Energetics of laying and incubation in birds: studies of swallows Hirundo rustica, dippers Cinclus cinclus and Japanese quail Coturnix coturnix

Abstract

The doubly labelled water technique was validated for captive-bred, laying Japanese quail (Coturnix coturnix), and used to measure the energy expenditure of free-living laying swallows (Hirundo rustica). Swallows had a slightly higher energy expenditure during laying than during incubation or nestling-rearing. Energy expenditure and the net cost of incubation increased sharply for female dippers (Cinclus cinclus) when clutches were enlarged to 6 eggs.

Lipophilic dyes were used to determine the rate of ovarian follicular growth and the volume of yolk deposjted in a 24 h period by captive quail and free-living swallows. Rates of yolk deposition were related to daily energy expenditure in quail, but not in swallows.

Balances placed under swallow nests recorded an increase in female mass from 5 d before the first egg was laid. Female mass peaked on the evening before the first egg and declined as eggs were laid. Mass changes during laying were equal to the mass of the oviduct and developing ova. However, body composition also changed, as a lipid reserve was built up in the final 4 d before the first egg was laid, whilst body water content declined. This substantially increased the peak energy requirement for biosynthesis in a laying swallow. The lipid reserve was catabolized during the remainder of the laying period. The lipid reserve was likely to serve as an insurance against a drop in food intake during laying. Shortage of food on the day before the first egg was laid led to a reduction in clutch size for some swallows. There was no evidence for use of a protein reserve by laying swallows.

A model was developed from which it was predicted that egg production by swallows, and probably all other insectivorous birds, would be constrained by energy rather than crude lipid or protein requirements. It was concluded that laying patterns and clutch sizes were sometimes constrained by food availability during egg-laying, and that an upper limit to clutch size could be set by the capacity of an incubating bird to cover the eggs.