TL;DR: In this paper, the parent adjusts its effort in relation to prevailing environmental conditions in order to maximize the output of young in its lifetime, rather than measurable in terms of adult survival and recruitment of young.
Abstract: 1. Energetics of reproduction in birds is reviewed with the question in mind how the parent adjusts its effort in relation to prevailing environmental conditions in order to maximize the output of young in its lifetime. Emphasis is on proximate controls, rather than ultimate factors measurable in terms of adult survival and recruitment of young. 2. The decision to breed or not to breed is clearly related to body condition of the female, presumably because of the implications this has for survival. 3. Laying date and clutch size are likewise under the influence of female condition and can hence be modified by experiments involving supplementary feeding. Natural variation in these features may often be related to territory quality. 4. How the bird decides whether or not to commence a second brood is not clear, but in the Great Tit the habitat-related difference in incidence of second broods is functionally understandable when survival probabilities of birds at different times are considered. 5. A distinction is made between a "capital" and "income" model for translatting rates of change of female body condition into appropriate decisions on laying date and clutch size and experiments are suggested that discriminate between the two. 6. Lack's view that brood size is in an evolutionary sense adjusted in order to balance food requirement and foraging capacity of the parents is accepted, and growth rates in nidicolous birds are analysed to ascertain if a finer adjustment exists superimposed on the integer steps of brood adjustment. Critical for this analysis are groups of birds where broods of one are common, since only in these circumstances is growth adjustment the only strategy open to the parents. In common with other animals, growth rate is related to mature body size but within a category of adult weight clear examples can be found for retardation of growth rate in pelecaniform and charadriiform species with singleton broods. 7. Since daily energy requirement is related to nestling size and growth rate, retardation of growth is explicable as a strategy only in terms of reducing the daily commitment of the parents, not reducing the total cost of producing a nestling. 8. An additional economy in growth is to reduce the contribution of fat to the nestling body. 9. Implied in Lack's view of brood size is a limitation of parental foraging capacity, and the last section of the paper is devoted to exploration of the proximate factors delimiting what Royama terms the optimal working capacity of parents feeding young. Observations of parent starlings confronted with manipulated brood size suggest a limit on the time that can be devoted to energetically extravagant flight activity, rather than a shortage of absolute time. Beyond the limit to which stressed parents can be made to fly, body weight declines. 10. Preliminary data on energy metabolized daily by parents confronted with large broods conforms to the simplified view that parental effort on a sustained basis equates to energy mobilization equivalent to 4 B.M.R. units and it is suggested that this level of energy expenditure represents a proximal decision substrate for determining the optimal working capacity of the parent. 11. The paper ends with a plea for more research on the proximate controls of avian reproduction, and calls attention to the central importance of the protein bank to parental body condition.
TL;DR: Experimental approaches to testing the hypothesis that birds adaptively limit clutch size for the sake of enhanced survival are likely to provide the strongest inferences.
Abstract: Lack (1947) hypothesized that clutch size in nidicolous birds has evolved by natural selection to correspond with the maximum number of young that, on average, the parents can feed. Although the hypothesis gained wide acceptance in subsequent years, the evidence is equivocal and inconsistencies remain (Klomp, 1970; Cody, 1971; von Haartman, 1971; Hussell, 1972). Those cases in which the most productive brood size is larger than the most common do not support the implied concept of direct limitation of clutch size by food supply. Furthermore, the interpretation of brood manipulation experiments that support Lack's hypothesis is open to question, since the results do not distinguish between food supply limits in the environment and possible adaptive limits upon parental feeding behavior (Cody, 1971; Hussell, 1972). Mountford (1968) suggested that incorrect formulation of predictions was responsible for some apparent contradictions. Lack recognized that selection should favor the clutch size that maximizes fitness, but overemphasized the direct influence of environmental factors upon clutch size as a single trait. A more comprehensive perspective incorporates interactions between clutch size and other life history features as well (Fisher, 1958; see review in Stearns, 1976). Pertinent models predict a most common clutch that is smaller than the most productive, under the assumption that rearing larger broods places greater stress upon parents and reduces their chances of surviving to breed again (Williams, 1966; Charnov and Krebs, 1974). The assumption of a trade-off between clutch size and adult survivorship within populations is largely untested. Observations from naturally occurring brood sizes have not yielded any consistent relationship between clutch size and parental survival (e.g., Kluyver, 1963; Perrins, 1965; Lack, 1966, p. 109). However, since variation in parental ability (e.g., efficiency in gathering food for egg formation or for feeding nestlings) may contribute to adaptive modification of clutch size (cf. Klomp, 1970), the search for such a relationship is confounded. If parents that normally initiate larger clutches are more capable of rearing them, the young in.those broods are not necessarily disadvantaged (cf. Perrins and Moss, 1975), nor are those parents necessarily less likely to survive than parents raising smaller broods. Greater weight losses among parents with larger broods (Hussell, 1972; Winkel and Winkel, 1976; Bryant, 1979) and lower probabilities of initiating a second brood after a large first brood (Kluyver, 1963; Pinkowski, 1977) provide indirect evidence that rearing large broods is stressful physiologically, but have yet to be linked to differential survival. Recently, Bryant (1979) found differences in survival between singleand double-brooded female House Martins (Delichon urbica), but these differences were not related to the brood sizes reared. Experimental approaches to testing the hypothesis that birds adaptively limit clutch size for the sake of enhanced survival are likely to provide the strongest inferences (cf. Ricklefs, 1973, p. 426; Stearns, 1976, p. 42). Artificial manipulation can extend brood sizes beyond limits currently observed within a population. However, if parental ability is reflecterd in individulm] clutch sizes. nqirpnts 278
TL;DR: A simple model of clutch size in nidicolous birds is developed in which the cost of reproduction is the risk of predation to both the parent and its dependent young, and a predation-risk perspective may offer considerable insight into the evolution of clutchsize in birds.
