Do insects have cognitive maps?
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Biomedical subjects
Publications and source records attributed to R Wehner.
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Insects can perceive the pattern of polarized light (e-vector pattern) in the sky and use it as a compass. To accomplish this navigational task they employ a specialized part of the retina in which the polarization analysers (ultraviolet receptors) are arranged in a way that mimics, by and large, the distribution of e-vector directions in the sky. By sweeping this 'matched polarization filter's across the sky, the insect translates the complex spatial information provided by the celestial e-vector patterns into rather simple temporal modulations of summed receptor outputs. This mechanism provides a striking example of peripheral coding by matched filtering in sensory systems.
A method for estimating progesterone in rat plasma using gas-liquid chromatography (GLC) with electron-capture detection is described. By means of a column chromatographic technique for the extraction of progesterone from plasma, it is possible to isolate the steroid in such a specific way that, contrary to previously published methods, no additional chromatographic purification and separation steps are needed prior to the final GLC analysis. As a derivative for electron-capture detection we used the 3,20-di-O-pentafluorobenzyloxime of progesterone. This derivative is superior to those used hitherto with regard to sensitivity and stability. The high overall recovery (mean value, 89.4%) makes this method especially convenient for research work when only little plasma is available, or when the concentration of progesterone is very low.
An easy and rapid column chromatographic method for the extraction of steroid hormones from plasma is presented. It permits the nearly quantitative separation of the steroids in a single step with smaller expenditure of time and work as compared to the usual liquid-liquid extraction. Problems of emulsions are eliminated. Furthermore, fractionated separation of hormones from plasma is possible. A simple procedure for the selective extraction of estriol (isolated from estrone and estradiol) is described.
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Individual, isolated rhabdoms from dark-adapted crayfish (Orconectes, Procambarus) were studied with a laterally incident microbeam that could be placed in single stacks of microvilli. Concentration gradients of metarhodopsin along the lengths of microvilli were produced by local bleaches, accomplished by irradiation with small spots of orange light at pH 9 in the presence of glutaraldehyde or formaldehyde. No subsequent redistribution of pigment was observed in the dark, indicating an absence of translational diffusion. On the basis of comparison with other systems, glutaraldehyde, but not formaldehyde (0.75%), would be expected to prevent diffusion of protein in the membrane. Under the same conditions photodichroism is observed, indicating an absence of free Brownian rotation. Photodichroism is larger in glutaraldehyde than in formaldehyde, suggesting that the bifunctional reagent quiets some molecular motion that is present after treatment with formaldehyde. Quantitative comparison of photodichroism with mathematical models indicates that the pigment absorption vectors are aligned within +/- 50 degrees of the microvillar axes and are tilted into the surface of the membrane at an average value of about 20 degrees. The photoconversion of rhodopsin to metarhodopsin is accompanied by an increase in molar extinction of about 20% at the lambda maxand a reorientation of the absorption vector by several degrees. The transition moment either tilts further into the membrane or loses some of its axial orientation, or both. The change in orientation is 3.5 time larger in formaldehyde than in glutaraldehyde.
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Single Golgi impregnated visual cells and their axons were treated from the retina to the first synaptic layer (lamina) in serial electron microscopic sections. This analysis of the retina-lamina projection was undertaken in the upper dorso-median eye region which is known to be involved in the perception of polarized light. For identification of individual visual cells and their fibres a numbering system was used which relates the number of each of the nine visual cells within one retinula to the transverse axis of the rhabdom (TRA) (Fig. 1). Because of the twist of the retinula along its course to the basement membrane (Fig. 6), individual visual cells change their position relative to any eye-constant co-ordinate system. Each axon bundle originating from one 9-celled retinula performs a 180 degrees-rotation before entering the lamina (Fig. 2). The direction of rotation (clockwise or counter-clockwise), which may differ even between adjacent bundles, is related to the two mirror-image types of rhabdoms in the corresponding retinulae and is opposite to the direction of rhabdom twist. Thus, even in small groups of the in total 5500 ommatidia in the eye of the bee, two types of retinulae exist which can be characterized by the geometry of the rhabdoms as well as by the direction of rotation of the retinulae and the axon bundles (Fig. 1). Visual cell numbers 1, 2, and 9, the microvilli of which are oriented in the direction of TRA, form three long visual fibres terminating in the second synaptic layer (medulla). In cross sections of laminar pseudocartridges they appear as the smallest fibre profiles arranged in a symmetrical line of the pseudocartridge bundle (=the transverse axis of the pseudocartridge; TPA) (Fig. 4). The remaining six fibres (cell numbers 3-8) only project to the lamina (short visual fibres; svf's). Two of them (cell numbers 5 and 6), which are the largest cells in the proximal retinula and have their microvilli perpendicularly arranged to TRA (Fig. 1), give rise to the two thickest axons of the underlaying pseudocartridge. In cross sections, t he connecting line of these two axons is orthogonally oriented to TPA (Fig. 5). A model was developed, in which all long visual fibres originate from ultraviolet receptors and in which the polarization sensitivity of the basal ninth cell is enhanced by the twist of the rhabdom. Finally, this model is discussed in light of behavioral experiments revealing the ultraviolet receptors as the only cells involved in the detection of polarized light.
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