Song pattern recognition and an auditory time window in the female bushcricket Ancistrura nigrovittata (Orthoptera: Phaneropteridae)

Mate finding in the phaneropterid bushcricket Ancistrura nigrovittata is achieved by a duet, where the female replies with a short sound to the male song. In experiments with artificial song models we analysed the parameters necessary for eliciting a female response. A verse of the male song consists of a group of 5–9 syllabes which after an interval of about 400 ms is followed by a final syllable. The female response was shown to depend on two processes: (i) recognition of the syllable group as belonging to a conspecific male and (ii) perception of the final syllable as a trigger. Critical parameters for the recognition process are the duration of syllables and syllable pauses, as well as the number of syllables in a group. However, even with an optimal syllable group, the response probability still depends on the interval between the syllable group and the final syllable. The female only responds when the final syllable of the male song occurs within a 250 ms long time window begining approximately 250 ms after the end of the male's syllable group. Her reply consists of a single tick, which follows the male's final syllable with a latency of only 25 ms.     

Responses and song pattern copying of Omega-type I-neurons in the cricket,Gryllus bimaculatus,at different prothoracic temperatures

Summary Omega-type I-neurons (ON/1) (Fig. 1A) were recorded intracellularly with the prothoracic ganglion kept at temperatures of either 8–9°, or 20–22° or 30–33 °C and the forelegs with the tympanal organs kept at ambient temperature (20–22 °C). The neurons were stimulated with synthetic calling songs (5 kHz carrier frequency) with syllable periods (SP in ms) varying between 20 and 100, presented at sound intensities between 40 and 80 dB SPL. The amplitude and duration of spikes as well as response latency decreased at higher temperatures (Figs. 1 B, 2, 6). At lower prothoracic temperatures (8–9 °C) the neuron's responses to songs with short SP (20 ms) failed to copy single syllables, or with moderate SP (40 ms) copied the syllable with low signal to noise ratio (Fig. 3). The auditory threshold of the ON/1 type neuron, when tested with the song model, was temperature-dependent. At 9° and 20 °C it was between 40 and 50 dB SPL and at 33 °C it was less than 40 dB SPL (Fig. 4). For each SP, the slope of the intensity-response function was positively correlated with temperature, however, at low prothoracic temperatures the slope was lower for songs with shorter SPs (Fig. 5). The poor copying of the syllabic structure of the songs with short SPs at low prothoracic temperatures finds a behavioral correlate because females when tested for phonotaxis on a walking compensator responded best to songs with longer SPs at a similar temperature.Abbreviations epsps excitatory postsynaptic potentials - ON/1 omega-type I-neuron - SP syllable period - SPL sound pressure level     

Song Learning in the Anna Hummingbird (Calypte anna)

Songs of wild male Anna hummingbirds (Calypte anna) consist of syllables grouped into phrases. Nearest neighbors tend to share similar syllable types, rhythms and syntax. Songs from different localities contain different syllable types, syntax and repetition indices. A male raised by hand in isolation produced a song consisting of highly variable syllable types of a wide frequency range. The song was simple in structure, and syllables were not grouped into phrases. Three males raised by hand as a group sang songs containing two stereotyped syllable types sung in alternating sequence and without phrase structure. These three males shared syllable types and syntax. The data from our study indicate that despite its relatively simple syrinx the Anna hummingbird learns syllable types, frequency, rhythm and syntax (as do oscines with their more complex syringes) during the song development process.     

Acoustic pattern recognition in a grasshopper: processing in the time or frequency domain?

Males of the grasshopper Chorthippus biguttulus produce songs which consist of the stereotyped and rhythmic iteration of a sound unit (termed syllable) separated by distinct syllable pauses. Virgin females respond to this signal, and to similar artificial signals, with song phrases of their own. In behavioural experiments the response probability of virgin females can be measured with artificial acoustic stimuli. The stimuli consisted of an amplitude modulated noise the envelope of which was altered. We investigated several hypotheses on the mechanisms of conspecific song recognition with special emphasis on the question whether recognition occurs in the frequency domain or in the time domain. (1) Females of Ch. biguttulus required only the first five Fourier components of the envelope function (corresponding to 50 Hz for a fundamental frequency of 10 Hz) to detect the syllable/pause structure. In addition, they detected small gaps within syllables if the signal contained at least ca. 15 Fourier components (corresponding to a frequency of 150 Hz). Further experiments showed that the correct phase information of the Fourier components is necessary for recognition, indicating that pattern recognition is not achieved merely on the basis of band pass filtering. (2) A cross correlation between the signal and an assumed internal template yields only inconsistent predictions of the response probabilities. (3) The recognizer system probably works in the time domain, possibly by direct comparison of adjacent syllable and pause durations. It is not yet clear whether the duration of a syllable is evaluated with respect to the preceding or succeeding pause. We emphasize that the neural recognizer of the grasshopper does not only examine a signal for its similarity to an internal template, but that it also takes into account features that indicate an incorrect signal. This may be a general feature of neuronal pattern recognition systems which have been shaped by natural selection. Received: 4 October 1997 / Accepted in revised form: 26 August 1998     

