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The “superpower” of invisibility is a reality and a necessity for many animals that live in featureless environments like the open ocean, where there is nowhere to hide. How do animals achieve invisibility? Many animals match their color patterns to their background, but this strategy is limited when the background scene is dynamic. Transparency allows organisms to match any background all the time. However, it is challenging for an organism to maintain transparency across its entire body volume. To be transparent, tissues must minimize light scattering, both at the surface and within. Until recently, it has been unclear how clear animals with complex bodies (such as many crustaceans with hard cuticles, thick muscles, and other internal organs) minimize such light scattering. This is especially challenging in an environment where light can come from many directions: reflections from downwelling sunlight and bioluminescent searchlights from predators. This review summarizes several recent discoveries of multiple unique adaptations for minimizing light scattering both on the exterior cuticle surface and throughout the body volume of transparent crustaceans, as well as the potential tradeoffs and challenges associated with transparent camouflage.

This feature issue on Light and Color in Nature focuses on the interplay of light with the landscape and other naked-eye phenomena and describes a wide range of natural optical phenomena that can be seen in our natural environment. Results are reported on the experimental and theoretical investigations undertaken to improve understanding of these phenomena.

Continental slope canyons off the United States Atlantic coast remain poorly studied, and in particular, the distributions of pelagic organisms in waters overlying these unique environments are not well documented. During the Early Career Scientist Deep Submergence Training cruise, AT36-EAGER, the distribution of organisms in the water column overlying Hydrographer Canyon, which cuts through the northwestern Atlantic continental margin, was investigated through daytime midwater observations using HOV Alvin (AD4831) at three depths. Mixed swarms of krill and Themisto sp. amphipods were observed at all depths surveyed. Observations centered at 250 m were also dominated by chaetognaths, copepods, and Phronima sp. amphipods, while at 500 and 750 m, the assemblages were dominated by the fishes in the families Paralepididae, Nemichthyidae, and Mytophidae. Additionally, measurements of methane, nitrous oxide, optical properties (absorbance and fluorescence), dissolved organic carbon, and base-extracted particulate organic carbon were made to better characterize the hydrography and biogeochemistry over Hydrographer Canyon. This study was aided by the use of telepresence to communicate between ship and shore-based researchers, and the expedition marks the first use of SMS messaging to communicate between the submersible and the ship. This study demonstrates the capabilities and utility of using Alvin for conducting water column science.

Whole-body transparency, an effective camouflage strategy in many aquatic species, can be disrupted by environmental and/or physiological stressors. We found that tail-flip escape responses temporarily disrupt the transparency of the anemone shrimp Ancylomenes pedersoni. After as few as three tail flips, the previously transparent abdominal muscle became cloudy. Eliciting additional tail flips to the point of exhaustion (16±1 s.e.m.; n=23) resulted in complete opacity, though the original transparency returned after 20–60 min of inactivity. We hypothesized that an exercise-induced increase in blood volume between muscle fibers creates regions of low refractive index fluid between high refractive index muscles, thereby increasing light scattering. We documented pre- and post-contraction perfusion by injecting Alexa Fluor 594 wheat germ agglutinin that labeled sarcolemmal surfaces and endothelial cells in contact with hemolymph and found more hemolymph perfused through the abdominal tissue post-exercise, presumably owing to more capillaries opening. In addition, we altered salinity (to 55‰ and 8‰), perforated the abdomen and injected a vasodilator. All three treatments increased both perfusion and opacity, lending further support to our hypothesis that increased hemolymph perfusion to the abdomen is one mechanism that can disrupt a shrimp's transparency. The fact that transparent shrimp at rest have little to no evidence of perfusion to their abdominal musculature (unlike the opaque shrimp Lysmata pederseni, which had more perfusion even at rest) indicates that they may experience significant physiological trade-offs in order to maintain their transparency; specifically, limiting blood flow and thereby reducing oxygen delivery may result in reduced performance.

Certain insect body and wing surfaces interact with light in unique ways to reflect circularly polarized light or to produce colorful interference patterns. Our study characterizes these optical signatures with the goal of testing whether insects can perceive these optical signatures (i.e. have matched filters).

Summary Transparent zooplankton and nekton are often nearly invisible when viewed under ambient light in the pelagic zone [1–3]. However, in this environment, where the light field is directional (and thus likely to cause reflections), and under the bioluminescent searchlights of potential predators, animals may be revealed by reflections from their body surface [4–7]. We investigated the cuticle surfaces of seven species of hyperiids (Crustacea; Amphipoda) using scanning electron microscopy and found two undocumented features that may reduce reflectance. We found that the legs of Cystisoma spp. (n = 5) are covered with an ordered array of nanoprotuberances 200 ± 20 nm SD in height that function optically as a gradient refractive index material [6, 8, 9]. Additionally, we observed that Cystisoma and six other species of hyperiids are covered with a monolayer of homogenous spheres (diameters ranging from 52 ± 7 nm SD on Cystisoma spp. to 320 ± 15 nm SD on Phronima spp.). Optical modeling using effective medium theory and transfer matrix methods demonstrated that both the nanoprotuberances and the monolayers reduce reflectance by as much as 100-fold, depending on the wavelength and angle of the incident light and the thickness of the gradient layer. Even though we only consider surface reflectance and not internal light scattering, our study demonstrates that these nanoprotuberances and spheres can improve crypsis in a featureless habitat where the smallest reflection can render an animal vulnerable to visual predation.

