Wing-flick Signals in the Courtship of the African Cave Cricket,Phaeophilacris spectrum

The male of the African cave cricket Phaeophilacris spectrum (Saltatoria: Grylloidea: Phalangopsidae) possesses tegmina without stridulatory organs, and both females and males lack tympanal organs. Therefore acoustic communication in the usual sense, which is typical for most crickets, is absent in this species. However, adaptations of the wing articulation allow the males to flick their wings forward over their heads. During courtship these movements are performed in series of 4–5 wing-flicks at a rate of 8–12 Hz (called wing-flick series = WFS) which elicit low-frequency air movements. Such signals may have taken over the functional significance of an acoustic ‘courtship song’. A quantitative analysis of the cricket's behaviour showed that in terms of frequency and total duration, ‘wing-flick series’ and ‘rocking’ behaviour are the most prominent signals of the male's courtship display. Moreover, analysis of the spatial relationships between male and female demonstrates that the wing-flicking is directed towards the female. The response of a female to a wing-flicking, courting male seems mainly to be calmed down and to reach a passive, receptive state, necessary for subsequent copulation. Females confronted with a male's courtship display exhibited a significantly higher threshold to disturbing stimuli than uncourted ones.     

Frequency as a releaser in the courtship song of two crickets,Gryllus bimaculatus (de Geer) and Teleogryllus oceanicus: a neuroethological analysis

1. The courtship behavior of male field crickets, Gryllus bimaculatus (De Geer) and Teleogryllus oceanicus, is a complex, multimodal behavioral act that involves acoustic signals (a courtship song; Fig. 1A,B). The dominant frequency is 4.5 kHz for T. oceanicus song (Fig. 1A) and 13.5 kHz for G. bimaculatus (Fig. IB).
2. When courting males are deprived of their courtship song by wing amputation, their courtship success declines markedly but is restored when courting is accompanied by tape-recordings of their courtship songs or a synthetic courtship song with only the dominant frequency of the natural song; other naturally occurring frequency components are ineffective for restoring mating success (Figs. 4, 5).
3. It has been suggested that an identified auditory interneuron, AN2, plays a critical role in courtship success. Chronic recordings of AN2 in an intact, tethered female show that AN2's response to the natural courtship song and synthesized songs at 4.5 and 13.5 kHz is similar in T. oceanicus. By contrast, in G. bimaculatus, AN2's response to the natural courtship song and synthesized song at 13.5 kHz, but not at 4.5 kHz, is similar (Figs. 2,3).
4. In behavioral experiments, playback of a 30 kHz synthetic courtship song in G. bimaculatus does not restore courtship success, yet this same stimulus elicits as strong a response from AN2 as does the normal courtship song (Fig. 6). Thus, contrary to earlier work by others, we conclude AN2 is not, by itself, a critical neural link in the courtship behavior of these two species of crickets.

     

Digger wasp against crickets. II. An airborne signal produced by a running predator

Summary Females of the digger wasp Liris niger Fabr. hunt crickets to provide food for their offspring by running with high velocity on the ground (>20–50 cm/s). Crickets are able to detect the running wasps by the air particle movement generated by the predator. We measured signals produced by running wasps using a microphone sensitive to air particle velocity. The wasps generated single air puffs with peak air particle velocities of 1–2 cm/s measured close to the running wasp. We measured frequency spectra of the signals containing only components below 50 Hz, with increasing intensities towards lower frequencies, especially below 10 Hz.We measured the air particle movement generated by artificially moved wasps, crickets or a styrofoam dummy of similar size to investigate the effect of velocity and shape of the moving object upon the composition of the signal. The velocity of movement appeared to be important for the intensity and frequency composition of the air particle movement. The shape of the moved body had an influence on the intensity but only little effect on the frequency spectrum. Measurements with a thermistor anemometer showed that a moving object caused air currents lasting longer than 100 ms after passing or approaching the probe. The air particle movements generated by hunting wasps are entirely sufficient with respect to intensity and frequency range to be registered by the filiform hair sensilla upon the cerci of crickets.     

Signals for Survival in the Lives of Crickets

Two behavioral acts of undoubted survival value are predatoravoidance and mate choice. In field crickets both are mediatedby acoustic signals containing high frequency spectral energy.Nocturnally active bats use ultrasonic echolocation signalsto detect and locate their prey, which includes insects thatdisperse by flying at night. Many insects have developed ultrasoundavoidance behaviors in flight, in order to elude bats. In crickets,an auditory interneuron that is excited by ultrasound has beenidentified and shown to initiate the avoidance behavior; itis a putative bat-detector cell. Male field crickets produceacoustic signals during their courtship (females are mute).Courtship song appears to facilitate mating success (copulation),for its absence in courtship diminishes the likelihood of copulation.The possible role of the bat-detector neuron in courtship behavioris considered because it is activated by courtship signals aswell as bat-like ultrasound. The role of behavioral contextshapes the participation of neurons in the neural networks thatunderlie a given behavior.     

