Ecological drivers of ultraviolet colour evolution in snakes.

Ultraviolet (UV) colour patterns invisible to humans are widespread in nature. However, research bias favouring species with conspicuous colours under sexual selection can limit our assessment of other ecological drivers of UV colour, like interactions between predators and prey. Here we demonstrate widespread UV colouration across Western Hemisphere snakes and find stronger support for a predator defence function than for reproduction. We find that UV colouration has evolved repeatedly in species with ecologies most sensitive to bird predation, with no sexual dichromatism at any life stage. By modelling visual systems of potential predators, we find that snake conspicuousness correlates with UV colouration and predator cone number, providing a plausible mechanism for selection. Our results suggest that UV reflectance should not be assumed absent in "cryptically coloured" animals, as signalling beyond human visual capacities may be a key outcome of species interactions in many taxa for which UV colour is likely underreported.

Pollinator shift ensures reproductive success in a camouflaged alpine plant.

BACKGROUND AND AIMS: There are intrinsic conflicts between signalling to mutualists and concealing (camouflaging) from antagonists. Like animals, plants also use camouflage as a defence against herbivores. However, this can potentially reduce their attractiveness to pollinators. METHODS: Using Fritillaria delavayi, an alpine camouflaged plant with inter-population floral colour divergence, we tested the influence of floral trait differences on reproduction. We conducted pollination experiments, measured floral morphological characteristics, estimated floral colours perceived by pollinators, analysed floral scent and investigated reproductive success in five populations. KEY RESULTS: We found that the reproduction of F. delavayi depends on pollinators. Under natural conditions, a flower-camouflaged population had 100 % fruit set and similar seed set to three out of four yellow-flowered populations. Bumblebees are important pollinators in the visually conspicuous yellow-flowered populations, whereas flies are the only pollinator in the flower-camouflaged population, visiting flowers more frequently than bumblebees. The camouflaged flowers cannot be discriminated from the rock background as perceived by pollinators, but may be located by flies through olfactory cues. CONCLUSIONS: Collectively, our results demonstrate that the flower-camouflaged population has different reproductive traits from the visually conspicuous yellow-flowered populations. A pollinator shift from bumblebees to flies, combined with high visitation frequency, compensates for the attractiveness disadvantage in camouflaged plants.

The physical and cellular mechanism of structural color change in zebrafish.

Many animals exhibit remarkable colors that are produced by the constructive interference of light reflected from arrays of intracellular guanine crystals. These animals can fine-tune their crystal-based structural colors to communicate with each other, regulate body temperature, and create camouflage. While it is known that these changes in color are caused by changes in the angle of the crystal arrays relative to incident light, the cellular machinery that drives color change is not understood. Here, using a combination of 3D focused ion beam scanning electron microscopy (FIB-SEM), micro-focused X-ray diffraction, superresolution fluorescence light microscopy, and pharmacological perturbations, we characterized the dynamics and 3D cellular reorganization of crystal arrays within zebrafish iridophores during norepinephrine (NE)-induced color change. We found that color change results from a coordinated 20° tilting of the intracellular crystals, which alters both crystal packing and the angle at which impinging light hits the crystals. Importantly, addition of the dynein inhibitor dynapyrazole-a completely blocked this NE-induced red shift by hindering crystal dynamics upon NE addition. FIB-SEM and microtubule organizing center (MTOC) mapping showed that microtubules arise from two MTOCs located near the poles of the iridophore and run parallel to, and in between, individual crystals. This suggests that dynein drives crystal angle change in response to NE by binding to the limiting membrane surrounding individual crystals and walking toward microtubule minus ends. Finally, we found that intracellular cAMP regulates the color change process. Together, our results provide mechanistic insight into the cellular machinery that drives structural color change.

Female cichlids mate with novel androgen receptor mutant males that lack coloration.