Temperature coupling in cricket acoustic communication

Summary Temperature effects on calling song production and recognition were investigated in the North American field cricket, Gryllus firmus. Temporal parameters of field-recorded G. firmus calling song are strongly affected by temperature. Chirp rate and syllable rate increase, by factors of 4 and 2, respectively, as linear functions of temperature over the range in which these animals sing in the field (12°–30 °C). Temperature affects syllable duration to a lesser extent, and does not influence calling song carrier frequency. Female phonotactic preference, measured on a spherical treadmill in the laboratory, also changes with temperature such that warmer females prefer songs with faster chirp and syllable rates. Best phonotaxis, measured as accuracy of orientation to the sound source, and highest walking velocity, occur in response to temperature-matched songs at 15°, 21°, and 30 °C. Experiments under semi-natural conditions in an outdoor arena revealed that females perform phonotaxis at temperatures as low as 13 °C. Taken together, the song and phonotaxis data demonstrate that this communication system is temperature coupled. A strategy is outlined by which temperature coupling may be exploited to test hypotheses about the organization of neural networks subserving song recognition.Abbreviations CP chirp period - SP syllable period - SD syllable duration     

Airflow and pressure during canary song: direct evidence for mini-breaths

Summary Male canaries (Serinus canaria) produce songs of long duration compared to the normal respiratory cycle. Each phrase in a song contains repetitions of a particular song syllable, with repetition rates for different syllables ranging from 3 to 35 notes/s. We measured tracheal airflow and air sac pressure in order to investigate respiratory dynamics during song.Song syllables (11–280 ms) are always accompanied by expiratory tracheal airflow. The silent intervals (15–90 ms) between successive syllables are accompanied by inspiration, except for a few phrases where airflow ceases instead of reversing. Thus, the mini-breath respiratory pattern is used most often by the five birds studied and pulsatile expiration is used only occasionally.Songs and phrases accompanied by minibreaths were of longer duration than those accompanied by pulsatile expiration, presumably because the animal's finite vital capacity is not a limiting factor when the volume of air expired for one note is replaced by inspiration prior to the next. Pulsatile expiration was used for only a few syllable types from one bird that were produced at higher repetition rates than syllables accompanied by mini-breaths. We suggest that male canaries switch to pulsatile expiration only when the syllable repetition rate is too high (greater than about 30 Hz) for them to achieve mini-breaths.Changes in syringeal configuration that may accompany song are discussed, based on the assumption that changes in the ratio of subsyringeal (air sac) pressure to tracheal flow rate reflect changes in syringeal resistance.     

Songs and the Function of Song Elements in Four Duetting Bushcricket Species (Ensifera, Phaneropteridae, Barbitistes)

The structure of male songs and the timing of female replies with respect to the male songs are described for four species of the palaearctic bushcricket genus Barbitistes (B. constrictus, B. ocskayi, B. serricauda, B. yersini). In a male song, 3 to 16 syllables form a chirp followed by a trigger syllable after a longer interval. The trigger syllable releases a female reply with a latency of 30 to 50 ms in all four species. In B. serricauda songs, there is no clearly separated trigger syllable. Instead, the first syllable of a chirp functions as a trigger syllable. Some B. serricauda males may produce a short female-type syllable just at the moment, when a female would reply. The possible function of such a syllable is acoustical mimicry. When comparing at least two song parameters, each species occupies a specific combination of values. According to the overlap of parameters a close phylogenetic relationship between B. constrictus and B. serricauda and between B. ocskayi and B. yersini is assumed. This interpretation is compared with a hypothesis based on morphological investigations.     

