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R R Hoy

Publications and source records attributed to R R Hoy.

At least 55 records · Page 3Linked to original sources
PubMed1990

Evolutionary innovation in behavior and speciation: opportunities for behavioral neuroethology.

Populations of animals differentiate by speciation. How speciation takes place, for long a puzzle for evolutionary biologists, should be regarded as a research opportunity for neuroethologists and comparative behaviorists. It is now clear that behavior may play an important part in the process of speciation. The existence of sexually dimorphic anatomy and behavior in many animals has provided a rich subject for investigation by comparative neurobiologists and behaviorists, who emphasize analysis of proximate mechanisms that generate dimorphism in phenotype. However, sexual dimorphisms also figure prominently in the theory of sexual selection. Sexual selection is viewed as a primary behavioral mechanism in the process of speciation. Some examples of 'explosive' speciation are presented, and the putative role that sexual selection plays in these cases is reviewed. A consideration of any evolutionary selective process must include genetics, and I will briefly summarize studies that indicate that even complex behavioral systems such as sexual selection, which sociobiologists refer to as a 'strategy', can have a relatively simple genetic basis.

Animals↗
PubMed1990

Ultrasound-induced yaw movements in the flying Australian field cricket (Teleogryllus oceanicus).

An ultrasonic stimulus induced negative phonotactic steering in the yaw axis of tethered, flying Australian field crickets. The forewings, hindwings and twisting of the thorax generated the forces which induced the yaw turn. However, abdominal ruddering did not contribute to yaw turns. Each aspect of the yaw steering response depended upon the stimulus intensity. At higher ultrasonic intensities, the magnitude and average angular velocity increased while the latency of the yaw turn decreased. Each of these factors varied in a graded manner, revealing that this behavior is more complex than a simple reflex.

Animals↗
PubMed1989

Audition in the praying mantis, Mantis religiosa L.: identification of an interneuron mediating ultrasonic hearing.

1. The praying mantis possesses a single ear located in the ventral midline of the metathorax. We have studied the mantis' auditory nervous system using both extracellular and intracellular techniques and have identified anatomically and physiologically a mirror-image pair of interneurons (MR-501-T3) in the metathoracic ganglion that mediates ultrasonic hearing. 2. MR-501-T3 is tuned broadly to ultrasound with best sensitivity (55-60 dB SPL) between 25 and 45 kHz. Its tuning matches closely that of the whole tympanal nerve. 3. The physiological responses of MR-501-T3 are characterized by: (1) a phasic-tonic firing pattern with a distinctive initial burst at 500-800 spikes/s; (2) minimum latencies of 8-12 ms; (3) no spontaneous activity; (4) sigmoid intensity response curves with a small (10 dB) dynamic range; (5) accurate coding of stimulus duration and of repetition rates up to 60 pps. 4. The ascending axon of MR-501-T3 conducts action potentials at 4 m/s, a rate comparable with some giant fiber systems. 5. MR-501-T3 shows no directional capability. Sound from right and left produce identical responses in both cells of the pair. Neither cutting one tympanal nerve nor removing one hemi-ear leads to different responses in the two cells indicating that they must receive a common input, either from the auditory afferents or from interneurons. We present evidence that the two cells are not directly connected. 6. MR-501-T3 is a large, symmetrical cell with its processes primarily in the intermediate neuropil (lateral ring tract). Its integration segment crosses the midline in the supramedian commissure, and the cell body lies dorsally near the entrance of the leg nerve. The axon travels in the dorsal lateral tract and is one of the largest (17 microns) in the connective. 7. Given the strong anatomical similarities between MR-501-T3 and the G and B cells of the locust, these cells may be homologous. 8. We present arguments based on our physiological results and existing behavioral data that MR-501-T3 is part of an ultrasonic warning/escape system in the mantis. As in moths, lacewings, and crickets, this system may provide a defense against nocturnally foraging bats.

Acoustic Stimulation↗
PubMed1989

Integration of ultrasound and flight inputs on descending neurons in the cricket brain.

