aiming to evaluate how this trait responds to different anthropogenic disturbances, we assess wing colour aspects of fruit-feeding butterflies sampled in Atlantic Forest remnants. These remnants, with well-defined understory and canopy, are crossed by roads and trails acting as artificial edges and besiege exotic pine silviculture areas of different ages, representing landscapes commonly found in the subtropical Atlantic Forest. Through standardised photographs, we obtained colour measurements of brightness, saturation, contrast, colour diversity and heterogeneity, plus the presence of eyespots and iridescence for dorsal and ventral wing surfaces of 47 butterfly species, and evaluated their distribution and abundance in the distinct environments. We hypothesise colour variables will differ in each environment due to their biotic and abiotic differences, being less diverse in anthropized ones due to the pressures generated by disturbances. As expected, different natural environments have diverse and unique colour traits that are not found in anthropized ones; however, artificial edges present brighter and even more diverse colours. Pine silviculture areas, despite differing succession stages, all have decreases in colour diversity. We demonstrate that different anthropogenic actions can lead the most colourful species to disappear. Therefore, we argue butterfly colouration can be seen as a bioindicator, representing a useful form of functional biodiversity, providing conservation status and facilitating communication with the general public.
Butterfly scales are among the richest natural sources of optical nanostructures, which produce structural color and iridescence. Several recurring nanostructure types have been described, such as ridge multilayers, gyroids and lower lamina thin films. While the optical mechanisms of these nanostructure classes are known, their phylogenetic distributions and functional ranges have not been described in detail. In this Review, we examine a century of research on the biological production of structural colors, including their evolution, development and genetic regulation. We have also created a database of more than 300 optical nanostructures in butterflies and conducted a meta-analysis of the color range, abundance and phylogenetic distribution of each nanostructure class. Butterfly structural colors are ubiquitous in short wavelengths but extremely rare in long wavelengths, especially red. In particular, blue wavelengths (around 450 nm) occur in more clades and are produced by more kinds of nanostructures than other hues. Nanostructure categories differ in prevalence, phylogenetic distribution, color range and brightness. For example, lamina thin films are the least bright; perforated lumen multilayers occur most often but are almost entirely restricted to the family Lycaenidae; and 3D photonic crystals, including gyroids, have the narrowest wavelength range (from about 450 to 550 nm). We discuss the implications of these patterns in terms of nanostructure evolution, physical constraint and relationships to pigmentary color. Finally, we highlight opportunities for future research, such as analyses of subadult and Hesperid structural colors and the identification of genes that directly build the nanostructures, with relevance for biomimetic engineering.
Multiple global trends and drivers have resulted in a steep escalation of tech-socio-economic inequities in basic human needs across the industrialized as well as industrializing nations. This escalation is paralleled by the growing trend of novel and simple frugal innovations for meeting basic human needs, which are applied across various communities in the world towards bridging gaps of inequity. It is noteworthy that frugal innovations are abundantly observed in the biological designs in nature. This paper is aimed at understanding the methodology of frugal engineering behind the resulting frugal manufacturing innovations through discovering the cross-section of frameworks of biological designs in nature and equitable social innovations. Authors have applied the framework of biological designs as these designs are observed to deliver multifunctionality, resilience, and sustainability which are key to a frugal and equitable innovation platform and achieved by the frugal engineering process. As water is one of the most basic human needs, this paper uses water as an illustrative example to understand the frugal engineering process. Authors discuss designs in nature from cactus, tree roots, and human skin, and design parallels in related frugal innovations namely in fog capturing nets, ice-stupa, and Zeer (pot-in-a-pot), respectively, for equitable water access. The authors propose a resulting methodology for frugal engineering.
