Avian retinal oil droplets : dietary manipulation of colour vision?

Avian vision is highly developed, with bird retinas containing rod and double-cone photoreceptors, plus four classes of single cones subserving tetrachromatic colour vision. Cones contain an oil droplet, rich in carotenoid pigments (except VS/ultraviolet-sensitive cones), that acts as a filter, substantially modifying light detected by the photoreceptor. Using dietary manipulations, we tested the effects of carotenoid availability on oil droplet absorbance properties in two species: Platycercus elegans and Taeniopygia guttata. Using microspectrophotometry, we determined whether manipulations affected oil droplet carotenoid concentration and whether changes would alter colour discrimination ability. In both species, increases in carotenoid concentration were found in carotenoid-supplemented birds, but only in the double cones. Magnitudes of effects of manipulations were often dependent on retinal location. The study provides, to our knowledge, the first experimental evidence of dietary intake over a short time period affecting carotenoid concentration of retinal oil droplets. Moreover, the allocation of carotenoids to the retina by both species is such that the change potentially preserves the spectral tuning of colour vision. Our study generates new insights into retinal regulation of carotenoid concentration of oil droplets, an area about which very little is known, with implications for our understanding of trade-offs in carotenoid allocation in birds.

Visual pigments, cone oil droplets and ocular media in four species of estrildid finch

A microspectrophotometric study was conducted on the retinal photoreceptors of four species of bird: cut-throat finches (Amadina fasciata), gouldian finches (Erythrura gouldiae), white-headed munias (Lonchura maja) and plum-headed finches (Neochmia modesta). Spectral characteristics of the photoreceptors in all four species were very similar. Rods contained a medium-wavelength-sensitive visual pigment with a wavelength of maximum absorbance at 502-504 nm. Four spectrally distinct types of single cone contained a visual pigment with wavelength of maximum absorbance at either 370-373 nm (ultraviolet-sensitive), 440-447 nm (short-wavelength-sensitive); 500 nm (medium-wavelength-sensitive) or 562-565 nm (long-wavelength-sensitive). Oil droplets in the ultraviolet-sensitive single cones showed no detectable absorption between 330 nm and 800 nm. Oil droplets in the short-, medium-, and long-wavelength-sensitive single cones had cut-off wavelengths at 415-423 nm, 510-520 nm and 567-575 nm, respectively. Double cones contained the visual pigment with wavelength of maximum absorbance at 562-565 nm observed in long-wavelength-sensitive single cones. Only the principal member of the double cone pair contained an oil droplet (P-type, cut-off wavelength at 414-489 nm depending on species and retinal location). Spectral transmittance of the intact ocular media of each species was measured along the optic axis. Wavelengths of 0.5 transmittance for all species were very similar (316-318 nm).

Visual pigments, oil droplets, ocular media and cone photoreceptor distribution in two species of passerine bird: the blue tit (Parus caeruleus L.) and the blackbird (Turdus merula L.)

The spectral absorption characteristics of the retinal photoreceptors of the blue tit (Pal trs caeruleus) and blackbird (Turdus merula) were investigated using microspectrophotometry. The retinae of both species contained rods, double cones and four spectrally distinct types of single cone. Whilst the visual pigments and cone oil droplets in the other receptor types are very similar in both species, the wavelength of maximum sensitivity (lambda(max)) of long-wavelength-sensitive single and double cone visual pigment occurs at a shorter wavelength (557 nm) in the blackbird than in the blue tit (563 nm). Oil droplets located in the long-wavelength-sensitive-single cones of both species cut off wavelengths below 570-573 nm, theoretically shifting cone peak spectral sensitivity some 40 nm towards the long-wavelength end of the spectrum. This raises the possibility that the precise lambda(max) of the long-wavelength-sensitive visual pigment is optimised for the visual function of the double cones. The distribution of cone photoreceptors across the retina, determined using conventional light and fluorescence microscopy also varies between the two species and may reflect differences in their visual ecology.