In response to ultrasonic stimuli, tethered flying crickets perform evasive steering movements that are directed away from the sound source (negative phonotaxis). In this study we have investigated the responsiveness to ultrasound of neurons that descend from the cricket brain, and whether flight activity facilitates the responsiveness of these neurons. 1. Ultrasonic stimuli evoke descending activity in the cervical connectives both ipsilateral and contralateral to the sound source. 2. Both the amount of descending activity and the latency of this response in the cervical connectives are linearly correlated with ultrasonic stimulus intensity, regardless of the cricket's behavioral state. 3. Flight activity significantly increases the amount of descending activity evoked by ultrasound at all stimulus intensities, and significantly decreases the latency of the response in the cervical connectives compared with non-flying crickets. Flight activity, however, does not significantly affect the activity in an interneuron (Int-1) carrying ultrasound input to the brain. Thus, the increase in the amount of descending activity produced during flight activity is due to the integration of input from Int-1 and the flight motor system to ultrasound-sensitive neurons in the cricket brain. 4. Descending units recorded in the cervical connectives originate in the cricket brain. A reduction in the amount of descending activity is correlated with a decrease in the magnitude of the negative phonotactic response of the abdomen during flight activity, suggesting that these descending units play a role in eliciting negative phonotaxis.

Acoustic Stimulation↗
PubMed1988

Hawaiian courtship songs: evolutionary innovation in communication signals of Drosophila.

In Hawaii, flies of the genus Drosophila have undergone spectacular adaptive radiation, resulting in the evolution of more than 500 species of Drosophila that are found nowhere else on earth. This taxonomic uniqueness is reflected in behavior and morphology. Hawaiian Drosophila sing songs, as do continental Drosophila; however, the Hawaiian songs have diverged strongly in form and mechanism of production. The click-song of D. fasciculisetae's (Maui) has a carrier frequency an order of magnitude higher than those reported in familiar continental species, such as D. melanogaster (170 hertz). Drosophila fasciculisetae's song resembles a cicada's more than a fly's song. The song of D. cyrtoloma (Maui) has a complex pulse rhythm more typical of crickets than flies. The pulse song of D. silvestris (Hawaii) closely resembles that of D. melanogaster in both pulse rhythm and carrier frequency, but D. melanogaster sings by vibrating its wings, whereas D. silvestris sings through abdominal vibrations. These mechanisms are radical departures from the continental wing song mechanism and are further examples of the remarkable behavioral innovation that has occurred in the Drosophila of Hawaii during their evolutionary transit through these islands.

Animal Communication↗
PubMed1988

Effect of auditory deafferentation on the synaptic connectivity of a pair of identified interneurons in adult field crickets.

In adult crickets, Teleogryllus oceanicus, unilateral auditory deafferentation causes the medial dendrites of an afferent-deprived, identified auditory interneuron (Int-1) in the prothoracic ganglion to sprout and form new functional connections in the contralateral auditory neuropil. The establishment of these new functional connections by the deafferented Int-1, however, does not appear to affect the physiological responses of Int-1's homolog on the intact side of the prothoracic ganglion which also innervates this auditory neuropil. Thus it appears that the sprouting dendrites of the deafferented Int-1 are not functionally competing with those of the intact Int-1 for synaptic connections in the remaining auditory neuropil following unilateral deafferentation in adult crickets. Moreover, we demonstrate that auditory function is restored to the afferent-deprived Int-1 within 4-6 days following deafferentation, when few branches of Int-1's medial dendrites can be seen to have sprouted. The strength of the physiological responses and extent of dendritic sprouting in the deafferented Int-1 progressively increase with time following deafferentation. By 28 days following deafferentation, most of the normal physiological responses of Int-1 to auditory stimuli have been restored in the deafferented Int-1, and the medial dendrites of the deafferented Int-1 have clearly sprouted and grown across into the contralateral auditory afferent field. The strength of the physiological responses of the deafferented Int-1 to auditory stimuli and extent of dendritic sprouting in the deafferented Int-1 are greater in crickets deafferented as juveniles than as adults. Thus, neuronal plasticity persists in Int-1 following sensory deprivation from the earliest juvenile stages through adulthood.

Acoustic Stimulation↗
PubMed1988

Kinematic and aerodynamic aspects of ultrasound-induced negative phonotaxis inflying Australian field crickets (Teleogryllus oceanicus).

Negative phonotaxis is elicited in flying Australian field crickets, Teleogryllus oceanicus, by ultrasonic stimuli. Using upright tethered flying crickets, we quantitatively examined several kinematic and aerodynamic factors which accompany ultrasound-induced negative phonotactic behavior. These factors included three kinematic effects (hindwing wingbeat frequency, hindwing elevation and depression, and forewing tilt) and two aerodynamic effects (pitch and roll). 1. Within two cycles following a 20 dB suprathreshold ultrasonic stimulus, the hindwing wingbeat frequency increases by 3-4 Hz and outlasts the duration of the stimulus. Moreover, the relationship between the maximum increase in wingbeat frequency and stimulus intensity is a two-stage response. At lower suprathreshold intensities the maximum wingbeat frequency increases by approximately 1 Hz; but, at higher intensities, the maximum increase is 3-4 Hz. 2. The maximum hindwing elevation angle increases on the side ipsilateral to the stimulus, while there was no change in upstroke elevation on the side contralateral to the stimulus. 3. An ultrasonic stimulus affects forewing tilt such that the forewings bank into the turn. The forewing ipsilateral to the stimulus tilts upward while the contralateral forewing tilts downward. Both the ipsilateral and contralateral forewing tilt change linearly with stimulus intensity. 4. Flying crickets pitch downward when presented with a laterally located ultrasonic stimulus. Amputation experiments indicate that both the fore and hindwings contribute to changes in pitch but the pitch response in an intact cricket exceeds the simple addition of fore and hindwing contributions. With the speaker placed above or below the flying cricket, the change is downward or upward, respectively.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1987

