Biomedical subjects

D G Stavenga

Publications and source records attributed to D G Stavenga.

At least 19 recordsLinked to original sources

PubMed2007

Sexual dichroism and pigment localization in the wing scales of Pieris rapae butterflies.

The beads in the wing scales of pierid butterflies play a crucially important role in wing coloration as shown by spectrophotometry and scanning electron microscopy (SEM). The beads contain pterin pigments, which in Pieris rapae absorb predominantly in the ultraviolet (UV). SEM demonstrates that in the European subspecies Pieris rapae rapae, both males and females have dorsal wing scales with a high concentration of beads. In the Japanese subspecies Pieris rapae crucivora, however, only the males have dorsal wing scales studded with beads, and the dorsal scales of females lack beads. Microspectrophotometry of single scales without beads yields reflectance spectra that increase slightly and monotonically with wavelength. With beads, the reflectance is strongly reduced in the UV and enhanced at the longer wavelengths. By stacking several layers of beaded scales, pierid butterflies achieve strong colour contrasts, which are not realized in the dorsal wings of female P. r. crucivora. Consequently, P. r. crucivora exhibits a strong sexual dichroism that is absent in P. r. rapae.

Animals↗

PubMed2006

Light on the moth-eye corneal nipple array of butterflies.

The outer surface of the facet lenses in the compound eyes of moths consists of an array of excessive cuticular protuberances, termed corneal nipples. We have investigated the moth-eye corneal nipple array of the facet lenses of 19 diurnal butterfly species by scanning electron microscopy, transmission electron microscopy and atomic force microscope, as well as by optical modelling. The nipples appeared to be arranged in domains with almost crystalline, hexagonal packing. The nipple distances were found to vary only slightly, ranging from about 180 to 240 nm, but the nipple heights varied between 0 (papilionids) and 230 nm (a nymphalid), in good agreement with previous work. The nipples create an interface with a gradient refractive index between that of air and the facet lens material, because their distance is distinctly smaller than the wavelength of light. The gradient in the refractive index was deduced from effective medium theory. By dividing the height of the nipple layer into 100 thin slices, an optical multilayer model could be applied to calculate the reflectance of the facet lenses as a function of height, polarization and angle of incidence. The reflectance progressively diminished with increased nipple height. Nipples with a paraboloid shape and height 250 nm, touching each other at the base, virtually completely reduced the reflectance for normally incident light. The calculated dependence of the reflectance on polarization and angle of incidence agreed well with experimental data, underscoring the validity of the modelling. The corneal nipples presumably mainly function to reduce the eye glare of moths that are inactive during the day, so to make them less visible for predators. Moths are probably ancestral to the diurnal butterflies, suggesting that the reduced size of the nipples of most butterfly species indicates a vanishing trait. This effect is extreme in papilionids, which have virtually absent nipples, in line with their highly developed status. A similar evolutionary development can be noticed for the tapetum of the ommatidia of lepidopteran eyes. It is most elaborate in moth-eyes, but strongly reduced in most diurnal butterflies and absent in papilionids.

Animals↗

PubMed2006

Partial coherence and other optical delicacies of lepidopteran superposition eyes.

Superposition eyes are generally thought to function ideally when the eye is spherical and with rhabdom tips in the focal plane of the imaging optics of facet lenses and crystalline cones. Anatomical data as well as direct optical measurements demonstrate that the superposition eyes of moths and skippers often deviate severely from the expected ideal case. Part of the deviation has been attributed to diffraction at the single facet lens, which was taken to be an essential limit to spatial resolution, because light traveling through different facet lenses was assumed to be incoherent. By considering the two-dimensional facet lens lattice, it is here demonstrated that many facets within a superposition aperture transmit coherent light, allowing a much sharper image than possible with single facet lens diffraction. Partial coherence therefore is an important aspect of superposition imaging. It is argued that broadening of the photoreceptor acceptance angles occurs because of optical errors in the facet lens-crystalline cone system other than diffraction. The transmittance of the superposition aperture of moths and skippers is improved by the corneal nipple arrays of the facet lenses, but quantitative assessment shows that the effect is minor.