NO-synthase inhibitor prevents flight-induced activation of agressiveness and courtship in male

Previous experience of flying enhances the aggressiveness (Hofmann and Stevenson, 2000, Nature, 403: 613) and accelerates the courtship behaviour (Dyakonova and Krushinski, 2003, DAN, 390: 709-712) of crickets Gryllus bimacultus. We present evidence that these effects may be mediated by activation of nitric oxide synthesis. The effects of flying on fighting and courtship were largely abolished in crickets who received haemocoel injections of a nonspecific NO-synthase inhibitor LNNA. Unlike this, LNNA exerted no significant effects on aggressive and courtship behaviour of nonflown males.     

Air-movement `signals' are not required for female mounting during courtship in the cricket Teleogryllus oceanicus

Wing movements associated with stridulation by the male during cricket courtship generate air movements that have been proposed to serve as signals to the female. We assessed this putative signaling role by interfering with the presumed communications channel in two ways: (1) by removing the female's cerci, which are candidate sensory organs for signal reception, and (2) by trimming the male's forewings and thus manipulating the signal itself. We measured the effects of these treatments on the probability and latency with which females mounted courting males. We found that neither treatment affected female mounting behavior. This was true both for old, highly motivated females and for younger females, which are less highly motivated and possibly more selective. We conclude that air movements play little or no role as signals that release female mounting behavior during courtship. Accepted: 11 June 1998     

Removal of both antennae influences the courtship and aggressive behaviors in male crickets

To test whether insect antennae are necessary for eliciting courtship and aggression toward appropriate partners, we antennectomized adult male crickets (Gryllus bimaculatus) and observed their behavior toward other antennectomized males and intact females. At 7 days after removal of both antennae, pairs of antennectomized males were placed together; 80% displayed courtship behavior, generating courtship song by rubbing their forewings together, toward other antennectomized males, and 20% displayed aggressive behavior. Only 45% courted intact females. No intact males courted antennectomized males, and 80% displayed aggressive behavior. All intact males courted females. The results for males with one antenna removed were essentially the same as for intact males. These findings indicate that a high proportion of male crickets with both antennae removed court other males and fail to display male-male aggression, demonstrating that removal of antennae from male crickets induces male-male courtship and that an antenna is necessary for the expression of male-male aggression. Moreover, brain serotonin (5-hydroxytryptamine; 5-HT) levels in male crickets were significantly reduced at 7 days after removal of antennae. The reduction of 5-HT was detected primarily in the central body of the brain. Thus, 5-HT in the central body of the male cricket brain may be involved in the behavioral changes.     

Acoustic communication and sexual selection in Orthoptera (Insecta)

Comparison of calling and courtship songs and mating strategies in different groups of Orthoptera shows that acoustic signals in bush crickets (Tettigonioidea) are used for searching conspecific mates at a distance, and song evolution is primarily driven by the acoustic surroundings and simultaneously singing other species. The role of sexual selection in the evolution of acoustic signals in bush crickets is much less important than in the evolution of other nonacoustic signals employed during direct contact. Acoustic signals of crickets (Grylloidea) are involved in both distant and short-range communication. Acoustic courtship signals, along with other courtship components, may offer the material for sexual selection, although, essentially, evolution of acoustic signals in crickets is determined by the acoustic surroundings. Acoustic communication in the grasshoppers of the subfamily Gomphocerinae is mainly a short-distance communication. Acoustic signals of Gomphocerinae are highly variable and elaborate in temporal parameters, and therefore can serve for evaluation of the mate "quality". Song evolution in this group is to a greater extent driven by sexual selection than by the acoustic surroundings and, therefore, could proceed faster than in other groups of Orthoptera and play a major role in speciation.     

Basic elements of behavior of the cricket <Emphasis Type="Italic">Phaeophilacris bredoides</Emphasis> Kaltenbach (Orthoptera,Gryllidae)

The intraspecific behavior of the non-singing cricket Phaeophilacris bredoides Kaltenbach, 1986, which has no tympanal system, stridulatory apparatus, and classical acoustic communication, was studied. Even though this cricket has no song, its intraspecific behavior can be differentiated into reproductive and agonistic (defensive and aggressive), as this was done before for singing crickets. The main elements and the sequence of the phases were described for reproductive behavior. The active role during copulation belongs to the male. Wingflicks and rocking movements of the male can function as a “song.” Wing-flicks apparently generate air movements that function as short-range signals during reproductive and aggressive behavior. Substrate-borne vibrations produced by rocking also seem to be associated with aggressive behavior. Antennal contacts form an important part of interaction between crickets of both sexes. Thus, intraspecific signaling is at least partly mediated by mechanosensory channels. The assumption about the possible direction of evolution in the singing and non-singing groups of crickets was made.     