A key challenge in animal behavior is disentangling the social stimuli that drive conspecific behaviors. For some species, like teleost fish, putative sexual signaling cues are inextricably linked to others, making it difficult to parse the precise roles distinct signals play in driving conspecific behaviors. In the African cichlid Astatotilapia burtoni, males are either dominant or subordinate, wherein bright coloration, territoriality, and courtship behavior inextricably correlate positively with rank. Here, we leveraged androgen receptor (AR) mutant male A. burtoni that lack dominance-typical coloration but not behavior to isolate the role of male coloration in driving female mating behaviors in this species. We found in independent behavioral assays that females behave aggressively towards AR mutant but not WT males, yet still mated with both types of males. Females showed enhanced activation of esr2b + cells in the hypothalamus when housed with either mutant or WT males and this activation scaled with spawning activities. Therefore, there is not a simple relationship between male coloration and female mating behaviors in A. burtoni, suggesting independent sensory mechanisms converge on hypothalamic esr2b cells to coordinate behavioral output.

Variation in Ventral Coloration Pattern and Aposematism in Tropical Warty Newts.

Aposematic coloration plays a crucial role in animal defense, and it is shaped by a complex interplay of factors such as physiological limitations and sexual and natural selection. Warty newts within the genus Paramesotriton exhibit significant variation in ventral coloration. In this study, we quantified the percentage of red ventral area to investigate aposematic ventral coloration in Paramesotriton deloustali and P. guangxiensis across eight populations in northern Vietnam. To assess the interaction between predators and the aposematic signals, we conducted experiments employing three types of clay replicas of newts: dorsal, red ventral, and black ventral models. Our findings revealed a significant variation in the red ventral area among different populations. Additionally, a significant correlation was detected between the red ventral area of the newt and the annual temperature range. In clay model experiments, a significant difference in predator attack rates was observed between dorsal and ventral clay models. Interestingly, there was no significant difference in attack rates between red and black ventral types. Our study suggested that the variation in the red ventral area of warty newts is probably influenced by multiple factors, including genetic constraints, sex, ambient environment, and diet. Furthermore, our results supported the effectiveness of displaying aposematic coloration as an antipredator defense mechanism in warty newts. However, variations in body size and the pressure of mammal predation might not play a significant role in determining aposematic coloration.

Evaluation of Color and Pigment Changes in Tomato after 1-Methylcyclopropene (1-MCP) Treatment.

The Polar Qualification System (PQS) was applied on hue spectra fingerprinting to describe color changes in tomato during storage. The cultivar 'Pitenza' was harvested at six different maturity stages, and half of the samples were subjected to gaseous 1-methylcyclopropene (1-MCP) treatment. Reference color parameters were recorded with a vision system colorimeter instrument, and the fruit pigment concentration was assessed with the DA-index®. Additionally, acoustic firmness (Stiffness) was measured. All acquired reference parameters were used to grade fruit in the supply chain. The applied 1-MCP treatments were used to control the ripening of climacteric horticultural produce. Both the DA-index® and stiffness values, presented as chlorophyll concentration and acoustic firmness, showed significant differences among maturity stages and treated and control samples and in their kinetics during storage. The machine vision parameter PQS-X was significantly affected by 1-MCP treatment (F = 10.18, p < 0.01), while PQS-Y was primarily affected by storage time (F = 18.18, p < 0.01) and maturity stage (F = 11.15, p < 0.01). A significant correlation was achieved for acoustic firmness with normalized color (r > 0.78) and PQS-Y (r > 0.80), as well as for the DA-index® (r > 0.9). The observed color changes agreed with the reference measurements. The significant statistical effect on the PQS coordinates suggests that hue spectra fingerprinting with this data compression technique is suitable for quality assessment based on color.

Melanocytes in regenerative medicine applications and disease modeling.

Melanocytes are dendritic cells localized in skin, eyes, hair follicles, ears, heart and central nervous system. They are characterized by the presence of melanosomes enriched in melanin which are responsible for skin, eye and hair pigmentation. They also have different functions in photoprotection, immunity and sound perception. Melanocyte dysfunction can cause pigmentary disorders, hearing and vision impairments or increased cancer susceptibility. This review focuses on the role of melanocytes in homeostasis and disease, before discussing their potential in regenerative medicine applications, such as for disease modeling, drug testing or therapy development using stem cell technologies, tissue engineering and extracellular vesicles.