The midline metathoracic ear of the praying mantis, Mantis religiosa.

The praying mantis, Mantis religiosa, is unique in possessing a single, tympanal auditory organ located in the ventral midline of its body between the metathoracic coxae. The ear is in a deep groove and consists of two tympana facing each other and backed by large air sacs. Neural transduction takes place in a structure at the anterior end of the groove. This tympanal organ contains 32 chordotonal sensilla organized into three groups, two of which are 180 degrees out of line with the one attaching directly to the tympanum. Innervation is provided by Nerve root 7 from the metathoracic ganglion. Cobalt backfills show that the auditory neuropile is a series of finger-like projections terminating ipsilaterally near the midline, primarily near DC III and SMC. The auditory neuropile thus differs from the pattern common to all other insects previously studied.

Animals↗
PubMed1987

Postsynaptic inhibition mediates high-frequency selectivity in the cricket Teleogryllus oceanicus: implications for flight phonotaxis behavior.

The frequency selectivity of the identified auditory interneuron, Int-1, in the cricket Teleogryllus oceanicus was examined using intracellular recording and staining techniques. Previous behavioral assays showed that crickets discriminate the low frequencies of the species calling song (4-5 kHz) from the high frequencies contained in the vocalizations of insectivorous bats (Nolen and Hoy, 1986a). Int-1 was excited by frequencies between 3 and 40 kHz, being similar, therefore, to the tympal organ (ear) in its broad range sensitivity; however, it responded differentially to high and low frequencies in terms of the number of action potentials evoked per stimulus tone pulse, the average discharge rate, and the latency of response. It was especially responsive to ultrasound (greater than 20 kHz), discharging at rates up to 400 spikes/sec (average rate), with 10 msec latencies; its response to pulses of the calling song was less than 150 spikes/sec, with 30 msec latencies. Int-1's dynamic range for ultrasound was also quite large, about 50 dB, compared to 20 dB for the calling song frequency. In addition, it responded well to trains of short, batlike pulses of ultrasound. These results are consistent with previous behavioral experiments showing that during flight, Int-1 was both necessary and sufficient for the ultrasound avoidance steering behavior (Nolen and Hoy, 1984), as long as it discharged above a rate of 180 spikes/sec. Ultrasound readily produced such high rates, whereas calling song rarely did; ultrasound reliably evoked avoidance steering over a wide dynamic range, while tone pulses of the calling song rarely did so (Nolen and Hoy, 1986a). A unique source of ipsilaterally mediated inhibition, tuned to the calling song frequency, accounted for the poor response to calling song and hence the neuron's high-frequency selectivity, and the behavioral and physiological effects of 2-tone suppression of high frequencies by the calling song (Nolen and Hoy, 1986b). These results further strengthen Int-1's proposed role as a "bat-detector" during flight and suggest only a limited role in other contexts such as social behavior.

Action Potentials↗
PubMed1986

Regeneration of normal afferent input does not eliminate aberrant synaptic connections of an identified auditory interneuron in the cricket, Teleogryllus oceanicus.

