Color plays an important role in insect life—many insects forage on colorful flowers and/or have colorful bodies. Accordingly, most insects have multiple spectral types of photoreceptors in their eyes, which gives them the capability to see colors. However, insects cannot perceive colors in the same way as human beings do because their eyes and brains differ substantially. An insect was the first nonhuman animal whose ability to discriminate colors has been demonstrated - in the beginning of the 20th century, von Frisch showed that the honeybee, Apis mellifera, can discriminate blue from any shade of gray. This method, called the gray-card experiment, is an accepted “gold standard” for the proof of color vision in animals. Insect species differ in the combinations of photoreceptors in their eyes, with peak sensitivities in ultraviolet (UV) and/or blue, green, and sometimes red parts of the spectrum. The number of photoreceptor spectral types can be as little as one or two, as in the grasshopper Phlaeoba and the beetle Tribolium, and as many as 10 and more in some species of butterflies and dragonflies. However, not all spectral receptor types are necessarily used for color vison. For example, the butterfly Papilio xuthus uses only four of its eight photoreceptors for color vision. Some insects have separate channels for processing chromatic and achromatic (lightness) information. In the honeybee, the achromatic channel has high spatial resolution and is mediated only by long-wavelength sensitive, or “green,” photoreceptors alone, whereas the spatial resolution of chromatic vision is low and mediated by all three spectral types of photoreceptors. Whether other insects have a similar separation of chromatic and achromatic vision remains uncertain. In contrast to vertebrates, insects do not use distinct sets of photoreceptors for nocturnal vision, and some nocturnal insects can see color at night. Insect photoreceptors are inherently polarization sensitive because of their microvillar organization. Therefore, some insects cannot discriminate changes in polarization of light from changes in its spectral composition. However, many insects sacrifice polarization sensitivity to retain reliable color vision. For example, in the honeybee, polarization sensitivity is eliminated by twisting the rhabdom in most parts of its compound eye except for the dorsal rim area that is specialized in polarization vision. Insects experience color constancy and color-contrast phenomena. Although in humans these aspects of vision are often attributed to cortical processing of color, simple models based on photoreceptor adaptation may explain color constancy and color induction in insects. Color discriminations can be evaluated using a simple model, which assumes that it is limited by photoreceptor noise. This model can help to predict discrimination of colors that are ecologically relevant, such as flower colors for pollinating insects. However, despite the fact that many insects forage on flowers, there is no evidence that insect pollinator vision coevolved with flower colors. The diverse color vision in butterflies appears to adaptively facilitate the recognition of their wing colors.
Natalie Hempel de Ibarra and Misha Vorobyev
Cynthia M. Harley and Mark K. Asplen
Annelid worms are simultaneously an interesting and difficult model system for understanding the evolution of animal vision. On the one hand, a wide variety of photoreceptor cells and eye morphologies are exhibited within a single phylum; on the other, annelid phylogenetics has been substantially re-envisioned within the last decade, suggesting the possibility of considerable convergent evolution. This article reviews the comparative anatomy of annelid visual systems within the context of the specific behaviors exhibited by these animals. Each of the major classes of annelid visual systems is examined, including both simple photoreceptor cells (including leech body eyes) and photoreceptive cells with pigment (trochophore larval eyes, ocellar tubes, complex eyes); meanwhile, behaviors examined include differential mobility and feeding strategies, similarities (or differences) in larval versus adult visual behaviors within a species, visual signaling, and depth sensing. Based on our review, several major trends in the comparative morphology and ethology of annelid vision are highlighted: (1) eye complexity tends to increase with mobility and higher-order predatory behavior; (2) although they have simple sensors these can relay complex information through large numbers or multimodality; (3) polychaete larval and adult eye morphology can differ strongly in many mobile species, but not in many sedentary species; and (4) annelids exhibiting visual signaling possess even more complex visual systems than expected, suggesting the possibility that complex eyes can be simultaneously well adapted to multiple visual tasks.