Animals↗

PubMed2004

Butterfly wing colours: scale beads make white pierid wings brighter.

The wing-scale morphologies of the pierid butterflies Pieris rapae (small white) and Delias nigrina (common jezabel), and the heliconine Heliconius melpomene are compared and related to the wing-reflectance spectra. Light scattering at the wing scales determines the wing reflectance, but when the scales contain an absorbing pigment, reflectance is suppressed in the absorption wavelength range of the pigment. The reflectance of the white wing areas of P. rapae, where the scales are studded with beads, is considerably higher than that of the white wing areas of H. melpomene, which has scales lacking beads. The beads presumably cause the distinct matt-white colour of the wings of pierids and function to increase the reflectance amplitude. This will improve the visual discrimination between conspecific males and females.

Animals↗

PubMed2004

Angular and spectral sensitivity of fly photoreceptors. III. Dependence on the pupil mechanism in the blowfly Calliphora.

A wave optics model for the facet lens-rhabdomere system of fly eyes is used to analyze the dependence of the angular and spectral sensitivity of R1-6 photoreceptors on the pupil mechanism. This assembly of light-absorbing pigment granules in the soma interacts with the waveguide modes propagating in the rhabdomere. A fly rhabdomere carries two modes in the middle wavelength range and four modes at short wavelengths, depending on the rhabdomere diameter and the angle of the incident light flux. The extension of the mode to outside the rhabdomere strongly depends on wavelength, and this dependence plays a determinant role in the light control function of the pupil. The absorbance spectrum of the pigment in the pupil granules is severely depressed at short wavelengths by waveguide effects, resulting in a distinct blue peak. Accordingly, pupil closure suppresses the photoreceptor's spectral sensitivity much more in the blue-green than in the UV. The pupil only narrows the angular sensitivity at short wavelengths. The geometrical size of the rhabdomere governs the angular sensitivity of fly photoreceptors in the dark-adapted state, but diffraction takes over in the fully light-adapted state.

Animals↗

PubMed2003

Angular and spectral sensitivity of fly photoreceptors. II. Dependence on facet lens F-number and rhabdomere type in Drosophila.

A wave-optical model for the integrated facet lens-rhabdomere system of fly eyes is used to calculate the effective light power in the rhabdomeres when the eye is illuminated with a point light source or with an extended source. Two rhabdomere types are considered: the slender rhabdomeres of R7,8 photoreceptors and the wider, but tapering R1-6 rhabdomeres. The angular sensitivities of the two rhabdomere types have been calculated as a function of F-number and wavelength by fitting Gaussian functions to the effective light power. For a given F-number, the angular sensitivity broadens with wavelength for the slender rhabdomeres, but it stays approximately constant for the wider rhabdomeres. The integrated effective light power increases with the rhabdomere diameter, but it is for both rhabdomere types nearly independent of the light wavelength and F-number. The results are used to interpret the small F-number of Drosophila facet lenses. Presumably the small head puts a limit to the size of the facet lens and favors a short focal length.

Algorithms↗

PubMed2002

Angular and spectral sensitivity of fly photoreceptors. I. Integrated facet lens and rhabdomere optics.

Three optical components of a fly's eye determine the angular sensitivity of the photoreceptors: the light diffracting facet lens, the wave-guiding rhabdomere and the light-absorbing visual pigment in the rhabdomere. How the integrated optical system of the fly eye shapes the angular sensitivity curves is quantitatively analyzed in five steps: (1) scalar diffraction theory for low Fresnel-number lenses is applied to four different facet lenses, with diameter 10, 20, 40, and 80 micro m, respectively, assuming a constant F-number of 2.2; (2) optical waveguide theory is used to calculate waveguide modes propagating in circular cylindrical rhabdomeres with diameter 1.0, 2.0, and 4.0 micro m, respectively; (3) the excitation of waveguide modes is studied with the tip of the waveguide positioned in the focal plane as well as outside this plane; (4) the light absorption from the various propagated modes by the visual pigment in the rhabdomere is calculated as a function of the angle of the incident light wave; and (5) the angular sensitivity of the photoreceptor is obtained by normalizing the total light absorption. Four wavelengths are considered: 300, 400, 500 and 600 nm. The analysis shows that the wavelength dependency of the lens diffraction is strongly compensated by that of the waveguide modes, an effect which is further enhanced by the decrease in light absorption when the mode number increases. The angular sensitivity of fly photoreceptors is robust to defocus and largely wavelength independent for all except very slender rhabdomeres.

