The POL area of the honey bee's eye: behavioural evidence

ABSTRACT. The uppermost dorsal part of the honey bee's compound eye contains a group of c. 150 specialized ommatidia. The photoreceptors of these ommatidia are characterized by a number of anatomical and physiological peculiarities which suggest that they have functional significance for the detection of polarized skylight. Here, we show by painting out different parts of the eye and recording the bee's behavioural responses that the specialized photoreceptors at the dorsal margin of the eye are indeed necessary for detecting polarized skylight and deriving compass information from celestial e-vector patterns. Hence, this group of specialized ommatidia can be called the POL area of the bee's compound eye.     

Pore canals in the cornea of a functionally specialized area of the honey bee's compound eye

Summary The fine structure of the cornea in an anatomically and functionally specialized part of the honey bee's compound eye (dorsal rim area) was examined by light microscopy, transmission electron and scanning electron microscopy. Under incident illumination the cornea appears grey and cloudy, leaving only the centers of the corneal lenses clear. This is due to numerous pore canals that penetrate the cornea from the inside, ending a few m below the outer surface. They consist of (1) a small cylindrical cellular evagination of a pigment cell (proximal), and (2) a rugged-walled, pinetree-shaped extracellular part (distal). The functional significance of these pore canals is discussed. It is concluded that their light scattering properties cause the wide visual fields of the photoreceptor cells measured electrophysiologically in the dorsal rim area, and that this is related to the way this eye region detects polarization in skylight.     

Cognitive physiology: moving the mind's eye before the head's eye

Under natural conditions, shifts of spatial attention are often followed by matching eye movement. Recent evidence suggests that this close coupling is reflected in the ability of the same cortical area to shift eye position and the locus of attention.     

Computation and physiology of sensory-motor processing in eye movements

Performance in sensory-motor behaviors guides our understanding of many of the key computational functions of the brain: the representation of sensory information, the translation of sensory signals to commands for movement, and the production of behavior. Eye movement behaviors have become a valuable testing ground for theories of neural computation because the neural circuitry has been well characterized and the mechanical control of the eye is comparatively simple. Here I review recent studies of eye movement behaviors that provide insight into sensory-motor computation at the single neuron and systems levels. They show that errors in sensory estimation dominate eye movement variability and that the motor system functions to reduce the behavioral impact of its own intrinsic noise sources.     

Hearing in honeybees: the mechanical response of the bee's antenna to near field sound

In the dance language, honeybees use airborne near field sound signals to inform their nestmates of the location of food sources. In behavioral experiments it has recently been shown that Johnston's organ, a chordotonal organ located in the pedicel of the antenna, is used to perceive these sound signals. In the present study the mechanical response of the antennal flagellum to stimulation with near field sound signals was investigated using laser vibrometry. The absolute amplitudes of antennal deflection with acoustical stimulation, the response to sounds of different displacement and velocity amplitudes, the shape of movement of the flagellum, the mechanical frequency response and the mechanical directional sensitivity of the auditory sense organ of the honeybee are described. Using pulsed stimuli simulating the dance sounds it is shown that the temporal pattern of the dance sound is resolved on the level of antennal vibrations.     

Determination of optical parameters of the porcine eye and development of a simulated model

The eye is a very sophisticated system of optical elements for the preeminent sense of vision. In recent years, the number of laser surgery to correct the optical aberration such as myopia or astigmatism has significantly increased. Consequently, improving the knowledge related to the interactions of light with the eye is very important in order to enhance the efficiency of the surgery. For this reason, a complete optical characterization of the porcine eye is presented in this study. Kubelka‐Munk and Inverse Adding‐Doubling methods were applied to spectroscopy measurement to determine the absorption and scattering coefficients. Furthermore, the refractive index has been measured by ellipsometry. The different parameters were obtained for the cornea, lens, vitreous humor, sclera, iris, choroids and eyelid in the visible and infrared region. Thereafter, the results are implemented in a COMSOL Multiphysics® software to create an eye model. This model gives a better understanding of the propagation of light in the eye by adding optical parts such as the iris, the sclera or the ciliary bodies. Two simulations show the propagation of light from the cornea to the retina but also from the sclera to the retina. This last possibility provides a better understanding of light propagation during eye laser surgery such as, for example, transscleral cyclophotocoagulation. Figure: Eye simulation models allow the development of new laser treatments in a simple and safe way for patients. To this purpose, the creation of an eye simulated model based on optical parameters obtained from experimental data is presented in this study. This model will facilitate the understanding of the light propagation inside the porcine eye.     

Emmetropization and optical development of the eye of the barn owl (Tyto alba)

1. We have studied the development of the refractive state in young barn owls (Tyto alba pratincola). Strikingly, the eyes had severe refractive errors shortly after lid opening (which occurred around day 14 after hatching; average from 6 owls: 13.83 ± 1.47 days). Refractive errors vanished in the subsequent one or two weeks (Fig. 1, Fig. 2).
2. Refractive errors did not differ by more than 1 diopter (D) in both eyes of an individual (Fig. 2). Thus, non-visual control of eye growth was sufficient to produce non-random refractions. However, visual input was finally required to adjust the optical system to emmetropia.
3. Using in-vivo A-scan ultrasonography of ocular dimensions (Fig. 4A), photokeratometric measurements of corneal radius of curvature (Fig. 4B), and frozen sections of excised eyes (Fig. 3), we developed paraxial schematic eye models which described age-dependent changes in ocular parameters and were applicable through the ages from lid opening to fledging (Table 1). A schematic eye for the adult barn owl (European subspecies: Tyto alba alba) is also provided. Eye sizes in an adult owl of the American (Tyto alba pratincola) and the European subspecies (T. alba alba) were similar despite of different body weights (500 g and 350 g, respectively).
4. The schematic eyes were used to test which ocular parameters might have caused the recovery from refractive errors. However, none of the ocular dimensions measured underwent obvious changes in their growth curves as visual input became available. Apparently, coordinated growth of several ocular components produced emmetropia.
5. From the schematic eye model, the developmental changes in image brightness and image magnification were calculated (Fig. 5). In barn owl eyes, image size was not quite as extreme as in the tawny owl or the great horned owl. However, the image was larger and the f/number was lower than in diurnal birds of comparable weight (pigeon, chicken). The observation supports a conclusion that image size is maximised in owls to permit a higher degree of photoreceptor convergence for higher light sensitivity at dusk while spatial acuity remains comparable to diurnal birds with smaller eyes.

     

Arrangement of optical axes and spatial resolution in the compound eye of the female blowfly Calliphora

We determined the optical axes of ommatidia in the wild-type female blowfly Calliphora by inspecting the deep pseudopupil in large parts of the compound eye. The resulting map of optical axes allowed us to evaluate the spatial resolution in different parts of the eye in terms of interommatidial angles as well as the density of optical axes, and to estimate the orientation of ommatidial rows along the hexagonal eye lattice. The optical axes are not homogeneously distributed over the eye. In the frontal visual field the spatial resolution is about two times higher than in its lateral part and about three times higher as compared to the eye's dorsal pole region. The orientation of the ommatidial rows along the eye lattice is not the same for different regions of the eye but changes in a characteristic way. The inter-individual variability in the orientation of the ommatidial rows is estimated to be smaller than 8 degrees . The characteristic arrangement of the ommatidial lattice is discussed as an adaptation for efficient evaluation of optic flow as induced during self-motions of the animal.