Physiological adjustments of the budgerygah (Melopsittacus undulatus) to dehydrating conditions.
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Aspartate aminotransferase (AST), alanine aminotransferase (ALT), creatine kinase (CK) and lactate dehydrogenase (LDH) concentrations were measured in 73 budgerigars. In spite of increases in the serum concentration of AST and ALT the loss of enzyme activity from the injection site occurred at about the same rate for all four enzymes tested. The rate constants, calculated in birds administered muscle extract via the intravenous or the intramuscular route of injection, disclosed that in subjects treated intramuscularly the serum enzyme elevation was dependent on the elimination half-life of the enzyme and partly on the rate at which the particular enzyme was absorbed into the bloodstream. Moreover, serum enzyme elevations correlated inversely with total clearance. To determine the aetiology of blood enzyme elevation in budgerigars, for diagnostic purposes, the absorption rate, total clearance and elimination half-life of each of these four enzymes are deemed valuable components.
Conditioning studies involving administration of biochemical or pharmacological agents are often stressful to experimental animals. Some of these stressful manipulations were examined for their effect on the avoidance conditioning of the budgerigar to sound stimuli. There were individual variations in the capacity of the birds to accommodate to these stresses, but generally the stress factors had negative effects on their subsequent conditioning, the degree of this effect depending upon the type and severity of stress. Simple anesthesia with ether had the least effect, intravenous injection being next in severity, while intracerebral injection was most inhibitory. The effects due to anesthesia and intravenous injection could be decreased by delaying the conditioning procedure for 1 to 2 hours following the treatment, but the effect of intracerebral injection was more prolonged. The retrieval of temporarily lost conditioned response through brief reconditioning was not affected by intracerebral injection, indicating that, once a memory trace was formed, subsequent emotional and physical stresses did not obliterate it irreversibly.
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Seminal plasma composition was studied in budgerigars. Semen was obtained from adult male budgerigars by applying gentle pressure to both sides of the cloaca. Pooled samples were centrifuged at 15,000 g for 2 min, and the seminal plasma separated for biochemical analysis. Osmolality, Na+, K+, Cl-, pH, glucose and fructose values were determined. The biochemical composition of budgerigar seminal plasma obtained in this study was: Osmolality 329.9 +/- 14.5 mOs/kg; Na+ 158.6 +/- 8.4 mEq/l; K+ 16.39 +/- 6.24 mEq/l; Cl- 109.2 +/- 7.4 mEq/l; pH 8.20 +/- 0.18 glucose 4.25 +/- 0.96 mmol/l; fructose 0.59 +/- 0.29 mmol/l. The results are discussed in relation to the values reported for the domestic fowl. This forms part of a reproductive biology study of non-domesticated avian species.
Backward, foward and simultaneous masking of brief tone pulses was examined in four parakeets. The pattern of non-simultaneous masking in parakeets is quite similar to that observed in humans with rates of threshold recovery faster for backward compared to forward masking conditions. These results may have relevance for the design of vocal signals and the coding of species-specific information in parakeets.
Three budgerigars and three zebra finches were tested for their ability to detect sinusoidal stimuli in the presence of broadband noise. Masked thresholds for 1, 2, and 4 KHz pure tones were measured with a fixed frequency condition, in which only one test frequency was presented in a session, and with an uncertain frequency condition, in which three signal frequencies were presented in random order in one session. The critical signal/noise ratios obtained in the fixed frequency condition were similar to those reported in a previous study (Okanoya and Dooling, 1987) for both species. When tested in the uncertain frequency condition, critical ratios for zebra finches increased for 1.5 dB at 1 KHz signal but remained unchanged 2 and 4 kHz. The critical ratios for budgerigars showed no difference at any frequency in the uncertain frequency condition. These results suggest that (1) budgerigars and zebra finches are similar in the degree to which attention factors are involved in the detection of signals in noise, and (2) the unusual shape of the budgerigar critical ratio function is not the result of central attentional processes.
Temporal gap detection thresholds were obtained for two species of birds, budgerigars and zebra finches, which are known to have different auditory filter bandwidths. Both species showed gap detection thresholds of about 2.5 msec for broadband noise stimuli. Comparing octave bands of noise centered at 1, 3, and 5 kHz, zebra finches showed the smallest gap thresholds for the noise band centered at 5 kHz whereas budgerigars showed the smallest gap detection thresholds for the noise band centered at 3 kHz. The results from zebra finches are generally consistent with filter theories of auditory spectro-temporal perception whereas the result from budgerigars are not. In aggregate, these comparative data suggest the relation between spectral and temporal resolving power in these two species may involve different mechanisms.
Among avian species that learn their vocalizations, budgerigars have been thought to be unusual in receiving auditory input to the vocal control circuitry of the telencephalon via an auditory component of nucleus basalis (NB), rather than via Field L2, the end station of the classical thalamofugal pathway. In the present study, however, the application of neural tracing techniques showed the two auditory pathways to be reciprocally connected within the telencephalon. The implication is, therefore, that auditory information essential for vocal learning and vocal production in the budgerigar is derived from the primary as well as the secondary auditory pathway.
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Contact call-driven zenk (zif268, egr1, NGF1A, Krox 24) mRNA expression was mapped with in situ hybridization histochemistry in a vocal learning parrot, the budgerigar (M. undulatus). Relative to controls, call stimulation induced high zenk mRNA expression in all auditory areas including those closely associated with the vocal system within the anterior forebrain (Brauth et al. (2001) J. Comp. Neurol. 432, 481; (2002) Learn. Memory 9, 76). Thus there is a high correspondence between the distributions of neurons exhibiting contact call-driven zenk protein and mRNA expression in budgerigars. Field L2a, an area reported previously to express only perinucleolar zenk protein localization (Brauth et al. (2002) Learn. Memory 9, 76) also showed zenk mRNA expression.