Vocal Plasticity – are Pied Flycatchers,Ficedula Hypoleuca,Open‐Ended Learners?

In some songbird species, large song repertoires are advantageous in female attraction, whereas song sharing with neighbours may give an advantage in male–male competition. Open‐ended learners, with the ability to memorize new song elements throughout their lives, may learn from territorial neighbours and thus benefit from increasing both repertoire size and song sharing. A distinction needs to be made between true adult song learning, i.e. memorization of novel song elements, and vocal plasticity resulting in changes in the use of previously memorized elements, such as the use of hidden repertoires or increased production of previously rare syllable types. We assessed the ability of adult pied flycatcher Ficedula hypoleuca males to learn previously unheard song elements and to change their song production in response to playback of unfamiliar, conspecific song, emulating a singing neighbour. After a 1‐week playback treatment, three out of 20 subjects had learned foreign song elements, providing evidence from the wild that pied flycatchers are true open‐ended learners. However, the syllable sharing with the playback stimulus repertoires had not changed, and the males’ repertoires had decreased rather than increased. Hence, we did not find support for increased syllable sharing with neighbours or increased repertoire size as functions of adult song learning in pied flycatchers. Because pied flycatcher song seems to serve mainly for mate attraction, copying of attractive syllable types is a possible alternative function of adult song learning in this species.     

Seasonality in song behaviour revisited: seasonal and annual variants and invariants in the song of the domesticated canary (Serinus canaria)

The song of the domesticated canary (Serinus canaria) is one of the most widely used models to study the neural correlates of behavioural plasticity and the mechanisms of female mate choice. However, only few studies have described the song behaviour in detail and monitored their changes throughout the year, and these data are restricted to the “Waterslager” strain. Here, we studied the song characteristics of the male common domesticated canary at different times of the year, the spring breeding and autumnal non-breeding season, and monitored the birds' songs up to the following breeding season. During breeding, males have increased plasma levels of testosterone, and songs are on average longer and consist of fewer non-repeated syllable types compared to the non-breeding season. When subsequent seasons are compared, song duration and the proportion of non-repeated syllable types change seasonally but not across years. Repertoire size remains constant throughout seasons although syllable types are exchanged. Syllable carry-over is significantly higher from one breeding season to the next than between the breeding and non-breeding season. Further, the repertoire of the breeding season contains more potentially sexually attractive syllable types than that of the non-breeding season. These data show that overall song structure is retained throughout the year while seasonality occurs in the temporal pattern and in repertoire composition.     

C4B3 allotype with a novel Ch phenotype

The fourth component of complement (C4) has two classes of protein, C4A and C4B, both of which have many allelic forms. The serological determinants Rodgers (Rg1, Rg2) and Chido (Ch1, Ch2, Ch3) are generally associated with C4A and C4B, respectively. The C4B3 allotype has been detected in a single Canadian family that expresses a novel Ch phenotype, Ch:–1, 2, –3. There was no information for the Rg determinants, as the C4A ^* 2B ^* 3 haplotype would normally express Rg on the C4A protein. Other C4B3 allotypes in informative families have different Ch phenotypes, and the relationships of these within extended major histocompatibility complex haplotypes are discussed in this paper.     

Relationship between the Sequencing and Timing of Vocal Motor Elements in Birdsong

Accurate coordination of the sequencing and timing of motor gestures is important for the performance of complex and evolutionarily relevant behaviors. However, the degree to which motor sequencing and timing are related remains largely unknown. Birdsong is a communicative behavior that consists of discrete vocal motor elements (‘syllables’) that are sequenced and timed in a precise manner. To reveal the relationship between syllable sequencing and timing, we analyzed how variation in the probability of syllable transitions at branch points, nodes in song with variable sequencing across renditions, correlated with variation in the duration of silent gaps between syllable transitions (‘gap durations’) for adult Bengalese finch song. We observed a significant negative relationship between transition probability and gap duration: more prevalent transitions were produced with shorter gap durations. We then assessed the degree to which long-term age-dependent changes and acute context-dependent changes to syllable sequencing and timing followed this inverse relationship. Age- but not context-dependent changes to syllable sequencing and timing were inversely related. On average, gap durations at branch points decreased with age, and the magnitude of this decrease was greater for transitions that increased in prevalence than for transitions that decreased in prevalence. In contrast, there was no systematic relationship between acute context-dependent changes to syllable sequencing and timing. Gap durations at branch points decreased when birds produced female-directed courtship song compared to when they produced undirected song, and the magnitude of this decrease was not related to the direction and magnitude of changes to transition probabilities. These analyses suggest that neural mechanisms that regulate syllable sequencing could similarly control syllable timing but also highlight mechanisms that can independently regulate syllable sequencing and timing.     

