title: Yolk steroids in great titParus majoreggs: variation and covariation between hormones and with environmental and parental factors.
abstract: Avian mothers can potentially alter the phenotypes of their offspring by varying the concentration of steroid hormones in their eggs. We explored variation in androstenedione (A4), testosterone (T), 5α-dihydrotestosterone (DHT), 17β-estradiol (E2), and corticosterone (CORT) in the yolks of 12 free-living great titParus majorclutches. We analyzed variation and covariation in greater detail than previous studies, using models for variation with laying sequence that take into account variable clutch size and comparing correlations between pairs of hormones at the within- and between-clutch levels. We also investigated relationships between hormone levels and various environmental, life history, and parental traits. For three of the five steroids, we found no significant correlates, but based on individual statistical tests (a) DHT varied between clutches with male age (1 year old vs older); (b) DHT and CORT were negatively correlated within clutches with the average temperature on the day (DHT and CORT) or 3 days (DHT only) preceding laying; and (c) DHT in the last egg of the clutch relative to the clutch mean was positively correlated with the interval between clutch completion and the onset of incubation (incubation delay). Relationships with ambient temperature and incubation delay have not previously been reported for any yolk hormone in birds. Intriguingly, the three relationships for DHT are consistent with more DHT being transferred to eggs in situations that could be more energetically challenging for the female. More research is needed to determine the generality of the patterns we found and to understand their functional significance.
  
  
  CONCLUSIONS
  The yolks of birds' eggs contain steroid hormones produced by the mother which can affect the development and behavior of the resultant chicks. We analyzed five steroid hormones in the yolks of wild great tits and show for the first time that yolk hormone levels are related to ambient temperature in the day(s) just before laying and, in the last-laid egg, with the day it is laid relative to the onset of incubation, and that the concentrations of pairs of yolk hormones can vary with each other in a different way between and within clutches. These results contribute insights into the ways in which yolk hormones may adaptively modify the chicks or may reflect physiological processes occurring in the mother.
authors.affiliation: Department of Animal Ecology, Netherlands Institute of Ecology (NIOO-KNAW), Droevendaalsesteeg 10, PO Box 50, 6700 AB Wageningen, The Netherlands.
authors.forename: C M
authors.initials: CM
authors.lastname: Lessells
authors.affiliation: Department of Animal Ecology, Netherlands Institute of Ecology (NIOO-KNAW), Droevendaalsesteeg 10, PO Box 50, 6700 AB Wageningen, The Netherlands ; Section of Ecology, Department of Biology, University of Turku, Turku, Finland.
authors.forename: S
authors.initials: S
authors.lastname: Ruuskanen
authors.affiliation: Center for Reproductive Biology, School of Biological Sciences, Washington State University, Pullman, WA 99164 USA.
authors.forename: H
authors.initials: H
authors.lastname: Schwabl
keywords: Ambient temperature; Avian; Birds; Hatching asynchrony; Incubation; Yolk steroids