Environmental colour pattern variation in Mediterranean Podarcis.

BACKGROUND: Lizards of the genus Podarcis are widespread in the Mediterranean region, including islands and island archipelagos. These small-bodied lizards have a predominantly protective green-brown colouration. However, some populations display unusual patterns, in which the colouration is predominantly blue or uniformly black. This study explores the factors that influence this chromatic variation, whether environmental (climate and island conditions) or evolutionary (phylogenetic trait conservatism). The colouration of 1400 individuals (27 species) was analysed in the CIELAB colour space. RESULTS: Pagel's λ indicated that colouration is weakly conserved within phylogenetic lineages. Although the island surface plays a key role in the chromatic variability of these lacertids, geographic isolation and climate hold less influence. The colouration of some small island populations tends to be uniform and dark, possibly due to intense intraspecific competition and lower predatory pressure. CONCLUSIONS: This study highlights the importance of island populations in understanding the processes that favour the emergence of extreme phenotypes in small ectothermic vertebrates.

Human Hair Graying Revisited: Principles, Misconceptions, and Key Research Frontiers.

Hair graying holds psychosocial importance and serves as an excellent model for studying human pigmentation and aging in an accessible miniorgan. Current evidence suggests that graying results from an interindividually varying mixture of cumulative oxidative and DNA damage, excessive mTORC1 activity, melanocyte senescence, and inadequate production of pigmentation-promoting factors in the hair matrix. Various regulators modulate this process, including genetic factors (DNA repair defects and IRF4 sequence variation, peripheral clock genes, P-cadherin signaling, neuromediators, HGF, KIT ligand secretion, and autophagic flux. This leads to reduced MITF- and tyrosinase-controlled melanogenesis, defective melanosome transfer to precortical matrix keratinocytes, and eventual depletion of hair follicle (HF) pigmentary unit (HFPU) melanocytes and their local progenitors. Graying becomes irreversible only when bulge melanocyte stem cells are also depleted, occurring later in this process. Distinct pigmentary microenvironments are created as the HF cycles: early anagen is the most conducive phase for melanocytic reintegration and activation, and only during anagen can the phenotype of hair graying and repigmentation manifest, whereas the HFPU disassembles during catagen. The temporary reversibility of graying is highlighted by several drugs and hormones that induce repigmentation, indicating potential target pathways. We advise caution in directly applying mouse model concepts, define major open questions, and discuss future human antigraying strategies.

Plumage color degradation indicates reproductive effort: an experiment.

Plumage color has traditionally been regarded as a static ornamental trait, but evidence is accumulating for significant color changes without molt that typically reduce the conspicuousness of ornamentation. In some species, the social partner seems to increase its reproductive investment if the color trait is experimentally enhanced, suggesting that color change could act as a signal. However, the information content of this signal is so far unclear. For example, birds in poor condition or making greater effort may deteriorate more severely. We used brood size manipulations to alter the reproductive effort of male and female collared flycatchers Ficedula albicollis. Both sexes showed less severe decline in some reflectance attribute of their white breast when their brood was experimentally reduced. In each sex, greater deterioration of the reflectance trait affected by the manipulation was accompanied by increased feeding rate by the partner. These feeding patterns do not prove, but are consistent with, a compensatory response by the partner to induced degradation. The manipulation effects on color change we detected confirm for the first time that plumage color deterioration can indicate current reproductive effort, thereby providing a potential fitness advantage to social partners that react to such deterioration.

Reversing Gray Hair: Inspiring the Development of New Therapies Through Research on Hair Pigmentation and Repigmentation Progress.