No Effect of Body Size on the Frequency of Calling and Courtship Song in the Two-Spotted Cricket,Gryllus bimaculatus

The relationship between body size and vocalization parameters has been studied in many animal species. In insect species, however, the effect of body size on song frequency has remained unclear. Here we analyzed the effect of body size on the frequency spectra of mating songs produced by the two-spotted cricket, Gryllus bimaculatus. We recorded the calling songs and courtship songs of male crickets of different body sizes. The calling songs contained a frequency component that peaked at 5.7 kHz. On the other hand, courtship songs contained two frequency components that peaked at 5.8 and 14.7 kHz. The dominant frequency of each component in both the calling and courtship songs was constant regardless of body size. The size of the harp and mirror regions in the cricket forewings, which are the acoustic sources of the songs, correlated positively with body size. These findings suggest that the frequency contents of both the calling and courtship songs of the cricket are unaffected by whole body, harp, or mirror size.     

Tonic and modulatory subsystems of the complex gravity receptor system in crickets, Gryllus bimaculatus

The gravity receptor system of crickets Gryllus bimaculatus is composed of antennal, cercal and leg subsystems. The cercal gravity receptors are the club-shaped sensilla. Each of these subsystems elicits compensatory head movements during passive roll.The extent of compensatory head movements depends on the strength of the gravitational stimulus applied to the leg subsystem. Amputation of 2 legs never causes a decrease in reflex amplitude. Unilateral amputation of 1 to 3 legs always induces a roll movement of the head to the intact body side. Therefore, the leg gravity receptor system exerts a modulatory and tonic effect on the neck muscles.The gravity receptors of 1 cercus or 1 antenna only elicit compensatory head movements. They exert no tonic effect on the neck muscles.The results are discussed with respect to (i) the proposed connectivity of the cercus-neck muscle pathway, (ii) mutual inhibitory interactions between the sensory pathways originating in the leg gravity receptors, and (iii) the influence of non-gravitationally induced excitation on the occurrence of compensatory head movements during passive roll of the crickets.     

The role of chemoreception in sex recognition by male crickets: Acheta domesticus and Teleogryllus oceanicus

ABSTRACT. Contact chemoreception is important in female recognition by Teleogryllus oceanicus (Le Guillou) males. Antennal contact of female conspecifics, body regions, detached antennae and conditioned substrate elicited mostly courtship responses including courtship songs. Aggressive acts were produced only in response to male conspecifics. Male body regions, detached antennae and conditioned substrate elicited very few courtship or aggressive acts and no songs. This suggests that one or several communication modes, in addition to chemical communication, are necessary to elicit aggressive responses. Acheta domesticus (L.) males cannot rely upon chemical cues for recognition of either sex. Responses to conspecifics suggest that A. domesticus males produce aggressive acts immediately after antennal contact with either sex. Aggressive response to males usually persists, but response to females often switches to courtship. Responses to body regions, detached antennae, and conditioned substrate were few, with courtship and aggressive responses elicited by both male- and female-generated stimuli. The importance of contact chemoreception in cricket communication is suggested by (1) failure of hexane-washed antennae to elicit aggressive or courtship acts, and (2) males spending more time in contact with body regions and conditioned substrates than with corresponding controls. Lack of response to male or female odour-laden air suggests that chemical signals are used by males only if directly contacted. Chemical and other signals supplement the obvious use of acoustic signals for intra- and intersexual communication in these crickets. The importance of multimodal communication in sex recognition is discussed.     

Courtship song and immune function in the field cricket Gryllus bimaculatus

It has been assumed that sexual ornaments have evolved to reveal males' health and vigour for females. Choosy females may indirectly use ornaments as an indicator of the presence and effectiveness of genes for resistance against parasites. In this study we tested whether females of the Mediterranean field cricket, Gryllus bimaculatus, can use courtship song as a cue for choosing males with high immunocompetence, measured as encapsulation rate of nylon implants and lytic activity of haemolymph. We found that female crickets preferred courtship songs from males with a high encapsulation rate. Female crickets also had a tendency to prefer courtship songs with high tick rate and long high-frequency tick duration. These preferred song components were positively correlated with encapsulation rate, but negatively correlated with lytic activity of the male. In contrast to previous studies of crickets, there was no correlation between male weight and encapsulation rate or lytic activity. There is some evidence in another cricket species that the ability to encapsulate pathogens is heritable. Thus, in light of this study it seems possible that by preferring males according to their courtship song, females might benefit by increasing the parasite resistance of their offspring.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 503–510.     