In the cricket, Teleogryllus oceanicus, the dendritic arborizations of an identified auditory interneuron (Int-1) are normally restricted to the ipsilateral auditory neuropile; unilateral deafferentation causes the medial portion of the dendritic field to sprout across the midline and make functional connections with the contralateral auditory neuropile (Hoy et al., '78: Soc. Neurosci. Abstr. 4:115, '85: Proc. Natl. Acad. Sci. USA 82:7772-7786; Hoy and Moiseff, '79: Soc. Neurosci. Abstr. 5:163). We have found that regeneration of the auditory afferents also results in an aberrant pattern of innervation of Int-1. Crickets were unilaterally deafferented during postembryonic development by crushing or cutting the auditory nerve. Regeneration of afferent-to-Int-1 connections was tested behaviorally. Of 86 nerve-crushed crickets tested as adults in the behavioral assay, 66% showed functional regeneration of the afferents. Similar results were obtained from the nerve-cut group. However, morphological investigations demonstrated that most of the regenerates still retained the aberrant contralateral dendritic projection. Electrophysiological recordings from these Int-1s showed that not only are some of them driven by their regenerated auditory afferents (the normal pathway) but that they retain their excitability via their contralateral dendrites (the aberrant pathway). This demonstrates that reinnervation of Int-1 by its normal afferent pool neither causes retraction nor prevents the formation of connections made with foreign, contralateral afferents. When the contralateral afferent pool was removed after Int-1 had sprouted, the sprouts remained present, but preliminary results suggest that if the contralateral afferents are removed before Int-1 is deafferented, sprouts are not formed. The results are discussed in relation to the roles of competition and conservation of membrane area in controlling synapse formation.

Animals↗
PubMed1986

The cyclopean ear: a new sense for the praying mantis.

The praying mantis, thought to be deaf, possesses a sensitive and specialized acoustic sense. Neural recordings show that the auditory system responds primarily to ultrasound between 25 and 45 kilohertz with thresholds of 55 to 60 decibels. Other insects with auditory tympana possess paired, laterally placed ears; the mantis has only a single ear that is located in the ventral midline between the metathoracic legs. Some species of mantis abruptly and dramatically alter their flight path when stimulated with ultrasonic pulses, suggesting a behavioral response to insectivorous echo-locating bats.

Animals↗
PubMed1986

Phonotaxis in flying crickets. I. Attraction to the calling song and avoidance of bat-like ultrasound are discrete behaviors.

The steering responses of three species of field crickets, Teleogryllus oceanicus, T. commodus, and Gryllus bimaculatus, were characterized during tethered flight using single tone-pulses (rather than model calling song) presented at carrier frequencies from 3-100 kHz. This range of frequencies encompasses the natural songs of crickets (4-20 kHz, Fig. 1) as well as the echolocation cries of insectivorous bats (12-100 kHz). The single-pulse stimulus paradigm was necessary to assess the aversive nature of high carrier frequencies without introducing complications due to the attractive properties of repeated pulse stimuli such as model calling songs. Unlike the natural calling song, single tone-pulses were not attractive and did not elicit positive phonotactic steering even when presented at the calling song carrier frequency (Figs. 2, 3, and 9). In addition to temporal pattern, phonotactic steering was sensitive to carrier frequency as well as sound intensity. Three discrete flight steering behaviors positive phonotaxis, negative phonotaxis and evasion, were elicited by appropriate combinations of frequency, temporal pattern and sound intensity (Fig. 12). Positive phonotactic steering required a model calling song temporal pattern, was tuned to 5 kHz and was restricted to frequencies below 9 kHz. Negative phonotactic steering, similar to the 'early warning' bat-avoidance behavior of moths, was produced by low intensity (55 dB SPL) tone-pulses at frequencies between 12 and 100 kHz (Figs. 2, 3, and 9). In contrast to model calling song, single tone-pulses of high intensity 5-10 kHz elicited negative phonotactic steering; low intensity ultrasound (20-100 kHz) produced only negative phonotactic steering, regardless of pulse repetition pattern. 'Evasive', side-to-side steering, similar to the 'last-chance' bat-evasion behavior of moths was produced in response to high intensity (greater than 90 dB) ultrasound (20-100 kHz). Since the demonstration of negative phonotactic steering did not require the use of a calling song temporal pattern, avoidance of ultrasound cannot be the result of systematic errors in localizing an inherently attractive stimulus when presented at high carrier frequencies. Unlike attraction to model calling song, the ultrasound-mediated steering responses were of short latency (25-35 ms) and were produced in an open loop manner (Fig. 4), both properties of escape behaviors.(ABSTRACT TRUNCATED AT 400 WORDS)

Animal Communication↗
PubMed1986

Phonotaxis in flying crickets. II. Physiological mechanisms of two-tone suppression of the high frequency avoidance steering behavior by the calling song.