Hair graying is a common and visible sign of aging resulting from decreased or absence of melanogenesis. Although it has been established that gray hair greatly impacts people's mental health and social life, there is no effective countermeasure other than hair dyes. It has long been thought that reversal of gray hair on a large scale is rare. However, a recent study reported that individual gray hair darkening is a common phenomenon, suggesting the possibility of large-scale reversal of gray hair. In this article, we summarize the regulation mechanism of melanogenesis and review existing cases of hair repigmentation caused by several factors, including monoclonal antibodies drugs, tyrosine kinase inhibitors (TKIs), immunomodulators, other drugs, micro-injury, and tumors, and speculate on the mechanisms behind them. This review offers some insights for further research into the modulation of melanogenesis and presents a novel perspective on the development of clinical therapies, with emphasis on topical treatments.

Carotenoid ornaments and the spandrels of physiology: a critique of theory to explain condition dependency.

Even as numerous studies have documented that the red and yellow coloration resulting from the deposition of carotenoids serves as an honest signal of condition, the evolution of condition dependency is contentious. The resource trade-off hypothesis proposes that condition-dependent honest signalling relies on a trade-off of resources between ornamental display and body maintenance. By this model, condition dependency can evolve through selection for a re-allocation of resources to promote ornament expression. By contrast, the index hypothesis proposes that selection focuses mate choice on carotenoid coloration that is inherently condition dependent because production of such coloration is inexorably tied to vital cellular processes. These hypotheses for the origins of condition dependency make strongly contrasting and testable predictions about ornamental traits. To assess these two models, we review the mechanisms of production of carotenoids, patterns of condition dependency involving different classes of carotenoids, and patterns of behavioural responses to carotenoid coloration. We review evidence that traits can be condition dependent without the influence of sexual selection and that novel traits can show condition-dependent expression as soon as they appear in a population, without the possibility of sexual selection. We conclude by highlighting new opportunities for studying condition-dependent signalling made possible by genetic manipulation and expression of ornamental traits in synthetic biological systems.

Testing multiple hypotheses on the colour change of treefrogs in response to various external conditions.

Amphibians are famous for their ability to change colours. And a considerable number of studies have investigated the internal and external factors that affect the expression of this phenotypic plasticity. Evidence to date suggests that thermoregulation and camouflage are the main pressures that influence frogs' adaptive colour change responses. However, certain gaps in our knowledge of this phenomenon remain, namely: (i) how do frogs adjust their colour in response to continuously changing external conditions?; (ii) what is the direction of change when two different functions of colour (camouflage and thermoregulation) are in conflict?; (iii) does reflectance in the near-infrared region show thermally adaptive change?; and (iv) is the colour change ability of each frog an individual trait (i.e., consistent within an individual over time)? Using Dryophytes japonicus (Hylidae, Hyla), we performed a series of experiments to answer the above questions. We first showed that frogs' responses to continuously-changing external conditions (i.e., background colour and temperature) were not linear and limited to the range they experience under natural conditions. Second, when a functional conflict existed, camouflage constrained the adaptive response for thermoregulation and vice versa. Third, though both temperature and background colour induced a change in near-infrared reflectance, this change was largely explained by the high correlation between colour (reflectance in the visible spectrum) and near-infrared reflectance. Fourth, within-individual variation in colour change capacity (i.e., the degree of colour change an individual can display) was lower than inter-individual variation, suggesting individuality of colour change capacity; however, we also found that colour change capacity could change gradually with time within individuals. Our results collectively reveal several new aspects of how evolution shapes the colour change process and highlight how variation in external conditions restricts the extent of colour change in treefrogs.

Canalised and plastic components of melanin-based colouration: a diet-manipulation experiment in house sparrows.

Whether melanin-based plumage colouration accurately reflects a bird's quality is still controversial. To better understand potential mechanisms behind the observed variation in plumage colouration, we shifted our attention from a high-level expression of colour to low-level physiological phenomena by targeting the microstructure and pigment content of the feather. In a well-studied model system, the house sparrow (Passer domesticus), we combined an experimental manipulation of birds' physiological condition and availability of resources that are key to the production of the studied colouration (phenylalanine and tyrosine (PT). We found that feathers from sparrows fed with the control diet had noticeably lower values of brightness, suggesting a higher quality of the ornamental "blackness" in comparison to those sampled from birds fed with a PT-reduced diet. Electron paramagnetic resonance (EPR) spectroscopy detected higher melanin concentrations in samples from the control than the PT-reduced group. Our multi-level analysis excluded mechanisms such as barbule density and melanosomes' distribution, clearly pointing to the finest-level proxy of colour: the concentration of melanin in melanosomes themselves. Despite melanins being manufactured by birds endogenously, the efficiency of melanogenesis can be noticeably limited by diet. As a result, the birds' plumage colouration is affected, which may entail consequences in social signalling.