Tetrachromacy, oil droplets and bird plumage colours

There is a growing body of data on avian eyes, including measurements of visual pigment and oil droplet spectral absorption, and of receptor densities and their distributions across the retina. These data are sufficient to predict psychophysical colour discrimination thresholds for light-adapted eyes, and hence provide a basis for relating eye design to visual needs. We examine the advantages of coloured oil droplets, UV vision and tetrachromacy for discriminating a diverse set of avian plumage spectra under natural illumination. Discriminability is enhanced both by tetrachromacy and coloured oil droplets. Oil droplets may also improve colour constancy. Comparison of the performance of a pigeon's eye, where the shortest wavelength receptor peak is at 410 nm, with that of the passerine Leiothrix, where the ultraviolet-sensitive peak is at 365 nm, generally shows a small advantage to the latter, but this advantage depends critically on the noise level in the sensitivity mechanism and on the set of spectra being viewed.

Absorbance of retinal oil droplets of the budgerigar : sex, spatial and plumage morph-related variation

Intraspecific variation in photoreceptor physiology is known in several vertebrate taxa, but is currently unknown in birds, despite many avian traits varying intraspecifically, and avian visual ecology encompassing a wide range of environments and visual stimuli, which might influence spectral sensitivity. Avian retinal photoreceptors contain light absorbing carotenoid-rich oil droplets that affect vision. Carotenoids are also important plumage components. However, our understanding of the regulation of carotenoids in oil droplets remains rudimentary. Among birds, Melopsittacus undulatus has probably the best-studied colour vision, shows profound intraspecific variation in plumage colour, and increased plasma carotenoids during moult. We used microspectrophotometry to determine whether a relationship exists between oil droplet carotenoid concentration and plumage pigmentation, and tested for sex and spatial variation in droplet absorbance across the retina. Absorbance of one variety of P-type droplets was higher in males. No relationship was found between droplet absorbance and plumage colour. We found a spatial pattern of droplets absorbance across the retina that matched a pattern found in another parrot, and other avian species. Our work provides insights into the development and maintenance of retinal oil droplets and suggests a common mechanism and function for carotenoid deposition in the retina across bird species.

Coloured oil droplets enhance colour discrimination

The eyes of most diurnal reptiles and birds contain coloured retinal filters-oil droplets. Although these filters are widespread, their adaptive advantage remains uncertain. To understand why coloured oil droplets appeared and were retained during evolution, I consider both the benefits and the costs of light filtering in the retina. Oil droplets decrease cone quantum catch and reduce the overlap in sensitivity between spectrally adjacent cones. The reduction of spectral overlap increases the volume occupied by object colours in a cone space, whereas the decrease in quantum catch increases noise, and thus reduces the discriminability of similar colours. The trade-off between these two effects determines the total benefit of oil droplets. Calculations show that coloured oil droplets increase the number of object colours that can be discriminated, and thus are beneficial for colour vision.

Microspectrophotometry of visual pigments and oil droplets in a marine bird, the wedge-tailed shearwater Puffinus pacificus: Topographic variations in photoreceptor spectral characteristics

Microspectrophotometric examination of the retina of a procellariiform marine bird, the wedge-tailed shearwater Puffinus pacificus, revealed the presence of five different types of vitamin A(1)-based visual pigment in seven different types of photoreceptor. A single class of rod contained a medium-wavelength sensitive visual pigment with a wavelength of maximum absorbance (lambda(max)) at 502 nm. Four different types of single cone contained visual pigments maximally sensitive in either the violet (VS, lambda(max) 406 nm), short (SWS, lambda(max) 450 nm), medium (MWS, lambda(max) 503 nm) or long (LWS, lambda(max) 566 nm) spectral ranges. In the peripheral retina, the SWS, MWS and LWS single cones contained pigmented oil droplets in their inner segments with cut-off wavelengths (lambda(cut)) at 445 (C-type), 506 (Y-type) and 562 nm (R-type), respectively. The VS visual pigment was paired with a transparent (T-type) oil droplet that displayed no significant absorption above at least 370 run. Both the principal and accessory members of the double cone pair contained the same 566 nm lambda(max) visual pigment as the LWS single cones but only the principal member contained an oil droplet, which had a lambda(cut) at 413 nm. The retina had a horizontal band or 'visual streak' of increased photoreceptor density running across the retina approximately 1.5 mm dorsal to the top of the pecten. Cones in the centre of the horizontal streak were smaller and had oil droplets that were either transparent/colourless or much less pigmented than at the periphery. It is proposed that the reduction in cone oil droplet pigmentation in retinal areas associated with high visual acuity is an adaptation to compensate for the reduced photon capture ability of the narrower photoreceptors found there. Measurements of the spectral transmittance of the ocular media reveal that wavelengths down to at least 300 nm would be transmitted to the retina.

