Digit ratio varies with sex, egg order and strength of mate preference in zebra finches

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Abstract
The steroid environment encountered by developing vertebrates has important organizational effects on physiology and behaviour that persist throughout an organism's lifetime. Optimal allocation of maternal steroids to zygotes may be difficult to achieve because of the sexually antagonistic effects of steroids; thus, for example, a hormone environment beneficial to a developing male may be much less beneficial to a developing female. Research into the important topic of how mothers might adaptively adjust steroid titres experienced by particular young has been constrained by the difficulty of measuring the steroid environment experienced by the embryo at critical times in development. A potential approach to this problem has been suggested by research on variation in digit ratios in humans, where the ratio of the length of the second and fourth digits reflects the steroid environment experienced by the foetus; notably, digit 4 lengthens in response to androgens. In light of the conservative nature of homeobox genes regulating early development in tetrapods, we questioned whether a sex difference in digit ratio exists in a passerine bird, the zebra finch, Taeniopygia guttata castanotis, and whether observed variation in the ratio is consistent with the previously reported pattern that androgen allocation to zebra finch egg yolk declines across laying order. We established an aviary population of outbred, wild–type zebra finches, and allowed them to breed freely. Hatchlings were marked to correspond to their egg order, and their digit ratios were measured after birds reached adulthood. We found that digit ratio increased across egg order, which is consistent with a pattern of decreasing androgen allocation. Moreover, digit ratios differed between the sexes. We also investigated whether variation in digit ratio among adult females predicted variation in their performance in mate–choice tests. Digit ratio accounted for almost 50% of the variance in strength of female preference for an attractive male trait: specifically, females with higher (presumably less ‘androgenized’) ratios had stronger preferences for attractive males. Digit ratio may prove to be an extremely useful tool for addressing a wide range of questions about vertebrate differentiation and behaviour.

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