Song characteristics and morphological traits in four populations of the grasshopper Chorthippus biguttulus L.

We investigated four populations of the grasshopper Chorthippus biguttulus with respect to differences in morphological traits and characteristics of their communication signals. A special focus was laid on possible correlations between morphological and song traits of males that could be used by females to infer quality cues of potential mates. We also tested whether females exhibit preferences for males of their own population. Specific song features (onset accentuation, offset, syllable period) of males—but not of females—differed between populations. We observed size differences both in males and females from different populations, but the size ranks of the two sexes were not always correlated. Environmental factors appear to have a strong influence on different size traits, compared to genetic origin. In all populations a specific song feature, the accentuation of syllable onsets, showed a similar correlation with a morphological trait, hind leg size, but its correlation with other size indicators sometimes differed in sign. Females did not prefer songs of males from their own population. The best predictor for song attractiveness was—unexpectedly—not the onset accentuation but the offset depth.     

Temporal patterns of intra- and interspecific acoustic signals differ in two closely related species of Acanthoplus (Orthoptera: Tettigoniidae: Hetrodinae)

Males of the closely related African tettigoniids Acanthoplus discoidales and Acanthoplus longipes produce a long-lasting calling song and a short disturbance sound. The temporal patterns of the sounds were analysed in respect to species differences and song type differences. The calling songs of both species consist of impulses which are separated into verses of two syllables, with fewer impulses in the first syllable. A. longipes produces more impulses in each syllable than A. discoidales and has longer verse durations, verse intervals and syllable intervals. Also, the disturbance sounds, produced after mechanical stimulation, contain distinct verses of impulses. The disturbance sound of A. longipes has a higher number of impulses per verse than that of A. discoidales. The frequency spectra of the songs in both species have similar peak frequencies (around 12.5 kHz) and both species have their greatest hearing sensitivity in the range between 5 and 10 kHz. Females of A. longipes perform phonotaxis only to songs with a species-specific temporal pattern. By contrast, females of A. discoidales react positively to calling songs of both species.     

Separate localization of sound recognizing and sound producing neural mechanisms in a grasshopper

Summary In the two acridid speciesChorthippus parallelus andCh. montanus, the sound template by which females recognize male song varies with temperature, as does the song itself. At relatively high temperatures the females respond best to simulated songs with high syllable frequencies, and at lower temperatures songs with lower syllable frequencies are preferred.The temperature around the supraesophageal and metathoracic ganglia of female grasshoppers was monitored by implanted thermocouples, and either the head or the thorax was warmed selectively while the animal was free to move (within the imits of the wires). Then simulations of the conspecific song varying in syllable frequency corresponding to different song temperatures were presented, and the stridulatory responses of the animals were observed.The results were as follows. 1. Song recognition (in particular, the position of the peak of the response curve) depended on the temperature of the head. 2. The rate of stridulatory hindleg movement was determined by the temperature of the thoracic ganglia.This result provides strong evidence against the genetic coupling hypothesis.     

Effects of Song Experience and Song Quality on Immediate Early Gene Expression in Female Canaries (Serinus canaria)

Female songbirds are thought to make mate choices based on aspects of male song quality. Male canaries (Serinus canaria) produce songs with “special” syllables that have been shown to be highly salient to female listeners – eliciting high rates of sexual displays and enhanced immediate early gene (IEG) expression. Immunohistochemistry for the IEG ZENK was used to examine the effects of experience with these syllables on activity in the caudal mesopallium (CMM) and nidocaudal mesopallium (NCM), two auditory areas important in processing conspecific song. Photostimulated female canaries were housed in sound attenuated chambers and played pseudosongs containing either three special syllables or three non‐special syllables, an intro, and an outro sequence. Females that heard special syllable pseudosongs exhibited higher ZENK expression in CMM. To assess the effects of experience, photostimulated females were pair housed and exposed to playback of song with or without special syllables for 14 days. After transfer to individual housing, birds were played one of the aforementioned stimuli or silence. ZENK expression in CMM and NCM was equivalent for song with and without special syllables, but significantly lower for silence. Females who experienced song with special syllables had lower plasma estradiol concentrations after final song playback. This study indicates that CMM exhibits an IEG response bias to special syllables in limited acoustic contexts, but not in full song, which may contain additional biologically relevant information. Furthermore, estradiol concentrations may mediate changes in song responses, serving as a mechanism for modulating mate choice in differing song environments.     