The effects of two-tone stimuli on the high frequency bat-avoidance steering behavior of flying crickets (Teleogryllus oceanicus) were studied during tethered flight. Similarly, the effects of two-tone stimuli on the ultrasound sensitive auditory interneuron, Int-1, which elicits this behavior, were studied using intracellular staining and recording techniques. When a low frequency tone (3-8 kHz) was presented simultaneously with an aversive high frequency tone (in a two-tone stimulus paradigm), the high frequency avoidance steering behavior was suppressed. Suppression was optimal when the low frequency tone was between 4 and 5 kHz and about 10-15 dB louder than the high frequency tone (Figs. 2, 3). Best suppression occurred when the low frequency tone-pulse just preceded or overlapped the high frequency tone-pulse, indicating that the suppressive effects of 5 kHz could last for up to 70 ms (Fig. 4). The threshold for avoidance of the bat-like stimulus was elevated when model bat biosonar (30 kHz) was presented while the animal was performing positive phonotaxis toward 5 kHz model calling song, but only if the calling song intensity was relatively high (greater than 70-80 dB SPL) (Fig. 1). However, avoidance steering could always be elicited as long as the calling song was not more than 10 dB louder than the ultrasound (Fig. 1). This suppressive effect did not require performance of positive phonotaxis to the calling song (Fig. 2) and was probably due to the persistence of the suppressive effects of the 5 kHz model calling song (Fig. 4). The requirement for relatively high intensities of calling song suggest that the suppression of bat-avoidance by the calling song is not likely to be of great significance in nature. The high frequency harmonics of the male cricket's natural calling song overlap the lower frequency range used by insectivorous bats (10-20 kHz) and are loud enough to elicit avoidance behavior in a flying female as she closely approaches a singing male (Fig. 5). The high frequency 'harmonics' of a model calling song were aversive even if presented with a normally attractive temporal pattern (pulse repetition rate of 16 pps) (Fig. 6A). When the 5 kHz 'fundamental' was added to one of the high frequency 'harmonics', in a two-tone stimulus paradigm, this complex model calling song was attractive; the high frequency 'harmonic' no longer elicited the avoidance behavior (Fig. 6) and the animals steered toward the model CS. Thus, addition of 5 kHz to a high frequency harmonic of the calling song 'masked' the aversive nature of this stimulus.(ABSTRACT TRUNCATED AT 400 WORDS)

Animal Communication↗
PubMed1985

Dendritic sprouting and compensatory synaptogenesis in an identified interneuron follow auditory deprivation in a cricket.

We examined the effect of chronic afferent deprivation on an identified interneuron (Int-1) in the auditory system of the Australian field cricket Teleogryllus oceanicus. In normal intact crickets, the auditory afferents from each ear terminate ipsilaterally onto a single Int-1. Each bilaterally paired Int-1 is excited by ultrasound stimulation of its ipsilateral ear but not by the contralateral ear. Unilateral removal of an ear early in postembryonic development deprives the developing Int-1 of ipsilateral auditory innervation. Consequently, the ipsilateral dendrites of the deprived interneuron sprout, grow aberrantly across the ganglionic midline, and terminate specifically in the intact auditory neuropile of the contralateral (unlesioned) side, where they form functional synapses with the contralateral afferents. This unusual compensatory dendritic sprouting restores auditory function to the neuron. Thus, it is demonstrated that the dendritic shape of an identified Int, as well as its synaptic connectivity, is altered as a consequence of chronic sensory deprivation.

Afferent Pathways↗
PubMed1984

Initiation of behavior by single neurons: the role of behavioral context.

Flying crickets avoid sources of ultrasound, possibly echolocating bats, by making rapid steering movements that turn them away from the stimulus. Electrical stimulation of a single, identified sensory interneuron (Int-1) elicits avoidance steering; depressing its response to ultrasound abolishes avoidance steering. Int-1 is necessary and sufficient for this behavior but only while the cricket is in flight. Thus, the sufficiency of Int-1 for eliciting this behavior is contingent on behavioral context.

Acoustic Stimulation↗
PubMed1978

Steering responses of flying crickets to sound and ultrasound: Mate attraction and predator avoidance.

We investigated the steering responses of tethered, flying adult female crickets (Teleogryllus oceanicus) to acoustic stimulation. Crickets responded directionally to directional sound stimulation by bending their abdomens and hind legs to one side. We interpret this response as an attempt to turn. When stimulated with a model of conspecific calling song with a carrier frequency of from 3 to 9 kHz, crickets turned toward the sound source. We believe that this indicates a positive phonotactic response of flying females to calling, conspecific males. When offered a choice between conspecific song and the song of another species, females turned exclusively toward conspecific song, demonstrating that the response is species specific. The direction of the response is dependent on the carrier frequency of the song, and it demonstrates frequency discrimination. Females turned toward calling song when it was played at carrier frequencies from 3 to 9 kHz, but they turned away from the same song pattern played at carrier frequencies from 30 to 70 kHz. This negative phonotaxis to ultrasonic stimuli suggests that crickets, like some other nocturnal flying insects, can evade bats by acoustic detection.

Journal Article↗