Seeing in the dark: vision and visual behaviour in nocturnal bees and wasps.
Warrant Eric J
The Journal of experimental biology
In response to the pressures of predation, parasitism and competition for limited resources, several groups of (mainly) tropical bees and wasps have independently evolved a nocturnal lifestyle. Like their day-active (diurnal) relatives, these insects possess apposition compound eyes, a relatively light-insensitive eye design that is best suited to vision in bright light. Despite this, nocturnal bees and wasps are able to forage at night, with many species capable of flying through a dark and complex forest between the nest and a foraging site, a behaviour that relies heavily on vision and is limited by light intensity. In the two best-studied species - the Central American sweat bee Megalopta genalis (Halictidae) and the Indian carpenter bee Xylocopa tranquebarica (Apidae) - learned visual landmarks are used to guide foraging and homing. Their apposition eyes, however, have only around 30 times greater optical sensitivity than the eyes of their closest diurnal relatives, a fact that is apparently inconsistent with their remarkable nocturnal visual abilities. Moreover, signals generated in the photoreceptors, even though amplified by a high transduction gain, are too noisy and slow to transmit significant amounts of information in dim light. How have nocturnal bees and wasps resolved these paradoxes? Even though this question remains to be answered conclusively, a mounting body of theoretical and experimental evidence suggests that the slow and noisy visual signals generated by the photoreceptors are spatially summed by second-order monopolar cells in the lamina, a process that could dramatically improve visual reliability for the coarser and slower features of the visual world at night.
10.1242/jeb.015396
Pax6 in Collembola: Adaptive Evolution of Eye Regression.
Hou Ya-Nan,Li Sheng,Luan Yun-Xia
Scientific reports
Unlike the compound eyes in insects, collembolan eyes are comparatively simple: some species have eyes with different numbers of ocelli (1 + 1 to 8 + 8), and some species have no apparent eye structures. Pax6 is a universal master control gene for eye morphogenesis. In this study, full-length Pax6 cDNAs, Fc-Pax6 and Cd-Pax6, were cloned from an eyeless collembolan (Folsomia candida, soil-dwelling) and an eyed one (Ceratophysella denticulata, surface-dwelling), respectively. Their phylogenetic positions are between the two Pax6 paralogs in insects, eyeless (ey) and twin of eyeless (toy), and their protein sequences are more similar to Ey than to Toy. Both Fc-Pax6 and Cd-Pax6 could induce ectopic eyes in Drosophila, while Fc-Pax6 exhibited much weaker transactivation ability than Cd-Pax6. The C-terminus of collembolan Pax6 is indispensable for its transactivation ability, and determines the differences of transactivation ability between Fc-Pax6 and Cd-Pax6. One of the possible reasons is that Fc-Pax6 accumulated more mutations at some key functional sites of C-terminus under a lower selection pressure on eye development due to the dark habitats of F. candida. The composite data provide a first molecular evidence for the monophyletic origin of collembolan eyes, and indicate the eye degeneration of collembolans is caused by adaptive evolution.
10.1038/srep20800
Color vision and color formation in dragonflies.
Futahashi Ryo
Current opinion in insect science
Dragonflies including damselflies are colorful and large-eyed insects, which show remarkable sexual dimorphism, color transition, and color polymorphism. Recent comprehensive visual transcriptomics has unveiled an extraordinary diversity of opsin genes within the lineage of dragonflies. These opsin genes are differentially expressed between aquatic larvae and terrestrial adults, as well as between dorsal and ventral regions of adult compound eyes. Recent topics of color formation in dragonflies are also outlined. Non-iridescent blue color is caused by coherent light scattering from the quasiordered nanostructures, whereas iridescent color is produced by multilayer structures. Wrinkles or wax crystals sometimes enhances multilayer structural colors. Sex-specific and stage-specific color differences in red dragonflies is attributed to redox states of ommochrome pigments.
10.1016/j.cois.2016.05.014
Eye structure, activity rhythms, and visually-driven behavior are tuned to visual niche in ants.