Elisabetta Tolla, Jonathan H. Pérez, Ian C. Dunn, Simone L. Meddle, and Tyler J. Stevenson
Neuroendocrine mechanisms control the seasonal reproduction in birds and mammals. Seasonal reproduction is ubiquitous across vertebrate and invertebrate species, and its timing is extremely crucial in order to maximize offspring survival. The hypothalamus is the key brain region that integrates environmental cues. An endogenous circannual timer with oscillations that approximate one year is also localized in the hypothalamus. Successful timing of reproduction involves the combination of endogenous internal timers that are entrained by local environmental cues. Photoperiod, or the annual change in day length, is the primary cue most temperate animals use to predict future environmental conditions. Birds are able to detect light through photoreceptors located in the medio-basal hypothalamus. These photoreceptors are localized in neuroendocrine regions and regulate the key reproductive neuropeptide gonadotropin-releasing hormone (GnRH). In mammals, retinal photoreceptors transduce light information the suprachiasmatic nucleus in the hypothalamus, which then modulates the nocturnal duration of melatonin. Melatonin in mammals is crucial, as it regulates the neuroendocrine release of GnRH and downstream transitions across seasonal reproductive states. The tanycyte cells lining the third ventricle (3rdV) of the hypothalamus are the critical node for the integration of internal (i.e., circannual timing) and external (e.g., photoperiod) information necessary for the regulation of seasonal reproduction.
Color perception in macaque monkeys and humans depends on the visually evoked activity in three cone photoreceptors and on neuronal post-processing of cone signals. Neuronal post-processing of cone signals occurs in two stages in the pathway from retina to the primary visual cortex. The first stage, in in P (midget) ganglion cells in the retina, is a single-opponent subtractive comparison of the cone signals. The single-opponent computation is then sent to neurons in the Parvocellular layers of the Lateral Geniculate Nucleus (LGN), the main visual nucleus of the thalamus. The second stage of processing of color-related signals is in the primary visual cortex, V1, where multiple comparisons of the single-opponent signals are made. The diversity of neuronal interactions in V1cortex causes the cortical color cells to be subdivided into classes of single-opponent cells and double-opponent cells. Double-opponent cells have visual properties that can be used to explain most of the phenomenology of color perception of surface colors; they respond best to color edges and spatial patterns of color. Single opponent cells, in retina, LGN, and V1, respond to color modulation over their receptive fields and respond best to color modulation over a large area in the visual field.
Mindaugas Mitkus, Simon Potier, Graham R. Martin, Olivier Duriez, and Almut Kelber
Diurnal raptors (birds of the orders Accipitriformes and Falconiformes), renowned for their extraordinarily sharp eyesight, have fascinated humans for centuries. The high visual acuity in some raptor species is possible due to their large eyes, both in relative and absolute terms, and a high density of cone photoreceptors. Some large raptors, such as wedge-tailed eagles and the Old World vultures, have visual acuities twice as high as humans and six times as high as ostriches—the animals with the largest terrestrial eyes. The raptor retina has rods, double cones, and four spectral types of single cones. The highest density of single cones occurs in one or two specialized retinal regions: the foveae, where, at least in some species, rods and double cones are absent. The deep central fovea allows for the highest acuity in the lateral visual field that is probably used for detecting prey from a large distance. Pursuit-hunting raptors have a second, shallower, temporal fovea that allows for sharp vision in the frontal field of view. Scavenging carrion eaters do not possess a temporal fovea that may indicate different needs in foraging behavior. Moreover, pursuit-hunting and scavenging raptors also differ in configuration of visual fields, with a more extensive field of view in scavengers. The eyes of diurnal raptors, unlike those of most other birds, are not very sensitive to ultraviolet light, which is strongly absorbed by their cornea and lens. As a result of the low density of rods, and the narrow and densely packed single cones in the central fovea, the visual performance of diurnal raptors drops dramatically as light levels decrease. These and other visual properties underpin prey detection and pursuit and show how these birds’ vision is adapted to make them successful diurnal predators.