The courtship behaviour of the pipefish Nerophis lumbriciformis: reflections of an adaptation to intertidal life

The courtship behaviour of Nerophis lumbriciformis (Pisces: Syngnathidae) consists of three distinct phases (initial courtship, spawning, and embrace) marked by prominent behavioural changes. The first courtship phase is characterised by female quivering. Courtship activity increases from low to high levels of quivering, causing undulatory head movements in the female. In the second phase, the female transfers her eggs onto the male incubating ventral surface. During the final phase, the male wraps his body around the female (embrace). Females actively initiate courtship indicating that these pipefishes are courtship role reversed. The complete lack of swimming and vertical movements in the courtship behaviour of N. lumbriciformis, unlike in the behaviour of other syngnathid species, suggests an adaptation to intertidal conditions.     

Neurohormonal control of the mating interval in the male cricket,Gryllus bimaculatus DeGeer

Summary Between two mating acts of the male cricket (Gryllus bimaculatus), spermatophore protrusion (SP) and courtship stridulation (CS), there is a fixed time interval. This interval lasts about 1 h. During the period from SP to CS, the male cricket does not stridulate nor make any type of mating sound (post-spermatophore protrusion silence: PSPS) and tolerates external sensory stimuli. We examined the effects of injections of hemolymph and ganglia extracts on the interval. Extracts obtained from crickets which had just started CS (CS crickets) and those which had finished SP (SP crickets) were effective. The extracts were fractionated by ul trafiltration. Fractions with a molecular weight of less than 1 kdalton affected the length of the PSPS. The fractions from both the hemolymph and the mesothoracic ganglion of CS crickets shortened the PSPS. On the other hand, the fractions from the hemolymph and the brain of SP crickets lengthened the PSPS. We estimated, by gel filtration, the molecular weight of the effective fractions from the mesothoracic ganglion and the brain to be 100–200 daltons. We also examined the effects of biogenic amines on the PSPS. Octopamine shortened the PSPS, whereas serotonin lengthened it. The results in dicate that at least two neurohormones from the brain and the mesothoracic ganglion reciprocally control the elicitation of CS and provide an appropriate interval in the mating sequence of the male cricket. Octopamine and serotonin are possible candidates for these neurohormones.Abbreviations CP copulation - CS courtship stridulation - SP spermatophore protrusion - PSPS post-spermatophore protrusion silence     

Force Control Is Related to Low-Frequency Oscillations in Force and Surface EMG

Force variability during constant force tasks is directly related to oscillations below 0.5 Hz in force. However, it is unknown whether such oscillations exist in muscle activity. The purpose of this paper, therefore, was to determine whether oscillations below 0.5 Hz in force are evident in the activation of muscle. Fourteen young adults (21.07±2.76 years, 7 women) performed constant isometric force tasks at 5% and 30% MVC by abducting the left index finger. We recorded the force output from the index finger and surface EMG from the first dorsal interosseous (FDI) muscle and quantified the following outcomes: 1) variability of force using the SD of force; 2) power spectrum of force below 2 Hz; 3) EMG bursts; 4) power spectrum of EMG bursts below 2 Hz; and 5) power spectrum of the interference EMG from 10–300 Hz. The SD of force increased significantly from 5 to 30% MVC and this increase was significantly related to the increase in force oscillations below 0.5 Hz (R ² = 0.82). For both force levels, the power spectrum for force and EMG burst was similar and contained most of the power from 0–0.5 Hz. Force and EMG burst oscillations below 0.5 Hz were highly coherent (coherence = 0.68). The increase in force oscillations below 0.5 Hz from 5 to 30% MVC was related to an increase in EMG burst oscillations below 0.5 Hz (R ² = 0.51). Finally, there was a strong association between the increase in EMG burst oscillations below 0.5 Hz and the interference EMG from 35–60 Hz (R ² = 0.95). In conclusion, this finding demonstrates that bursting of the EMG signal contains low-frequency oscillations below 0.5 Hz, which are associated with oscillations in force below 0.5 Hz.     