Tribology of steel/steel interaction in oil-in-water mulsion; a rationale for lubricity

Oil droplets are dispersed in water by an anionic urfactant to form an emulsion. The lubricity of this emulsion in steel/steel interaction is explored in a ball on flat nanotribometer. The droplet size and charge are measured using dynamic light scattering, while the substrate charge density is estimated using the pH titration method. These data are combined to calculate the DLVO forces for the droplets generated for a range of surfactant concentration and two oil to water volume ratios. The droplets have a clear bi-modal size distribution. The study shows that the smaller droplets which experience weak repulsion are situated (at the highest DLVO barrier) much closer to the substrate than thebigger droplets, which experience the same DLVO force, are. We suggest that the smaller droplets thus play a more important role in lubricity than what the bigger droplets do. The largest volume of such small droplets occurs in the 0.5 mM-1 mM range of surfactant concentration and 1% oil to water volume ratio, where the coefficient of friction is also observed to be the least. (C) 2010 Elsevier Inc. All rights reserved.

Oil-in-water lecithin-based microemulsions as a potential delivery system for amphotericin B

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Localization and movement of mineral oil in plants by fluorescence and confocal microscopy

Fluorescence and confocal laser scanning microscopy were explored to investigate the movement and localization of mineral oils in citrus. In a laboratory experiment, fluorescence microscopy observation indicated that when a 'narrow' distillation fraction of an nC23 horticultural mineral oil was applied to adaxial and opposing abaxial leaf surfaces of potted orange [Citrus x aurantium L. (Sapindales: Rutaceae)] trees, oil penetrated steadily into treated leaves and, subsequently, moved to untreated petioles of the leaves and adjacent untreated stems. In another experiment, confocal laser scanning microscopy was used to visualize the penetration into, and the subsequent cellular distribution of, an nC24 agricultural mineral oil in C. trifoliata L. seedlings. Oil droplets penetrated or diffused into plants via both stomata and the cuticle of leaves and stems, and then moved within intercellular spaces and into various cells including phloem and xylem. Oil accumulated in droplets in intercellular spaces and within cells near the cell membrane. Oil entered cells without visibly damaging membranes or causing cell death. In a field experiment with mature orange trees, droplets of an nC23 horticultural mineral oil were observed, by fluorescence microscopy, in phloem sieve elements in spring flush growth produced 4-5 months and 16-17 months after the trees were sprayed with oil. These results suggest that movement of mineral oil in plants is both apoplastic via intercellular spaces and symplastic via plasmodesmata. The putative pattern of the translocation of mineral oil in plants and its relevance to oil-induced chronic phytotoxicity are discussed.

Modelling oil droplet absorption spectra and spectral sensitivities of bird cone photoreceptors

Birds have four spectrally distinct types of single cones that they use for colour vision. It is often desirable to be able to model the spectral sensitivities of the different cone types, which vary considerably between species. However, although there are several mathematical models available for describing the spectral absorption of visual pigments, there is no model describing the spectral absorption of the coloured oil droplets found in three of the four single cone types. In this paper, we describe such a model and illustrate its use in estimating the spectral sensitivities of single cones. Furthermore, we show that the spectral locations of the wavelengths of maximum absorbance (lambda(max)) of the short- (SWS), medium- (MWS) and long- (LWS) wavelength-sensitive visual pigments and the cut-off wavelengths (lambda(cut)) of their respective C-, Y- and R-type oil droplets can be predicted from the lambda(max) of the ultraviolet- (UVS)/violet- ( VS) sensitive visual pigment.

Cone photoreceptor oil droplet pigmentation is affected by ambient light intensity

