Joys and frustrations of doing research.
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Biomedical subjects
Publications and source records attributed to K D Roeder.
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Compared with more complex behavior patterns involving goal selection, evasive behavior of prey animals elicited by the presence of a natural predator is usually considered to be relatively inevitable and stereotyped. Noctuid moths, while flying in the field at night, show two types of evasive response pattern when their tympanic organs detect the ultrasonic cries of marauding insectivorous bats. Since sound is the sole communication channel between prey and predator, this situation invites neurophysiological and behavioral analysis. When, in the course of studying this behavior, moths are removed by stages from the natural circumstances of this interaction their evasion responses become much less invariant; that is, more evitable. The possible survival value of some evitability in this behavior under natural circumstances is discussed. The afferent nerve response of the noctuid auditory organ is highly stable; therefore, the source or sources of this evitability must lie down-stream in the moth central nervous system. Acoustic interneuronal systems within the thoracic ganglia and the brain have been examined for lapses in responsiveness and for other indications of transsynaptic instability that might correlate with the demonstrated behavioral evitability.
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Moths of the genus Celerio, hovering and feeding from blossoms, have been observed to react vigorously to high-pitched sounds. The acoustic receptor appears to involve the labial palps. Its characteristics have been determined from responses made, when the moth is exposed to artificial sound pulses, by an interneuron originating in its head. These responses have been found only in 4 out of 19 sphingid species tested.
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Insect-eating bats find their aerial food by sonar, through emitting ultrasonic chirps and locating sources of echoes. Certain moths have ears sensitive to these chirps and can detect bats well beyond the range of the bats' sonar. On hearing a distant bat, many moths turn and fly directly away from the source of ultrasound. Only one sense cell in each ear of a moth provides the primary nervous information for this response. This article describes my initial attempts to find out how a moth's central nervous system processes the train of chirps reaching its two ears. The ear of a restrained moth is exposed to a sequence of artifically generated ultrasonic pulses that approximates the cries made by a bat. This stimulus can be varied with respect to ultrasonic frequency (pitch), pulse intensity, pulse duration, the interval between pulses, and pulse-train duration. The more sensitive acoustic sense cell responds to all frequencies between about 15,000 and 80,000 cycles per second, but the signal that it transmits to the moth's central nervous system contains no measure of frequency within this range. However, this nerve signal reports variations in the other parameters of the stimulus. The acoustic fiber connects, in the central nervous system, with various nerve cells that transform the signal farther. The signal from a pulse-marker neuron contains no measures of pulse intensity or pulse duration, reporting only changes in interpulse interval and pulse-train duration. A train-marker neuron reports only the duration of the pulse train. The stimulus parameters may be likened to keys, each of which is necessary to gain admittance through a given door but becomes superfluous once this door has been passed. This analogy suggests one of the ways in which a signal is transformed in its passage through the nervous system, and how its specificity is assured in eliciting a given response. In addition to undergoing this kind of transformation, neural signals generated in the two directionally sensitive ears must be combined if a flying moth is to steer a course away from a distant bat. Neurons have been discovered in the central ganglia which summate signals from the right and left ears. Other neurons are inhibited in their activity by stimulation of one ear. The moth may combine signals from these neurons with motor-nerve information on the attitude of its own wings, which act as oscillating baffles modifying its directional acoustic sensitivity 20 to 40 times a second as it flaps an erratic path through the darkness.
A pair of thermistors forming part of a direct current bridge circuit was mounted in the wake of a moth in stationary flight. Differential changes in the thermistors' resistance provided a sensitive index of changes in the direction of the airstream as the insect made attempts to turn away from a source of ultrasonic pulses.
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