The evolution of sex similarities in social signals: Climatic seasonality is associated with lower sexual dimorphism and greater elaboration of female and male signals in antbirds (Thamnophilidae).

Selection on signals that mediate social competition varies with resource availability. Climate regulates resource availability, which may affect the strength of competition and selection on signals. Traditionally, this meant that more seasonal, colder, or dryer-overall harsher-environments should favor the elaboration of male signals under stronger male-male competition, increasing sexual dimorphism. However, females also use signals to compete; thus, harsher environments could strengthen competition and favor elaboration of signals in both sexes, decreasing sexual dimorphism. Alternatively, harsher environments could decrease sexual dimorphism due to scarcer resources to invest in signal elaboration in both sexes. We evaluated these contrasting hypotheses in antbirds, a family of Neotropical passerines that varies in female and male signals and occurs across diverse climatic regimes. We tested the association of sexual dimorphism of plumage coloration and songs with temperature, precipitation, and their seasonality. We found that greater seasonality is associated with lower sexual dimorphism in plumage coloration and greater elaboration of visual signals in both sexes, but not acoustic signals. Our results suggest that greater seasonality may be associated with convergent elaboration of female and male visual signals, highlighting the role of signals of both sexes in the evolution of sexual dimorphism.

Colours of stress in male Indian rock agamas predict testosterone levels but not performance.

Rapid physiological colour change offers dynamic signalling opportunities that can reveal distinct information to receivers in different contexts. Information content in dynamic colours, however, is largely unexplored. In males of the Indian rock agama (Psammophilus dorsalis), stressful events, including male-male agonistic interactions, induce a colour change, wherein the dorsal band turns yellow and the lateral bands turn orange. We aimed to determine whether these pigment-based dynamic colours convey information about individual quality. Using an agamid-specific visual model, we first quantified the chromatic and achromatic contrasts of each colour component displayed by males during handling stress, which induces the maximal response of aggression-typical colours. We then measured baseline testosterone levels, morphology (body mass and size), and performance measures (bite force and sprint speed) of these lizards. Chromatic contrasts of the dorsal yellow and lateral orange bands, individually and relative to each other (internal pair), were negatively correlated with testosterone levels, while the chromatic contrast of the internal pair was positively correlated with body condition. The lack of an association between colour contrasts and both bite force and sprint speed indicate that the conspicuousness of colours expressed during stressful events, such as agonistic interactions, do not reveal male performance ability. Despite our expectations of a positive relationship with testosterone, morphology (body condition), and performance (bite force, sprint speed), we find that for P. dorsalis, the conspicuousness of stress-induced colours provide only some information about individual quality. We speculate that the dynamicity of physiological colours may influence their function as content-containing signals in social interactions.

The physiological cost of colour change: evidence, implications and mitigations.

Animals benefit from phenotypic plasticity in changing environments, but this can come at a cost. Colour change, used for camouflage, communication, thermoregulation and UV protection, represents one of the most common plastic traits in nature and is categorised as morphological or physiological depending on the mechanism and speed of the change. Colour change has been assumed to carry physiological costs, but current knowledge has not advanced beyond this basic assumption. The costs of changing colour will shape the evolution of colour change in animals, yet no coherent research has been conducted in this area, leaving a gap in our understanding. Therefore, in this Review, we examine the direct and indirect evidence of the physiological cost of colour change from the cellular to the population level, in animals that utilise chromatophores in colour change. Our Review concludes that the physiological costs result from either one or a combination of the processes of (i) production, (ii) translocation and (iii) maintenance of pigments within the colour-containing cells (chromatophores). In addition, both types of colour change (morphological and physiological) pose costs as they require energy for hormone production and neural signalling. Moreover, our Review upholds the hypothesis that, if repetitively used, rapid colour change (i.e. seconds-minutes) is more costly than slow colour change (days-weeks) given that rapidly colour-changing animals show mitigations, such as avoiding colour change when possible. We discuss the potential implications of this cost on colour change, behaviour and evolution of colour-changing animals, generating testable hypotheses and emphasising the need for future work to address this gap.