The cone photoreceptors of many vertebrates contain spherical organelles called oil droplets. In birds, turtles, lizards and some lungfish the oil droplets are heavily pigmented and function to filter the spectrum of light incident upon the visual pigment within the outer segment. Pigmented oil droplets are beneficial for colour discrimination in bright light, but at lower light levels the reduction in sensitivity caused by the pigmentation increasingly outweighs the benefits generated by spectral tuning. Consequently, it is expected that species with pigmented oil droplets should modulate the density of pigment in response to ambient light intensity and thereby regulate the amount of light transmitted to the outer segment. In this study, microspectrophotometry was used to measure the absorption spectra of cone oil droplets in chickens (Gallus gallus domesticus) reared under bright (unfiltered) or dim (filtered) sunlight. Oil droplet pigmentation was found to be dependent on the intensity of the ambient light and the duration of exposure to the different lighting treatments. In adult chickens reared in bright light, the oil droplets of all cone types (except the violet-sensitive single cones, whose oil droplet is always non-pigmented) were more densely pigmented than those in chickens reared in dim light. Calculations show that the reduced levels of oil droplet pigmentation in chickens reared in dim light would increase the sensitivity and spectral bandwidth of the outer segment significantly. The density of pigmentation in the oil droplets presumably represents a trade-off between the need for good colour discrimination and absolute sensitivity. This might also explain why nocturnal animals, or those that underwent a nocturnal phase during their evolution, have evolved oil droplets with low pigment densities or no pigmentation or have lost their oil droplets altogether.

Fractal analysis of flocs formed in edible oil -water emulsions by image analysis techniques

Edible oil is an important contaminant in water and wastewater. Oil droplets smaller than 40 μm may remain in effluent as an emulsion and combine with other contaminants in water. Coagulation/flocculation processes are used to remove oil droplets from water and wastewater. By adding a polymer at proper dose, small oil droplets can be flocculated and separated from water. The purpose of this study was to characterize and analyze the morphology of flocs and floc formation in edible oil-water emulsions by using microscopic image analysis techniques. The fractal dimension, concentration of polymer, effect of pH and temperature are investigated and analyzed to develop a fractal model of the flocs. Three types of edible oil (corn, olive, and sunflower oil) at concentrations of 600 ppm (by volume) were used to determine the optimum polymer dosage and effect of pH and temperature. To find the optimum polymer dose, polymer was added to the oil-water emulsions at concentration of 0.5, 1.0, 1.5, 2.0, 3.0 and 3.5 ppm (by volume). The clearest supernatants obtained from flocculation of corn, olive, and sunflower oil were achieved at polymer dosage of 3.0 ppm producing turbidities of 4.52, 12.90, and 13.10 NTU, respectively. This concentration of polymer was subsequently used to study the effect of pH and temperature on flocculation. The effect of pH was studied at pH 5, 7, 9, and 11 at 30°C. Microscopic image analysis was used to investigate the morphology of flocs in terms of fractal dimension, radius of oil droplets trapped in floc, floc size, and histograms of oil droplet distribution. Fractal dimension indicates the density of oil droplets captured in flocs. By comparison of fractal dimensions, pH was found to be one of the most important factors controlling droplet flocculation. Neutral pH or pH 7 showed the highest degree of flocculation, while acidic (pH 5) and basic pH (pH 9 and pH 11) showed low efficiency of flocculation. The fractal dimensions achieved from flocculation of corn, olive, and sunflower oil at pH 7 and temperature 30°C were 1.2763, 1.3592, and 1.4413, respectively. The effect of temperature was explored at temperatures 20°, 30°, and 40°C and pH 7. The results of flocculation of oil at pH 7 and different temperatures revealed that temperature significantly affected flocculation. The fractal dimension of flocs formed in corn, olive and sunflower oil emulsion at pH 7 and temperature 20°, 30°, and 40°C were 1.82, 1.28, 1.29, 1.62, 1.36, 1.42, 1.36, 1.44, and 1.28, respectively. After comparison of fractal dimension, radius of oil droplets captured, and floc length in each oil type, the optimal flocculation temperature was determined to be 30°C. ^

Formulation characteristics of triacylglycerol oil-in-water emulsions loaded with ergocalciferol using microchannel emulsification