Acoustic signals in insects: A reproductive barrier and a taxonomic character

In singing insects, the song is an important component of the specific mate recognition system (SMRS). In communities of sympatric singing species, there is a partitioning of communication channels, the so-called “acoustic niches.” Within one community, the songs of different species always differ in temporal or frequency characters, i.e. occupy different acoustic niches. However, conspecific songs do not always act as an interspecific reproductive barrier, despite always being a SMRS component. The species that do not communicate acoustically due to allopatry, different timing of vocalization, inhabiting different biotopes, or unmatched food specializations can produce similar songs while forming reproductively isolated communities. Individuals of different sexes need not only to recognize a conspecific mate but also to evaluate its “quality.” The close-range signal (courtship song) provides more opportunities for choosing the “best” male than does the distant signal (calling song). In many species of Orthoptera, courtship includes not only acoustic but also vibrational, visual, chemical, and mechanical signals. An analysis of cricket songs showed the courtship songs to be on average more elaborate and variable than the calling songs. At the same time, due to the difference in mating behavior between the two groups, the acoustic component of courtship is used for mate quality evaluation to a greater extent in grasshoppers than in crickets. The courtship songs of grasshoppers are generally more elaborate in temporal structure than cricket songs; moreover, they may be accompanied by visual displays such as movements of various body parts. Thus, song evolution in grasshoppers is more strongly driven by sexual selection than that in crickets. According to the reinforcement hypothesis, the premating barrier between hybridizing species becomes stronger in response to reduced hybrid fitness. However, our behavioral experiments with two groups of hybridizing grasshopper species did not confirm the reinforcement hypothesis. We explain this, firstly, by a low level of genetic incompatibility between the hybridizing species and secondly, by high hybrid fitness when attracting a mate. A high competitive capability of hybrids may be accounted for by attractiveness of new elements in hybrid courtship songs. When we divide similar forms based on their songs, we in fact distinguish biological species using the criterion of their reproductive isolation. Acoustic differences between species are usually greater than morphological ones. Therefore, song analysis allows one to determine the real status of doubtful species-rank taxa, to distinguish species in a medley of sibling forms, and to reveal cryptic species in the cases when morphological studies fail to provide a univocal result. At the same time, songs are subject to intraspecific variation the range of which is different in different groups. Therefore, it is necessary to study which degree of difference corresponds to the species level before interpreting the status of some forms based on song comparisons. Besides, song similarities cannot indicate conspecificity of acoustically isolated forms; on the other hand, song differences between these forms prove that they are full-rank species.     

Modulations of EEG Beta Power during Planning and Execution of Grasping Movements

Although beta oscillations (≈ 13–35 Hz) are often considered as a sensorimotor rhythm, their functional role remains debated. In particular, the modulations of beta power during preparation and execution of complex movements in different contexts were barely investigated. Here, we analysed the beta oscillations recorded with electroencephalography (EEG) in a precued grasping task in which we manipulated two critical parameters: the grip type (precision vs. side grip) and the force (high vs. low force) required to pull an object along a horizontal axis. A cue was presented 3 s before a GO signal and provided full, partial or no information about the two movement parameters. We measured beta power over the centro-parietal areas during movement preparation and execution as well as during object hold. We explored the modulations of power in relation to the amount and type of prior information provided by the cue. We also investigated how beta power was affected by the grip and force parameters.We observed an increase in beta power around the cue onset followed by a decrease during movement preparation and execution. These modulations were followed by a transient power increase during object hold. This pattern of modulations did not differ between the 4 movement types (2 grips ×2 forces). However, the amount and type of prior information provided by the cue had a significant effect on the beta power during the preparatory delay. We discuss how these results fit with current hypotheses on the functional role of beta oscillations.     

Hair canopy of cricket sensory system tuned to predator signals

Filiform hairs located on the cerci of crickets are among the most sensitive sensors in the animal world and enable crickets to sense the faintest air movements generated by approaching predators. While the neurophysiological and biomechanical aspects of this sensory system have been studied independently for several decades, their integration into a coherent framework was wanting. In order to evaluate the hair canopy tuning to predator signals, we built a model of cercal population coding of oscillating air flows by the hundreds of hairs on the cerci of the sand cricket Gryllus bimaculatus (Insecta: Orthoptera). A complete survey of all hairs covering the cerci was done on intact cerci using scanning electronic microscopy. An additive population coding of sinusoid signals of varying frequencies and velocities taking into account hair directionality delivered the cercal canopy tuning curve. We show that the range of frequencies and velocities at which the cricket sensory system is best tuned corresponds to the values of signals produced by approaching predators. The relative frequencies of short (< 0.5 x 10(-3) m) and long hairs and their differing responses to oscillating air flows therefore enable crickets to detect predators in a time-frequency-intensity space both as far as possible and at close range.