Thermal Costs and Benefits of Replicated Color Evolution in the White Sands Desert Lizard Community.

AbstractTraits often contribute to multiple functions, complicating our understanding of the selective pressures that influence trait evolution. In the Chihuahuan Desert, predation is thought to be the primary driver of cryptic light coloration in three White Sands lizard species relative to the darker coloration of populations on adjacent dark soils. However, coloration also influences radiation absorption and thus animal body temperatures. We combined comparative physiological experiments and biophysical models to test for thermal consequences of evolving different color morphs in White Sands across the three species. While light and dark morphs have not evolved different physiological heat limits within species, differences in radiation absorption between morphs lead to body temperature differences that impact relative overheating risk and activity patterns. Moreover, for all three species, an idealized morph that matches the White Sands substrate would have considerably less activity time, by approximately 1 month, than existing light morphs. Overall, there are both benefits and costs to greater substrate matching, the balance of which may prevent the evolution of optimal crypsis. Our work highlights the importance of color in dictating thermal performance and the complexity inherent in understanding the evolution of coloration.

Star finches Neochmia ruficauda have a visual preference for white dot patterns: a possible case of trypophilia.

Many animals have polka dot patterns on their body surface, some of which are known to have signalling functions; however, their evolutionary origins remain unclear. Dot patterns can trigger a fear response (trypophobia) in humans and are known to function as aposematic signals in non-human animals, suggesting that dots may deserve attention for biological reasons. Interestingly in many birds, plumage dot patterns serve for social/sexual signalling. To understand their evolution, we have focused on the sensory bias hypothesis, which predicts the role of pre-existing sensory preference driven by natural selection in shaping signal design. Our previous phylogenetic comparative study supported the hypothesis and showed that diet-driven visual preference promoted the evolution of plumage patterns, as there was an evolutionary correlation between termite-eating (white roundish gregarious prey) and the presence of plumage dot patterns in species of the family Estrildidae. This suggests that these species possess an intrinsic preference for dots. To test this, we compared the responses of an Estrildid species with dot plumage pattern (star finch Neochmia ruficauda) towards simultaneously presented monochrome-printed white dot vs white stripe patterns under both food-deprived and -supplied conditions. Overall, star finches preferred dots to stripes. They showed foraging-like behaviours almost only toward dots when hungry and gazed at dots frequently even when food was available, suggesting both hunger-related and hunger-neutral dot preferences. These results are rather surprising, given how strongly the subjects were attracted to abstract dot patterns without organic structure, but provided good support for the sensory bias hypothesis.

Molecular parallelism in signaling function across different sexually selected ornaments in a warbler.

Extravagant ornaments are thought to signal male quality to females choosing mates, but the evidence linking ornament size to male quality is controversial, particularly in cases in which females prefer different ornaments in different populations. Here, we use whole-genome sequencing and transcriptomics to determine the genetic basis of ornament size in two populations of a widespread warbler, the common yellowthroat (Geothlypis trichas). Within a single subspecies, females in a Wisconsin population prefer males with larger black masks as mates, while females in a New York population prefer males with larger yellow bibs. Despite being produced by different pigments in different patches on the body, the size of the ornament preferred by females in each population was linked to numerous genes that function in many of the same core aspects of male quality (e.g., immunity and oxidative balance). These relationships confirm recent hypotheses linking the signaling function of ornaments to male quality. Furthermore, the parallelism in signaling function provides the flexibility for different types of ornaments to be used as signals of similar aspects of male quality. This could facilitate switches in female preference for different ornaments, a potentially important step in the early stages of divergence among populations.