Circularly polarized light (CPL) reflections are rare in nature. Only a few animal groups—most notably certain stomatopod crustaceans and certain beetles in the family Scarabaeidae—are known to reflect CPL from incident unpolarized light. Here, we examine five species of metallic scarabs in the genus Chrysina that, to the naked human eye, look remarkably similar. Using a spectropolarimetric reflectometer to characterize the complete Mueller matrix elements of the beetles’ elytral surfaces, we found that four of the five species were strongly left-handed circularly polarized (LHCP), and only one scarab species, Chrysina resplendens, had an overall lower degree of polarization and switched from LHCP to right-handed circularly polarized reflectance depending on wavelength.

Cephalopods possess unrivaled camouflage and signaling abilities that are enabled by their sophisticated skin, wherein multiple layers contain chromatophore pigment cells (as part of larger chromatophore organs) and different types of reflective cells called iridocytes and leucophores. The optical functionality of these cells (and thus cephalopod skin) critically relies upon subcellular structures partially composed of unusual structural proteins known as reflectins. Herein, we highlight studies that have investigated reflectins as materials within the context of color-changing coatings. We in turn discuss these proteins' multi-faceted properties, associated challenges, and future potential. Through our presentation of selected case studies, we hope to stimulate additional dialogue and spur further research on photonic technologies based on and inspired by reflectins.

Whole-body transparency, an effective camouflage strategy in many aquatic species, can be disrupted by environmental and/or physiological stressors. We found that tail-flip escape responses temporarily disrupt the transparency of the anemone shrimp

Blubber, the specialized hypodermis of cetaceans, provides thermal insulation through the quantity and quality of lipids it contains. Quality refers to percent lipid content; however, not all lipids are the same. Certain deep-diving cetacean groups possess blubber with lipids - wax esters (WE) - that are not typically found in mammals, and the insulative quality of 'waxy' blubber is unknown. Our study explored the influence of lipid storage class - specifically WE in pygmy sperm whales (Kogia breviceps; N=7) and typical mammalian triacylglycerols in short-finned pilot whales (Globicephala macrorhynchus; N=7) - on blubber's thermal properties. Although the blubber of both species had similar total lipid contents, the thermal conductivity of G. macrorhynchus blubber (0.20±0.01 W m(-1) °C(-1)) was significantly higher than that of K. breviceps (0.15±0.01 W m(-1) °C(-1); P=0.0006). These results suggest that lipid class significantly influences the ability of blubber to resist heat flow. In addition, because the lipid content of blubber is known to be stratified, we measured its depth-specific thermal conductivities. In K. breviceps blubber, the depth-specific conductivity values tended to vary inversely with lipid content. In contrast, G. macrorhynchus blubber displayed unexpected depth-specific relationships between lipid content and conductivity, which suggests that temperature-dependent effects, such as melting, may be occurring. Differences in heat flux measurements across the depth of the blubber samples provide evidence that both species are capable of storing heat in their blubber. The function of blubber as an insulator is complex and may rely upon its lipid class, stratified composition and dynamic heat storage capabilities.

SummaryBlubber, the specialized hypodermis of cetaceans, provides thermal insulation through the quantity and quality of lipids it contains. Quality refers to % lipid content; however, not all lipids are the same. Certain deep-diving cetacean groups possess blubber with lipids - wax esters (WE) - that are not typically found in mammals, and the insulative quality of “waxy” blubber is unknown. Our study explored the influence of lipid storage class – specifically WE in pygmy sperm whales (Kogia breviceps; n=7) and typical mammalian triacylglycerols (TAG) in short-finned pilot whales (Globicephala macrorhynchus; n=7) – on blubber’s thermal properties. Although the blubber of both species had similar total lipid contents, the thermal conductivity of G. macrorhynchus blubber (0.20 ± 0.01 Wm-1°C-1) was significantly higher than that of K. breviceps (0.15 ± 0.01 Wm-1°C-1; P=0.0006). These results suggest that lipid class significantly influences blubber’s ability to resist heat flow. In addition, because blubber’s lipid content is known to be stratified, we measured its depth-specific thermal conductivities. In K. breviceps blubber, the depth-specific conductivity values tended to vary inversely with lipid content. In contrast, G. macrorhynchus blubber displayed unexpected depth-specific relationships between lipid content and conductivity, which suggests that temperature-dependent effects, such as melting, may be occurring. Differences in heat flux measurements across the depth of the blubber samples provide evidence that both species are capable of storing heat in their blubber. The function of blubber as an insulator is complex and may rely upon its lipid class, stratified composition, and dynamic heat storage capabilities.