Ergocalciferol is one important form of vitamin D that is needed for proper functioning of the human metabolic system. The study formulates monodisperse food grade ergocalciferol loaded oil-in-water (O/W) emulsions by microchannel emulsification (MCE). The primary characterization was performed with grooved MCE, while the storage stability and encapsulating efficiency (EE) were investigated with straight-through MCE. The grooved microchannel (MC) array plate has 5 × 18 μm MCs, while the asymmetric straight-through MC array plate consists of numerous 10 × 80 μm microslots each connected to a 10 μm diameter circular MC. Ergocalciferol at a concentration of 0.2-1.0% (w/w) was added to various oils and served as the dispersed phase, while the continuous phase constituted either of 1% (w/w) Tween 20, decaglycerol monolaurate (Sunsoft A-12) or β-lactoglobulin. The primary characterization indicated successful emulsification in the presence of 1% (w/w) Tween 20 or Sunsoft A-12. The average droplet diameter increased slowly with the increasing concentration of ergocalciferol and ranged from 28.3 to 30.0 μm with a coefficient of variation below 6.0%. Straight-through MCE was conducted with 0.5% (w/w) ergocalciferol in soybean oil together with 1% (w/w) Tween 20 in Milli-Q water as the optimum dispersed and continuous phases. Monodisperse O/W emulsions with a Sauter mean diameter (d3,2) of 34 μm with a relative span factor of less than 0.2 were successfully obtained from straight-through MCE. The resultant oil droplets were physically stable for 15 days (d) at 4 °C without any significant increase in d3,2. The monodisperse O/W emulsions exhibited an ergocalciferol EE of more than 75% during the storage period.

Konjac- ja galaktoglukomannaanipohjaiset emulsiokalvot

Emulsiokalvolla tarkoitetaan kalvoa, joka on valmistettu haihduttamalla ylimääräinen vesi pois emulsiosta. Polysakkaridipohjainen emulsiokalvo koostuu kalvonmuodostuspolysakkaridista, rasvasta, emulgointiaineesta ja pehmittimestä. Kirjallisuusosassa selvitettiin, mitä raaka-aineita polysakkaridipohjaisissa emulsiokalvoissa käytetään ja mitkä tekijät vaikuttavat emulsiokalvojen vesihöyrynläpäisevyyteen ja mekaanisiin ominaisuuksiin. Tutkimuksen kokeellisen osan tavoitteena oli selvittää, miten konjac-glukomannaani (KGM) ja kuusen galaktoglukomannaani (GGM) soveltuvat emulsiokalvon raaka-aineiksi. Lisäksi selvitettiin, miten rasvan tyyppi ja rasvapitoisuus vaikuttavat GGM-KGM-pohjaisten emulsiokalvojen mekaanisiin ominaisuuksiin ja vesihöyrynläpäisevyyteen. Mehiläisvahasta, mäntyöljystä ja rypsiöljystä valmistettiin emulsiokalvot, joissa oli 30 %:n (paino-% GGM:sta) rasvapitoisuudet. Lisäksi mehiläisvahasta valmistettiin emulsiokalvot, joissa oli 10 ja 50 % mehiläisvahaa. Emulsiokalvoja verrattiin vertailukalvoon, jossa ei ollut rasvaa. Kalvoissa käytetty KGM:n ja GGM:n suhde oli 1:1. Kalvoista mitattiin vesihöyrynläpäisevyys ja -läpäisynopeus, vetolujuus, Youngin moduuli ja murtovenymä. Näiden lisäksi kalvojen poikkileikkaus kuvattiin pyyhkäisyelektronimikroskoopilla. GGM ja KGM soveltuvat emulsiokalvon raaka-aineiksi. Huoneenlämpötilassa kuivatuista kalvoista saatiin tasaisemman näköisiä kuin lämpökaapissa kuivatuista. Pyyhkäisyelektronimikroskooppikuvissa vahapisarat olivat öljypisaroita pienempiä, mikä mahdollisesti vaikutti siihen, että vahapisarat pysyivät paremmin kiinnittyneenä kalvomatriisissa. Öljypisaroiden koko oli kalvoissa noin 10 ?m ja vahapisaroiden 2–6 ?m. Vesihöyrynläpäisynopeus oli pienin 50 %:n mehiläisvahakalvolla (p < 0,05). Vesihöyrynläpäisevyys laski lineaarisesti mehiläisvahapitoisuuden suurentuessa. Öljykalvot ja 10 %:n mehiläisvahakalvo eivät eronneet tilastollisesti merkitsevästi vesihöyrynläpäisevyyden suhteen vertailukalvosta. Pienin vetolujuus ja Youngin moduuli oli 50 %:n mehiläisvahakalvolla. Vertailukalvo oli kestävin ja jäykin. Murtovenymän suhteen kalvot eivät eronneet toisistaan tilastollisesti merkitsevästi. Tutkimuksessa onnistuttiin valmistamaan GGM-KGM-pohjaisia emulsiokalvoja, jotka pidättivät vesihöyryä vertailukalvoa paremmin ja silti säilyttivät mekaaniset ominaisuutensa kohtuullisen hyvin.