Sex-specific spectral tuning for the partner's song in the duetting bushcricket Ancistrura nigrovittata (Orthoptera: Phaneropteridae)

The song of the male bushcricket Ancistrura nigrovittata consists of a sequence of verses. Each verse comprises a syllable group, plus, after about 400 ms a single syllable serving as a trigger for the female response song. The carrier frequency of the male song spectrum peaks at around 15 kHz, while the female song peaks at around 27 kHz. The thresholds of female responses to models of male songs are lowest for song frequencies between 12 and 16 kHz and therefore correspond to the male song spectrum. The threshold curve of the female response to the trigger syllable at different frequencies has the same shape as the tuning for the syllable group. Phonotactic thresholds of male Ancistrura nigrovittata to synthetic female responses at different frequencies are lowest between 24 and 28 kHz and thereby correspond to the female song spectrum and clearly differ from female response thresholds. Activity of the tympanic fibre bundle of both sexes is most sensitive between 15 and 35 kHz and therefore not specifically tuned to the partner's song. Individual behavioural thresholds have their minimum within 10 dB of the values of tympanic thresholds.     

Filtering of temporal parameters of the calling song by cricket females of two closely related species: a behavioral analysis

The phonotactic response of cricket females was investigated on a locomotion compensator to determine the temporal parameters of the male's calling song which are important for species recognition. Two sympatric species (Teleogryllus commodus, T. oceanicus) that show different syllable periods in the chirp and trill parts of their calling songs were used. By their responses T. commodus females exhibited two temporal filters for syllable periods, which were tuned to the species-specific syllable periods occurring during chirp and trill. For song recognition both filters had to be activated and for both a minimum number of three to five consecutive syllable periods was necessary. In contrast, T. oceanicus females showed only one sharply tuned filter corresponding to the chirp part of the male's calling song. This filter was sufficient for calling song recognition. Syllable periods of the trill part also influenced calling song recognition, but these played only a minor role. Carrier frequency was also important for positive phonotaxis. Calling song recognition by T. commodus females is largely based on central nervous processing, while for T. oceanicus both peripheral frequency filtering and central temporal filtering is important. Accepted: 17 January 1997     

Sexual differences in cricket calling song recognition

Summary Phonotactic behavior was studied in male crickets,Teleogryllus oceanicus. Tethered flying males were presented with electronically synthesized calling song models in a two-choice phonotaxis assay, and their song preferences were determined and compared with previous findings for females.Males are poorer at discriminating between songs than females; they do not display choice behavior as frequently as females, and the choices they do make are not as consistent as those of females (Figs. 3, 4). T. oceanicus calling song is composed of rhythmically different chirp and trill sections. The selectivity of males for these two components differs from that of females. Females prefer chirp to trill, but the opposite is true for males (Fig. 5B-F). Males are similar to females in that they prefer either a conspecific song model or its separate components to a heterospecific model (Fig. 5A, G, H).Behavioral and neural implications of these findings are discussed.     

Temporal resolution for calling song signals by female crickets, <Emphasis Type="Italic">Gryllus bimaculatus</Emphasis>

A behavioural gap detection paradigm was used to determine the temporal resolution for song patterns by female crickets, Gryllus bimaculatus. For stimuli with a modulation depth of 100% the critical gap duration was 6–8 ms. A reduction of the modulation depth of gaps to 50% led either to an increase or a decrease of the critical gap duration. In the latter case, the critical gap duration dropped to 3–4 ms indicating a higher sensitivity of auditory processing. The response curve for variation of pulse period was not limited by temporal resolution. However, the reduced response to stimuli with a high duty cycle, and thus short pause durations, was in accordance with the limits of temporal resolution. The critical duration of masking pulses inserted into pauses was 4–6 ms. An analysis of the songs of males revealed that gaps (5.8 ms) and masking pulses (6.9 ms) were at detectable time scales for the auditory pathway of female crickets. However, most of the observed temporal variation of song patterns was tolerated by females. Critical cues such as pulse period and pulse duty cycle provided little basis for inter-individual selection by females.