A comparison between two ways to measure minimum frequency and an experimental test of vocal plasticity in red-winged blackbirds in response to noise

in Behaviour
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We examined whether red-winged blackbirds modulate their vocalizations in response to experimental highway noise, alternating between ambient-control and noise-playback periods. Our measures of song duration were shorter, and with a lower value of freq5% (a measure of energy distribution), during noise-playback; however, we interpret these results as noise-induced artefacts. This apparent lack of vocal plasticity should be taken cautiously because we had a small sample size and most birds produced only one song type: song type-related vocal plasticity was unlikely to be found. We found no evidence of a shift in minimum frequency with noise when this was measured with a threshold method on power spectra, but it seemed to increase when measured by eye from spectrograms. Our results suggest that the by-eye practice can lead to bias, which is problematic as several previous studies have used this procedure. Use of the threshold method, over the by-eye practice, is encouraged.

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Figures
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    Noise spectrograms and power spectra. Upper panel shows spectrograms of 3 different 10 s segments (a, b and c) of recorded noise in a busy highway near the city of Davis, CA, USA. Lower panel show their corresponding power spectra. Note that, although most noise energy is present in low frequencies and decreases with frequency, there were also cases with relatively small peaks at higher frequencies (e.g., noise profile below ‘c’).

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    Example showing how we measured the song minimum frequency using the threshold method and by-eye on spectrograms. To obtain the song minimum frequency with the threshold method we first selected the song and obtained its peak frequency from a power spectrum (shown in the lower panel as ‘peak’); this peak frequency had an amplitude of 91.4 dB. We subtracted 20 dB from this amplitude to obtain 71.4 dB (shown with a horizontal line in the lower panel). In this example, this horizontal line intersects the power spectrum curve four times to the left of the peak (numbers 1, 2, 3 and 4); intersections 1, 2 and 3 resulted from song energy, and intersection 4 resulted from background noise energy. In this example there were three intersections to the right of the peak (numbers 1, 2 and 3), all resulting from song energy. We defined the minimum and maximum frequencies as those that corresponded to the last intersection, to the left (intersection 3) and right (intersection 3) of the peak respectively, that resulted from song energy (see text for further details). To obtain the minimum frequency by eye we placed the lower limit of the selection on the lowest song frequency as judged with our eyes (lower horizontal broken line in upper panel, broken vertical line at the left in the lower panel). To obtain the maximum song frequency we raised the upper limit of the cursor up to the point where no increment of energy (dB) could be detected (higher broken horizontal line in the upper panel, broken vertical line at the right in the lower panel, see text for more details). This figure also shows that in some cases the threshold method did not capture the lowest syllable in the song (indicated with an S in the upper panel), especially when the signal to noise ratio was suboptimum. Note that the amplitude values are not calibrated (Raven calculates decibels values relative to 1); thus y-axis values cannot be compared among recordings or studies, but they are suitable for relative comparisons of amplitude within recordings, and for measurements involving difference in amplitude, such as those described here.

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    Waveform (upper panel) and spectrogram (bottom panel) of a red-winged blackbird song recorded in Conaway Ranch, close to the City of Davis, CA, USA. Songs in this population are made of visually distinctive traces in the spectrogram (straight black lines below ‘Song duration’ are added to aid visualization); each of these were considered as a different syllable. Song energy fades out as frequency increases. This sound file was high-pass filtered at 1 kH and the figure was made with Avisoft-SAS Lab lite v. 5.2.09.

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    Red-winged blackbird calls. We identified 10 different call types in our population. Calls recorded in Conaway Ranch, close to the city of Davis, CA, USA. Cheer-like calls: (a) cheer, (b) harmonic cheer, (c) broken cheer. Check-like calls: (d) check, (e) check 2, (f) check 3, (g) check 4, (h) check 5, (i) check 6, (j) check 7.

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    Minimum frequency of songs measured with the by-eye practice (a) and with the threshold method (b). When we used the by-eye practice to obtain the song minimum frequency, we found a significant increase in song minimum frequency during the noise-playback treatment (p=0.011), but there was no significant effect when measured with the threshold method (p=0.229). Every two points linked with a line represent a male that sang both during the ambient noise (Low) and noise-playback (High) treatments.

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    Graphic examples of minimum frequencies measured by eye and threshold method in the same song type during ambient-control and noise playback periods. Song minimum frequencies obtained by eye (by-eye practice) during ambient-control (panel a) and noise-playback treatment (panel b), and the threshold method during ambient-control (panel c) and noise-playback treatment (panel d). The by-eye practice commonly scored higher minimum frequency values at higher noise levels (b) than at lower noise levels (a); the threshold method sometimes missed the lowest syllable of the song (c, d). Letter L above an arrow (a, c) represents the lowest syllable in the song. This syllable was masked by noise during the noise-playback treatment (b, d), and was not measured with the by-eye practice during the noise-playback treatment (b), or with the threshold method during the ambient-control and noise-playback treatments (c, d). Thus, one possible downside of the by-eye practice is bias, while a possible downside of the threshold method is missing the lowest frequency.

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    Minimum frequency of check-like calls measured with the by-eye practice and with the threshold method. When we measured the minimum frequency by-eye on spectrograms (a), we found a significant increment of check-like call minimum frequency during the noise-playback treatment (p=0.002), but not when we used the threshold method (p=0.927) (b). Every two points linked with a line represent a male that called both during the ambient-control (Low) and noise-playback (High) treatments.

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