Animals↗

PubMed2002

Colour in the eyes of insects.

Many insect species have darkly coloured eyes, but distinct colours or patterns are frequently featured. A number of exemplary cases of flies and butterflies are discussed to illustrate our present knowledge of the physical basis of eye colours, their functional background, and the implications for insect colour vision. The screening pigments in the pigment cells commonly determine the eye colour. The red screening pigments of fly eyes and the dorsal eye regions of dragonflies allow stray light to photochemically restore photoconverted visual pigments. A similar role is played by yellow pigment granules inside the photoreceptor cells which function as a light-controlling pupil. Most insect eyes contain black screening pigments which prevent stray light to produce background noise in the photoreceptors. The eyes of tabanid flies are marked by strong metallic colours, due to multilayers in the corneal facet lenses. The corneal multilayers in the gold-green eyes of the deer fly Chrysops relictus reduce the lens transmission in the orange-green, thus narrowing the sensitivity spectrum of photoreceptors having a green absorbing rhodopsin. The tapetum in the eyes of butterflies probably enhances the spectral sensitivity of proximal long-wavelength photoreceptors. Pigment granules lining the rhabdom fine-tune the sensitivity spectra.

Animals↗

PubMed2002

Opsin cDNA sequences of a UV and green rhodopsin of the satyrine butterfly Bicyclus anynana.

The cDNAs of an ultraviolet (UV) and long-wavelength (LW) (green) absorbing rhodopsin of the bush brown Bicyclus anynana were partially identified. The UV sequence, encoding 377 amino acids, is 76-79% identical to the UV sequences of the papilionids Papilio glaucus and Papilio xuthus and the moth Manduca sexta. A dendrogram derived from aligning the amino acid sequences reveals an equidistant position of Bicyclus between Papilio and Manduca. The sequence of the green opsin cDNA fragment, which encodes 242 amino acids, represents six of the seven transmembrane regions. At the amino acid level, this fragment is more than 80% identical to the corresponding LW opsin sequences of Dryas, Heliconius, Papilio (rhodopsin 2) and Manduca. Whereas three LW absorbing rhodopsins were identified in the papilionid butterflies, only one green opsin was found in B. anynana.

Amino Acid Sequence↗

PubMed2001

Retinal regionalization and heterogeneity of butterfly eyes.

The regional characteristics of the eyes of butterflies from different families have been surveyed using epi-illumination microscopy, utilizing the eyeshine visible due to the tapetum situated proximally to the rhabdom. All butterflies studied have a high spatial acuity in the frontal region. The facet diameter varies slightly across the eye, and the interommatidial angle and the eye parameter p are especially large dorsally. Whereas the ommatidial lattice is generally highly regular, the eyeshine colours distinctly depend on the species. Sometimes the eyeshine is locally uniform, but often it is heterogeneous. It is hypothesized that the regional characteristics as well as the local heterogeneity are adaptations that optimize spectral discrimination.

Animals↗

PubMed2000

Calcium imaging demonstrates colocalization of calcium influx and extrusion in fly photoreceptors.