Yilmaz Ayse,Aksoy Volkan,Camlitepe Yilmaz,Giurfa Martin
Frontiers in behavioral neuroscience
Insects have evolved physiological adaptations and behavioral strategies that allow them to cope with a broad spectrum of environmental challenges and contribute to their evolutionary success. Visual performance plays a key role in this success. Correlates between life style and eye organization have been reported in various insect species. Yet, if and how visual ecology translates effectively into different visual discrimination and learning capabilities has been less explored. Here we report results from optical and behavioral analyses performed in two sympatric ant species, Formica cunicularia and Camponotus aethiops. We show that the former are diurnal while the latter are cathemeral. Accordingly, F. cunicularia workers present compound eyes with higher resolution, while C. aethiops workers exhibit eyes with lower resolution but higher sensitivity. The discrimination and learning of visual stimuli differs significantly between these species in controlled dual-choice experiments: discrimination learning of small-field visual stimuli is achieved by F. cunicularia but not by C. aethiops, while both species master the discrimination of large-field visual stimuli. Our work thus provides a paradigmatic example about how timing of foraging activities and visual environment match the organization of compound eyes and visually-driven behavior. This correspondence underlines the relevance of an ecological/evolutionary framework for analyses in behavioral neuroscience.
10.3389/fnbeh.2014.00205
The eyes and vision of butterflies.
Arikawa Kentaro
The Journal of physiology
Butterflies use colour vision when searching for flowers. Unlike the trichromatic retinas of humans (blue, green and red cones; plus rods) and honeybees (ultraviolet, blue and green photoreceptors), butterfly retinas typically have six or more photoreceptor classes with distinct spectral sensitivities. The eyes of the Japanese yellow swallowtail (Papilio xuthus) contain ultraviolet, violet, blue, green, red and broad-band receptors, with each ommatidium housing nine photoreceptor cells in one of three fixed combinations. The Papilio eye is thus a random patchwork of three types of spectrally heterogeneous ommatidia. To determine whether Papilio use all of their receptors to see colours, we measured their ability to discriminate monochromatic lights of slightly different wavelengths. We found that Papilio can detect differences as small as 1-2 nm in three wavelength regions, rivalling human performance. We then used mathematical modelling to infer which photoreceptors are involved in wavelength discrimination. Our simulation indicated that the Papilio vision is tetrachromatic, employing the ultraviolet, blue, green and red receptors. The random array of three ommatidial types is a common feature in butterflies. To address the question of how the spectrally complex eyes of butterflies evolved, we studied their developmental process. We have found that the development of butterfly eyes shares its molecular logic with that of Drosophila: the three-way stochastic expression pattern of the transcription factor Spineless determines the fate of ommatidia, creating the random array in Papilio.
10.1113/JP273917
The evolution and development of eye size in flies.
Casares Fernando,McGregor Alistair P
Wiley interdisciplinary reviews. Developmental biology
The compound eyes of flies exhibit striking variation in size, which has contributed to the adaptation of these animals to different habitats and their evolution of specialist behaviors. These differences in size are caused by differences in the number and/or size of ommatidia, which are specified during the development of the retinal field in the eye imaginal disc. While the genes and developmental mechanisms that regulate the formation of compound eyes are understood in great detail in the fruit fly Drosophila melanogaster, we know very little about the genetic changes and mechanistic alterations that lead to natural variation in ommatidia number and/or size, and thus overall eye size, within and between fly species. Understanding the genetic and developmental bases for this natural variation in eye size not only has great potential to help us understand adaptations in fly vision but also determine how eye size and organ size more generally are regulated. Here we explore the genetic and developmental mechanisms that could underlie natural differences in compound eye size within and among fly species based on our knowledge of eye development in D. melanogaster and the few cases where the causative genes and mechanisms have already been identified. We suggest that the fly eye provides an evolutionary and developmental framework to better understand the regulation and diversification of this crucial sensory organ globally at a systems level as well as the gene regulatory networks and mechanisms acting at the tissue, cellular and molecular levels. This article is categorized under: Establishment of Spatial and Temporal Patterns > Regulation of Size, Proportion, and Timing Invertebrate Organogenesis > Flies Comparative Development and Evolution > Regulation of Organ Diversity.
10.1002/wdev.380