Male birdsong is generally regarded as a secondary sexual characteristic under the control of gonadal steroids1,2. Song typically waxes and wanes with the seasonal cycle of testicular growth and regression and decreases after adult castration. Testosterone therapy reinstates song, induces it in females, augments it in intact males, and spring testosterone profiles correlate with seasonal song production3-10. Thus, testosterone has been viewed as a major factor in song acquisition and production3,11-14 acting either directly, or after aromatization within the brain15,16. We show here, however, that song learning and early phases of the development of singing both take place in castrated male birds with no significant levels of testosterone in their blood plasma. Testosterone seems to be required for song crystallization, however. Oestradiol was unexpectedly still present after castration, evidently from a non-testicular source, throughout the period of male song acquisition.
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