During illumination, Ca(2+) enters fly photoreceptor cells through light-activated channels that are located in the rhabdomere, the compartment specialized for phototransduction. From the rhabdomere, Ca(2+) diffuses into the cell body. We visualize this process by rapidly imaging the fluorescence in a cross section of a photoreceptor cell injected with a fluorescent Ca(2+) indicator in vivo. The free Ca(2+) concentration in the rhabdomere shows a very fast and large transient shortly after light onset. The free Ca(2+) concentration in the cell body rises more slowly and displays a much smaller transient. After approximately 400 ms of light stimulation, the Ca(2+) concentration in both compartments reaches a steady state, indicating that thereafter an amount of Ca(2+), equivalent to the amount of Ca(2+) flowing into the cell, is extruded. Quantitative analysis demonstrates that during the steady state, the free Ca(2+) concentration in the rhabdomere and throughout the cell body is the same. This shows that Ca(2+) extrusion takes place very close to the location of Ca(2+) influx, the rhabdomere, because otherwise gradients in the steady-state distribution of Ca(2+) should be measured. The close colocalization of Ca(2+) influx and Ca(2+) extrusion ensures that, after turning off the light, Ca(2+) removal from the rhabdomere is faster than from the cell body. This is functionally significant because it ensures rapid dark adaptation.

Animals↗

PubMed2000

Calcium transients in the rhabdomeres of dark- and light-adapted fly photoreceptor cells.

The light response of fly photoreceptor cells is modulated by changes in free Ca(2+) concentration. Fly phototransduction and most processes regulating it take place in or very close to the rhabdomere. We therefore measured the kinetics and the absolute values of the free Ca(2+) concentration in the rhabdomere of fly photoreceptor cells in vivo by making use of the natural optics of the fly's eye. We show that Ca(2+) flowing into the rhabdomere after light stimulation of dark-adapted cells causes fast Ca(2+) transients that reach peak values higher than 200 microM in <20 msec. Approximately 500 msec later, the free Ca(2+) concentration has declined again to approximately 20 microM. The duration of the Ca(2+) transients becomes still shorter, and their size reduced, when the photoreceptor cell is light-adapted. This reduction in duration and size of the Ca(2+) transients is graded with the intensity of the adapting light. The kinetics and absolute values of the free calcium concentration found to occur in the rhabdomere are suitable to mediate the fast feedback signals known to act on the fly phototransduction cascade.

Adaptation, Ocular↗

PubMed1999

Does Ca2+ reach millimolar concentrations after single photon absorption in Drosophila photoreceptor microvilli?

The quantum bump, the elementary event of fly phototransduction induced by the absorption of a single photon, is a small, transient current due to the opening of cation-channels permeable to Ca2+. These channels are located in small, tube-like protrusions of the cell membrane, the microvilli. Using a modeling approach, we calculate the changes of free Ca2+ concentration inside the microvilli, taking into account influx and diffusion of Ca2+. Independent of permeability ratios and Ca2+ buffering, we find that the free Ca2+ concentrations rise to millimolar values, as long as we assume that all activated channels are located in a single microvillus. When we assume that as much as 25 microvilli participate in a single bump, the free Ca2+ concentration still reaches values higher than 80 microM. These very high concentrations show that the microvilli of fly photoreceptors are unique structures in which the Ca2+ signaling is even more extreme than in calcium concentration microdomains very close to Ca2+ channels.

Animals↗

PubMed1999

An ultraviolet absorbing pigment causes a narrow-band violet receptor and a single-peaked green receptor in the eye of the butterfly Papilio.

The distal photoreceptors in the tiered retina of Papilio exhibit different spectral sensitivities. There are at least two types of short-wavelength sensitive receptors: an ultraviolet receptor with a normal spectral shape and a violet receptor with a very narrow spectral bandwidth. Furthermore, a blue receptor, a double-peaked green receptor and a single-peaked green receptor exist. The violet receptor and single-peaked green receptor are only found in ommatidia that fluoresce under ultraviolet illumination. About 28% of the ommatidia in the ventral half of the retina exhibit the UV-induced fluorescence. The fluorescence originates from an ultraviolet-absorbing pigment, located in the most distal 70 microns of the ommatidium, that acts as an absorption filter, both for a UV visual pigment, causing the narrow spectral sensitivity of the violet receptor, and for a green visual pigment, causing a single-peaked green receptor.

