Over the past 50 years, songbirds have become a valuable model organism for scientists studying vocal communication from its behavioral, hormonal, neuronal, and genetic perspectives. Many advances in our understanding of vocal learning result from research using the zebra finch, a close-ended vocal learner. We review some of the manipulations used in zebra finch research, such as isolate housing, transient/irreversible impairment of hearing/vocal organs, implantation of small devices for chronic electrophysiology, head fixation for imaging, aversive song conditioning using sound playback, and mounting of miniature backpacks for behavioral monitoring. We highlight the use of these manipulations in scientific research, and estimate their impact on animal welfare, based on the literature and on data from our past and ongoing work. The assessment of harm-benefits tradeoffs is a legal prerequisite for animal research in Switzerland. We conclude that a diverse set of known stressors reliably lead to suppressed singing rate, and that by contraposition, increased singing rate may be a useful indicator of welfare. We hope that our study can contribute to answering some of the most burning questions about zebra finch welfare in research on vocal behaviors. Therefore, the interplay between corticosterone levels during development and long-term consequences of stress needs to be further studied.
Skin temperatureA stressor can increase the core body temperature, provoking a decrease of the surface temperature. This phenomenon known as stress-induced hyperthermia could be utilized to measure stress. Indeed, infrared thermography has been used to determine the effect of stressful procedures in birds (Edgar et al., 2013;Herborn et al., 2015;Jerem et al., 2015) . Contrary to the stress snapshot of corticosterone, this non-invasive imaging method might provide longitudinal measurements of stress.
Body weightBody weight can be used to assess body condition in birds, especially when multiple measurements are made over the course of a study (Brown, 1996;Dickens et al., 2009bDickens et al., , 2009a .