TASTE PAPILLAE OF THE BLOWFLY.
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Biomedical subjects
Publications and source records attributed to V G DETHIER.
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The electrical responses of the neurons associated with the various types of chemosensory hairs of the blowfly, Phormia regina Meigen, following stimulation by chemical and mechanical means have been studied. The singly innervated chemosensory hairs on the ovipositor, maxillary palpi, and antennae respond vigorously to chemical stimulation, but not to mechanical stimulation. The triply innervated chemosensory hairs on the labellum, tarsus, and wing have two neurons which respond only to chemical stimuli. The third neuron responds only to mechanical stimulation. The differential responses of the two chemosensory neurons to various chemical stimuli following the removal of the tip of the hair suggest that the structures responsible for chemoreception are located throughout the distal processes of these neurons. The response of the third neuron to mechanical stimulation is similar to the response recorded from the neuron associated with one type of tactile hair which responds to motion and not to steady deformation. Recordings have been made from the neurons associated with purely tactile hairs using the cut hair as an extension of the micropipette. The mechanosensory neuron of the wing chemosensory hair is capable of responding at the rate of at least 600 impulses per sec. and may serve to indicate changes in air flow over the wing surfaces during flight to enable the fly to correct the wing camber and attack angle.
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The response of the blowfly Phormia regina to stimulation by aldehydes in the vapor phase has been studied by means of a specially designed olfactometer. The median rejection threshold and the maximum acceptance threshold were selected as criteria of response. For both acceptance and rejection the distribution of thresholds in the population is normal with respect to the logarithm of concentration. When thresholds are expressed as molar concentrations, the values decrease progressively as chain length is increased. There is no attraction beyond decanal and no rejection beyond dodecanal. When thresholds are expressed as activities, most members of the aldehyde series are approximately equally stimulating at rejection and equally stimulating at acceptance. The relationship is most exact over the middle range of chain lengths. There is a tendency for the terminal members to stimulate at higher activities. These relationships are in close agreement with those which were found earlier to apply to the normal aliphatic alcohols. The similarity between the relative actions of the members of the two series suggests that the relation of equal olfactory stimulation at equal thermodynamic activities by homologous aliphatic compounds at least for homologues of intermediate chain length may be of rather general application in olfaction.
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The response of the blowfly Phormia regina to stimulation by alcohols in the vapor phase has been investigated by means of an olfactometer which permitted quantitative control of stimulus concentration. The median rejection threshold was selected as a criterion of response. As was true in the case of contact chemoreception, the distribution of thresholds in the fly population is normal with respect to the logarithm of concentration. In terms of molar concentration the alcohols are rejected at logarithmically decreasing concentration as chain length is increased. Beyond decanol there is no further stimulation. When thresholds are expressed as pressures and plotted against saturated vapor pressures on logarithmic coordinates, the data fit a line the slope of which is not significantly different from 1; i.e., the thresholds vary directly with vapor pressure. Individual threshold values, however, deviate significantly from this line. and the deviation must be ascribed to other factors which have not as yet been identified. When thresholds are expressed as activities, all alcohols are equally stimulating. It appears that the limiting process of olfaction, at least in so far as the normal alcohols are concerned, may involve an equilibrium process. Conformity to this concept is most exact for intermediate members of the series.
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Rejection thresholds of eight primary alcohols applied to the tarsal chemoreceptors of the blowfly Phormia regina Meigen and the ovipositor of Gryllus assimilis Fab. have been determined. Three different solvents for the alcohols have been used: water, ethylene glycol, and mineral oil. The comparative stimulating effectiveness of the alcohols assumes a different aspect with each different solvent. In oil the range of thresholds from methanol to octanol extends over less than one log unit as compared with the corresponding thresholds in water which extend over four log units. In glycol the thresholds extend over two and one half log units only. When water is employed as a solvent, the line which describes the relationship between threshold concentration and chain length of the compound exhibits a sharp break at or near butanol. No such discontinuity is evident when glycol or oil is employed as solvent. This is offered as evidence supporting the hypothesis that the limiting mechanism in tarsal chemoreception involves a two phase system whereby highly water-soluble compounds gain access to the receptor through an aqueous phase and the larger lipoid-soluble molecules chiefly through a lipoid phase. Additional facts which support this idea are gained from data which show that the inflection in the curve occurs at different points with different species of insects and is conspicuously absent in the case of man. When thresholds in aqueous solutions are converted from molar concentrations to activities, it is clear that the relation of equal physiological effect at equal thermodynamic activities does not apply here. The lower members of the series stimulate at progressively increasing activities up to pentanol and then at progressively decreasing activities. Furthermore, the ratio of water threshold to oil threshold exhibits no obvious agreement with the water/oil partition coefficients determined experimentally. These results indicate either that the limiting process of chemoreception in these insects does not depend upon the establishment of an equilibrium or that some kinetic effect is obscuring an underlying relationship which does so depend.
The rejection thresholds of the blowfly Phormia regina Meigen for a selected series of substituted aliphatic hydrocarbons have been determined and an analysis made of the effect on stimulation of increasing the number of like substitutions from one to two, of the interaction of different types of substituent, and of the position of the substituent in the molecule. The order of stimulating efficiency (reciprocal of rejection threshold) of the functional groups which have been studied is: Br > Cl ? CH(3) ? CHO > C=O > OH. There is a consistent decrease in stimulating effect following the introduction of a second hydroxyl group. The introduction of a second C=O has little effect. Introduction of a second halogen is generally followed by a decrease in threshold. Two different species of functional group, one of which tends to increase water solubility and the other to decrease it, tend to counter-balance each other. A single oxygen-containing functional group on the end of the molecule or two of these groups which are terminal reduce the stimulating effect less than when one or more of such groups are subterminal or located elsewhere within the molecule. Branching of the carbon chain decreases effectiveness. Sulfur and oxygen linkages also decrease effectiveness. In general, the length of the free alkyl group largely determines the stimulating effectiveness. Its power is modified to varying degrees by the nature of the attached polar groups. Of all chemical properties examined, solubility alone agrees consistently with the foregoing data. Threshold varies directly with the molar solubility of the compounds in water. The fact that the threshold values for individual compounds are frequently different from those which would be expected solely on the basis of the correlation between threshold and solubility in water suggests that other factors are also concerned in stimulation. These have not yet been identified.
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