Insect Color Vision

Author(s):  
Natalie Hempel de Ibarra ◽  
Misha Vorobyev

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.

1998 ◽  
Vol 15 (5) ◽  
pp. 787-797 ◽  
Author(s):  
M. KAMERMANS ◽  
D.A. KRAAIJ ◽  
H. SPEKREIJSE

Color vision is spectrally opponent, suggesting that spectrally opponent neurons, such as the horizontal cells in fish and turtle retinae, play a prominent role in color discrimination. In the accompanying paper (Kraaij et al., 1998), it was shown that the output signal of the horizontal cell system to the cones is not at all spectrally opponent. Therefore, a role for the spectrally opponent horizontal cells in color discrimination seems unlikely. In this paper, we propose that the horizontal cells play a prominent role in color constancy and simultaneous color contrast instead of in color discrimination. We have formulated a model of the cone/horizontal cell network based on measurements of the action spectra of the cones and of the feedback signal of the horizontal cell system to the various cone types. The key feature of the model is (1) that feedback is spectrally and spatially very broad and (2) that the gain of the cone synapse strongly depends on the feedback strength. This makes the synaptic gain of the cones strongly dependent on the spectral composition of the surround. Our model, which incorporates many physiological details of the outer retina, displays a behavior that can be interpreted as color constancy and simultaneous color contrast. We propose that the horizontal cell network modulates the cone synaptic gains such that the ratios of the cone outputs become almost invariant with the spectral composition of the global illumination. Therefore, color constancy appears to be coded in the retina.


2007 ◽  
Vol 24 (3) ◽  
pp. 277-290 ◽  
Author(s):  
M.T. VANLEEUWEN ◽  
C. JOSELEVITCH ◽  
I. FAHRENFORT ◽  
M. KAMERMANS

Color constancy is one of the most impressive features of color vision systems. Although the phenomenon has been studied for decades, its underlying neuronal mechanism remains unresolved. Literature indicates an early, possibly retinal mechanism and a late, possibly cortical mechanism. The early mechanism seems to involve chromatic spatial integration and performs the critical calculations for color constancy. The late mechanism seems to make the color manifest. We briefly review the current evidence for each mechanism. We discuss in more detail a model for the early mechanism that is based on direct measurements of goldfish outer retinal processing and induces color constancy and color contrast. In this study we extrapolate this model to primate retina, illustrating that it is highly likely that a similar mechanism is also present in primates. The logical consequence of our experimental work in goldfish and our model is that the wiring of the cone/horizontal cell system sets the reference point for color vision (i.e., it sets the white point for that animal).


1999 ◽  
Vol 202 (5) ◽  
pp. 497-511 ◽  
Author(s):  
E. Warrant ◽  
K. Bartsch ◽  
C. Günther

The fast-flying day-active hawkmoth Macroglossum stellatarum (Lepidoptera: Sphingidae) has a remarkable refracting superposition eye that departs radically from the classical principles of Exnerian superposition optics. Unlike its classical counterparts, this superposition eye is highly aspherical and contains extensive gradients of resolution and sensitivity. While such features are well known in apposition eyes, they were thought to be impossible in superposition eyes because of the imaging principle inherent in this design. We provide the first account of a superposition eye where these gradients are not only possible, but also produce superposition eyes of unsurpassed quality. Using goniometry and ophthalmoscopy, we find that superposition images formed in the eye are close to the diffraction limit. Moreover, the photoreceptors of the superposition eyes of M. stellatarum are organised to form local acute zones, one of which is frontal and slightly ventral, and another of which provides improved resolution along the equator of the eye. This angular packing of rhabdoms bears no resemblance to the angular packing of the overlying corneal facets. In fact, this eye has many more rhabdoms than facets, with up to four rhabdoms per facet in the frontal eye, a situation which means that M. stellatarum does not possess ommatidia in the accepted sense. The size of the facets and the area of the superposition aperture are both maximal at the frontal retinal acute zone. By having larger facets, a wider aperture and denser rhabdom packing, the frontal acute zone of M. stellatarum provides the eye with its sharpest and brightest image and samples the image with the densest photoreceptor matrix. It is this eye region that M. stellatarum uses to fixate flower entrances during hovering and feeding. This radical departure from classical Exnerian principles has resulted in a superposition eye which has not only high sensitivity but also outstanding spatial resolution.


(i) The dorsal eyes are sensitive to ultraviolet light, which is focused by the corneal lens into crystalline cones in the region where these taper progressively to columns across the clear zone. The action of these columns as light guides can be observed in fixed eyes embedded in polymerized resin. In life the light guide part of the column is surrounded by watery non-cellular haemolymph. (ii) Shadowing the eye surface with a thin wire (three facets wide) while recording from individual receptor units shows that ultraviolet light reaches each receptor by its own facet as in an apposition eye, and not, as in a superposition eye, by a group of many facets. (iii) As shown by the dye Lucifer Yellow injected from a microelectrode, the electrophysiological unit consists of all seven retinula cells in the rhabdom region. Consistent with this tight coupling of retinula cells there is no polarization sensitivity. The peak spectral sensitivity of all single units is at 345-365 nm in the ultraviolet. The acceptance angle is 2.0–2.5°. The sensitivity at the spectral peak to a point source on the optical axis of the unit is poor compared to that in other insects tested with the same equipment. (iv) The acceptance angles (∆ ρ ) in the dorsal eye are at the theoretical minimum for the facet diameter and wavelength from diffraction theory. Ultraviolet vision, therefore, has made possible a reduction in facet size but the interommatidial angle ∆ ϕ is greater than expected from the optimum sampling theory of the diffraction limited compound eye. In fact ∆ ρ ≈ ∆ ϕ ≈ 2°. (v) The dorsal eye is effectively a foveal region with greater sampling density and narrower receptive fields but less overlap of fields than the lateral eye. (vi) The square cones and yellow screening pigment strongly suggest that there is superposition by reflexion of yellow light that spreads between ommatidia across the clear zone. This yellow light might photoreisomerize the visual pigment. Attempts to prove this theory during the recording from single units have so far failed but no better function for the clear zone has been suggested.


2012 ◽  
Vol 2012 ◽  
pp. 1-6 ◽  
Author(s):  
Yuika Saito ◽  
Yoshiro Ohashi ◽  
Prabhat Verma

It is a general belief in apertureless near-field microscopy that the so-calledp-polarization configuration, where the incident light is polarized parallel to the axis of the probe, is advantageous to its counterpart, thes-polarization configuration, where the incident light is polarized perpendicular to the probe axis. While this is true for most samples under common near-field experimental conditions, there are samples which respond better to thes-polarization configuration due to their orientations. Indeed, there have been several reports that have discussed such samples. This leads us to an important requirement that the near-field experimental setup should be equipped with proper sensitivity for measurements withs-polarization configuration. This requires not only creation of effective s-polarized illumination at the near-field probe, but also proper enhancement of s-polarized light by the probe. In this paper, we have examined thes-polarization enhancement sensitivity of near-field probes by measuring and evaluating the near-field Rayleigh scattering images constructed by a variety of probes. We found that thes-polarization enhancement sensitivity strongly depends on the sharpness of the apex of near-field probes. We have discussed the efficient value of probe sharpness by considering a balance between the enhancement and the spatial resolution, both of which are essential requirements of apertureless near-field microscopy.


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