Abstract: A simple model of clutch size in nidicolous birds is developed in which the cost of reproduction is the risk of predation to both the parent and its dependent young. An analysis of the model shows that (1) conventional ideas of food limitation, though sufficient, are not necessary for the existence of an optimal clutch size; (2) predation as the sole cost of reproduction is adequate for the existence of an optimal clutch size; and (3) one need not expect a clutch-size-dependent physiological cost to a parent (leading to increased mortality) to be a major determinant of clutch size. In addition, several exper- imental and observational results that purport to demonstrate food limitation in breeding birds are also consistent with the idea of predation risk as the major cost of reproduction. Current ideas of clutch size determination and the costs of reproduction are considered in light of the above results, and possible syntheses are suggested. A predation-risk perspective may offer considerable insight into the evolution of clutch size in birds.
TL;DR: Results of a series of studies of organ growth in birds with widely varying growth rate capacities show that the growth pattern of the goose, a species which has a high growth rate capacity, is characterized by a rapid early development of the digestive organs and the liver whereas that of the quail and the turkey, species which have low growth rates, are characterized by an rapid earlyDevelopment of the pectorals and the feathers.
Abstract: An hypothesis has been formulated stating that the rate at which postnatal growth proceeds is at least partly determined by the distribution of growth between different organs. In order to test the hypothesis a series of studies of organ growth has been undertaken in some birds with widely varying growth rate capacities (geese, quail, turkeys, fieldfares and jackdaws). With regard to nidifugous birds the data show that the growth pattern of the goose, a species which has a high growth rate capacity, is characterized by a rapid early development of the digestive organs and the liver whereas that of the quail and the turkey, species which have low growth rate capacities, is characterized by a rapid early development of the pectorals and the feathers. The growth pattern of the nidicolous species, the fieldfare and the jackdaw which also show high growth rate capacities, is similar to that of the goose. These results are in close agreement with the hypothesis.
TL;DR: An index of food abundance was obtained and field experiments were performed to distinguish factors affecting variability in growth of nestlings and genetic variation in offspring and variation in pro- visioning abilities of parents may have been important components of within-population variation in growth regardless of where parents nested.
Abstract: We studied two populations of Tree Swallows (Tachycineta bicolor) that differed primarily in the amount of food available to the breeding birds. We obtained an index of food abundance and performed field experiments to distinguish factors affecting variability in growth of nestlings. The experiments were designed to detect the influence of the location of egg laying, incubation, and nestling rearing, type of parent (natural or foster), and year of breeding on nestling growth. Some broods were transferred between nests and raised by foster parents, and some clutches and broods were transferred between populations. Vari- ables were analyzed in two- and three-way factorial analyses of variance. The insect biomass index during the nestling period differed about 7-fold between loca- tions, regardless of year of breeding. Nestlings with more food grew and survived better. Type of parent (i.e. natural or foster) or prehatch factors such as location of incubation did not influence growth. The location where nestlings were raised, however, explained as much as 51% of the variation in growth, and genetic variation in offspring and variation in pro- visioning abilities of parents may have been important components of within-population variation in growth regardless of where parents nested. Received 11 February 1985, accepted 25 November 1985. THE main selective pressures responsible for reproductive performance in nidicolous birds are the availability of food, especially for the young and to a lesser extent for the laying fe- male, and the risk of predation on eggs, young, and parents (Lack 1968). Implicit in Lack's hy- pothesis is a limitation due to parental foraging ability. Nestling growth or survival has been shown (usually indirectly) to be affected by food supply in several bird species (van Balen 1973; von Bromssen and Jansson 1980; Bryant 1975, 1978a; Crossner 1977; Bryant and Gardiner 1979; Ross 1980; Prince and Ricketts 1981). Reduced availability of food markedly affects parental feeding success in some species (Dunn 1973, Bovino and Burtt 1979, Quinney and Smith 1980), and parents that raise large broods may show reduced survival or body mass compared with those that raise smaller broods (Hussell 1972, Askenmo 1977, Tinbergen 1981). Previously, the role of food abundance in re- production often has been obscured because adequate measurements of food supply are dif- ficult to obtain and because of the difficulties