A brood manipulation experiment on great tits Parus major was performed to study the effects of nestling age and brood size on parental care and offspring survival. Daily energy expenditure (DEE) of females feeding nestlings of 6 and 12 days of age was measured using the doubly-labeled water technique. Females adjusted their brooding behavior to the age of the young. The data are consistent with the idea that brooding behavior was determined primarily by the thermoregulatory requirements of the brood. Female DEE did not differ with nestling age; when differences in body mass were controlled for, it was lower during the brooding period than later. In enlarged broods, both parents showed significantly higher rates of food provisioning to the brood. Female DEE was affected by brood size manipulation, and it did not level off with brood size. There was no significant effect of nestling age on the relation between DEE and manipulation. Birds were able to raise a larger brood than the natural brood size, although larger broods suffered from increased nestling mortality rates during the peak demand period of the nestlings. Offspring condition at fledging was negatively affected by brood size manipulation, but recruitment rate per brood was positively related to brood size, suggesting that the optimal brood size exceeds the natural brood size in this population. Key words: brood size manipulation, doubly-labeled water technique, energy expenditure, great tits, parental care, Parus major. [Behav Ecol 10:598-606 (1999)] O ne of the central questions in life-history evolution is how the timing and intensity of reproduction is allocated over the lifetime of an individual. Since Lack (1947) suggested that clutch size in birds was adjusted to the number of offspring that the parents are able to raise, much work has been done to identify the factors limiting the intensity of reproduction. As a model, clutch size variation in birds has been studied intensively because it allows manipulation of the parental choice and estimation of the consequences in terms of effort or fitness (Stearns, 1992). To identify selection pressures we have to (1) relate effort to brood size, (2) relate brood size to brood fitness, (3) relate effort to parental fitness, and (4) relate total fitness to brood size. In the present paper, we concentrate on the first two questions.Parental effort has been defined by Williams (1966) as any behavior a parent may perform to enhance the reproductive value of the offspring of its current reproductive attempt at the cost of the parent's ability to invest in future offspring. The best way to explore the relationship between parental effort and brood size may be by experimental manipulations of reproductive effort (Gustafsson and Sutherland, 1988;Tinbergen and Daan, 1990;van Noordwijk and de Jong, 1986). As a manipulation of parental effort, experimental modifications of brood size in birds have been studied (Stearns, 1992), and parental provisioning is often used as a measure of parental effort. Research has ...