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Summary1. High breeding density can cause elevated plasma androgen levels in adult birds. Since maternal androgens are deposited into egg yolk, high breeding density may result in elevated yolk androgen levels as well. 2. The relationship between breeding density and yolk androgen levels was examined in the European Starling, Sturnus vulgaris . The concentration and total content of yolk androstenedione and yolk testosterone were measured in eggs from 24 clutches distributed across nine different colonies of nestboxes. 3. Yolk androstenedione and testosterone levels were significantly higher in colonies where a greater proportion of nestboxes had active nests. 4. Yolk testosterone levels were significantly higher, and yolk androstenedione levels were marginally higher, in colonies with a greater absolute number of active nests. 5. Yolk androgen levels were not related to the number of active nests in adjacent nestboxes. 6. We conclude that female starlings nesting in colonies with higher breeding densities transfer more androgen to their eggs. 7. This relationship may be mediated by increased interfemale aggression, particularly towards floater females searching for mates or nests to brood parasitize, in high-density colonies. Such a relationship between maternal environment and maternal yolk androgens may represent adaptive maternal modification of offspring phenotype or a non-adaptive physiological constraint which females cannot avoid.

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Yolk steroids of maternal origin have been proposed to influence genetic sex determination in birds, based on sex differences in yolk steroid concentrations of peafowl eggs incubated for 10 days. More recent reports dispute this proposal, as yolk steroids in eggs incubated for 3 days do not show such sex differences. To date, research examining this phenomenon has only analysed incubated eggs, although sex in avian species is determined before incubation begins. This may be a serious methodological flaw because incubation probably affects yolk steroid concentrations. Therefore, we investigated sex differences in yolk steroid concentrations of unincubated avian eggs. We withdrew yolk for steroid analysis from fresh, unincubated Japanese quail (Coturnix japonica) eggs by biopsy, and then incubated those eggs for 10 days, after which we harvested the embryonic material for genetic sexing and the incubated yolk for further steroid analysis. We found no sex differences in fresh Japanese quail eggs; however, sex differences were apparent in yolk steroids by day 10 of incubation, when female eggs had significantly more oestrogen in relation to androgen than male eggs. Concentrations of all yolk androgens decreased dramatically between laying and day 10 of incubation, whereas oestradiol (E2) concentrations increased marginally. Thus, yolk concentrations of androgens and E2 do not appear critical for avian sex determination.

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Chicks of obligate brood parasites employ a variety of morphological and behavioral strategies to outcompete nest mates. Elevated competitiveness is favored by natural selection because parasitic chicks are not related to their host parents or nest mates. When chicks of conspecific brood parasites (CBPs) are unrelated to their hosts, they and their parents would also benefit from traits that enhance competitiveness. However, these traits must be inducible tactics in CBPs, since conspecific brood parasitism (CBP) is facultative. Such tactics could be induced by resources passed to offspring through the egg. Thus, females engaging in CBP should allocate to their eggs resources that will enhance offspring competitiveness. We tested this prediction in a population of European starlings (Sturnus vulgaris) breeding in southern Sweden. Previous research showed that almost all CBPs in this population are floater females that have yet to breed in the current season. We identified putative brood parasitic eggs through monitoring egg laying and verified parasitism using protein fingerprinting. We then determined whether parasitic eggs were larger, larger-yolked, or had higher concentrations of yolk testosterone or androstenedione than control eggs. The 14 brood parasitic eggs laid in active nests (those with clutches of at least two eggs that were eventually incubated) did not differ from controls in any of these characteristics. Ten dumped eggs, laid in nonactive nest-boxes or on the ground, were smaller and smaller-yolked than control eggs but did not differ in yolk androgen concentrations. The failure of our prediction could be the result of high costs of investing in eggs, lack of competition-based benefits for chicks, or physiological constraints on egg manipulation.