A new combination of operant conditioning and psychophysical scaling procedures was used to study auditory perception in a small bird. In a same-different discrimination task, budgerigars learned to discriminate among pure tones that varied along one or more acoustic dimensions. Response latencies were used to generate a matrix of interstimulus similarities. Multidimensional scaling procedures were used to arrange these acoustic stimuli in a multidimensional space that supposedly reflects the bird's perceptual organization. For tones that varied in intensity, duration, and frequency simultaneously, budgerigars were much more sensitive to frequency changes. From a set of tones that varied only in intensity, it was possible to calculate the growth of loudness with intensity for the budgerigar. For tones that varied only in frequency, budgerigars showed evidence of an "acoustic fovea" for frequency change in the spectral region of 2-4 kHz. Budgerigars and humans also differed in their perceptual grouping of tone sequences that rise, fall, or remain constant in pitch. Surprisingly, budgerigars were much less responsive to pitch contour than were humans.
Operant conditioning and multidimensional scaling procedures were used to study auditory perception of complex sounds in the budgerigar. In a same-different discrimination task, budgerigars learned to discriminate among natural vocal signals. Multidimensional scaling procedures were used to arrange these complex acoustic stimuli in a two-dimensional space reflecting perceptual organization. Results show that budgerigars group vocal stimuli according to functional and acoustical categories. Studies with only contact calls show that birds also make within-category discriminations. The acoustic cues in contact calls most salient to budgerigars appear to be quite complex. There is a suggestion that the sex of the signaler may also be encoded in these calls. The results from budgerigars were compared with the results from humans tested on some of the same sets of complex sounds.
We trained budgerigars by operant conditioning to discriminate among a set of contact calls in a same-different task and analyzed response latencies from this task by using multidimensional-scaling (MDS) and cluster-analysis procedures. Humans listened to the same calls and indicated the similarity between pairs of calls by a direct rating procedure. An MDS program (sindscal) was used to arrange these complex acoustic stimuli in multidimensional space reflecting perceptual organization. Multiple regression techniques were used to identify the acoustic characteristics of contact calls that were correlated with the perceptual dimensions obtained from MDS. A number of spectral characteristics (e.g., peak frequency, rate of frequency modulation, and concentration of spectral energy) emerged as important for both budgerigars and humans, but the relative salience of these cues differed for the two species. Additional tests with two groups of budgerigars--cagemates and noncagemates--showed that experience with calls can change the salience of various acoustic characteristics used for perceptual organization and individual recognition.
Budgerigars have a complex vocal repertoire, some of which develops through learning. The authors examined the course of vocal development in budgerigars from hatching to about 4 weeks postfledging (approximately 85 days old). Food-begging calls showed changes in duration, peak frequency, bandwidth, and frequency modulation with age. Within a week of fledging, each bird produced a contact call bearing a strong resemblance to a shortened version of its patterned food-begging call. By 4 weeks postfledging, budgerigar contact call repertoires often contained more than one call type, and there was clear evidence of sharing and imitation among the calls of parents, fledglings, and other social companions. Perceptual testing showed that whereas acoustic variation in the structure of developing calls decreased both within and between nestling birds, the discrimination of these calls was easier for adult birds as young birds matured. These results suggest parallels with certain aspects of language development in humans.
Thresholds for detecting alterations in the timbre and harmonicity of complex harmonic signals were measured in zebra finches, budgerigars, and humans. The stimuli used in this experiment were designed to have particular salience for zebra finches by modeling them after natural zebra finch calls. All 3 species showed similar abilities for detecting an amplitude decrement in a single component of a harmonic complex. However, zebra finches and budgerigars were extraordinarily sensitive to the mistunings of single harmonics and exhibited significantly lower thresholds compared with humans at 2 different fundamental frequencies, 570 Hz and 285 Hz. Randomizing relative phases of components in a harmonic stimulus resulted in a significant increase in threshold for detecting mistunings in zebra finches but not in humans. Decreasing the duration of mistuned harmonic stimuli resulted in higher thresholds for both birds and humans. The overall superiority of birds in discriminating inharmonicity suggests that birds and mammals may use different strategies in processing these complex harmonic sounds.
The detection of signals in noise is important for understanding both the mechanisms of hearing and how the auditory system functions under more natural conditions. In humans, the auditory system gains some improvement if the signal and noise are separated in space (binaural masking release). Birds with small heads are at a disadvantage in separating noise and signal sources relative to large mammals, because interaural time differences are much smaller. Two binaural phenomena in budgerigars related to the detection of tones in noise were examined. Budgerigars show 8 dB of free-field binaural masking release when signal and noise are presented to their right side and correlated noise is presented to their left side. Budgerigars also show a spatial masking release of 9 dB when a signal and noise are separated in azimuth by 90 degrees. These results are similar to those found in humans and other mammals with much larger heads.
Avoidance conditioning and the method of limits were used to measure absolute auditory thresholds, masked thresholds, and critical ratios in 4 parakeets. The same procedure was then used to study frequency difference limens in 6 additional animals. The power spectrum and "constancy of intonation" of the parakeet call were also measured and related to the absolute and differential frequency sensitivity. The mechanism of frequency analysis in the parakeet ear was considered in relation to the present results and to the anatomical and functional differences between the avian and mammalian auditory systems.