Animals↗

PubMed1998

Light dependence of calcium and membrane potential measured in blowfly photoreceptors in vivo.

Light adaptation in insect photoreceptors is caused by an increase in the cytosolic Ca2+ concentration. To better understand this process, we measured the cytosolic Ca2+ concentration in vivo as a function of adapting light intensity in the white-eyed blowfly mutant chalky. We developed a technique to measure the cytosolic Ca2+ concentration under conditions as natural as possible. The calcium indicator dyes Oregon Green 1, 2, or 5N (Molecular Probes, Inc., Eugene, OR) were iontophoretically injected via an intracellular electrode into a photoreceptor cell in the intact eye; the same electrode was also used to measure the membrane potential. The blue-induced green fluorescence of these dyes could be monitored by making use of the optics of the facet lens and the rhabdomere waveguide. The use of the different Ca2+-sensitive dyes that possess different affinities for Ca2+ allowed the quantitative determination of the cytosolic Ca2+ concentration in the steady state. Determining the cytosolic Ca2+ concentration as a function of the adapting light intensity shows that the Ca2+ concentration is regulated in a graded fashion over the whole dynamic range where a photoreceptor cell can respond to light. When a photoreceptor is adapted to bright light, the cytosolic Ca2+ concentration reaches stable values higher than 10 microM. The data are consistent with the hypothesis that the logarithm of the increase in cytosolic Ca2+ concentration is linear with the logarithm of the light intensity. From the estimated values of the cytosolic Ca2+ concentration, we conclude that the Ca2+-buffering capacity is limited. The percentage of the Ca2+ influx that is buffered gradually decreases with increasing Ca2+ concentrations; at cytosolic Ca2+ concentration levels above 10 microM, buffering becomes minimal.

Adaptation, Ocular↗

PubMed1993

Simple exponential functions describing the absorbance bands of visual pigment spectra.

Literature data for visual pigment spectra are formally treated by assuming that the spectra consist of a summation of absorbance bands, that the shape of the bands is invariant according to the Mansfield-MacNichol transform and that this shape is described by simple exponential functions. A new template for constructing visual pigment spectra from peak wavelengths is derived.

Animals↗

PubMed1992

Eye regionalization and spectral tuning of retinal pigments in insects.

The spatial and spectral properties of an eye can often be directly linked to the behaviour and habitat of the animal. In a honey bee (Apis mellifera) society, the drones use the well-developed dorsal part of the eye to detect the queen against the sky during her nuptial flight. Recently it has become clear that the dorsal area of the drone's eye serves its task by cleverly combining a number of optical mechanisms, thus achieving a high spatial acuity as well as a high sensitivity precisely in the wavelength range of interest--the ultraviolet to blue range. Since the various optical specializations in the drone eye have now been recognized, they can be traced in the eyes of other species: thus, the drone eye serves as a model to give a better understanding of the relationship between structure and function of compound eyes in particular, but also of visual systems in general.

Animals↗

PubMed1990

Dioptrics of the facet lenses of male blowflies Calliphora and Chrysomyia.

1. The dioptrics of the facet lenses of two blowfly species, Calliphora erythrocephala and Chrysomyia megacephala, was investigated. Measurements were performed on facet lenses ranging in diameter from 20 to 80 microns. 2. The radius of curvature of the front surface of the facet lenses, measured by microreflectometry, increases approximately linearly with the facet lens diameter. 3. The optical path difference of the facet lens and water, measured by interference microscopy, depends on the distance from the optical axis according to a parabolic function. Average refractive index values, calculated from the optical path difference profile together with estimates of the thickness profile, are between 1.40 and 1.43, with the lowest values in the largest lenses. 4. The F-number calculated from the experimental data ranges from 1.5 to 2.2. It is argued that the range of effective F-numbers is 2.1-2.4.

Animals↗