Great tits provided with ad libitum food lay larger eggs when exposed to colder temperatures

The amount of nutrients deposited into a bird egg varies both between and within clutches of the same female. Larger eggs enhance offspring traits, but as a tradeoff, laying large eggs also infers energetic costs to the female. Income breeders usually lay larger eggs later in the season, when temperatures and food availability are higher. Egg size is thus affected by the daily amount of energy available to produce an egg under cold conditions, but it is less well known in how far temperature exerts direct effects on egg size. We show that great tit females Parus major with access to ad libitum food and breeding in climate‐controlled aviaries varied their egg investments. The size of an individual egg was best predicted by mean temperatures one week pre‐laying, with females laying larger, rather than smaller, eggs under colder conditions. Eggs increased in size over the season, but not significantly over the laying sequence. The degree of daily temperature fluctuation did not influence egg size. In addition to a substantial between‐female variation, sisters were more similar to each other than unrelated females, showing that egg size does also reflect heritable intrinsic female properties. Natural variation in egg size is thus not only determined by energy‐limitation, but also due to females allocating more resources to eggs laid in colder environments, thus increasing early survival of the chicks. That the positive correlation between temperature and egg investments that is found in a natural population is reversed under ad libitum food conditions demonstrates that wild great tits tradeoff own condition with survival prospects of their chicks as a function of available food, not ambient temperature.     

Seasonal sex–specific energy expenditure in breeding and non-breeding Palestine sunbirds Nectarinia osea

The timing of the chick‐rearing phase is known to have a profound effect on the reproductive success of birds. However, little is known about the energetic costs faced by the parents during different periods of the breeding season. These costs may have vital consequences for both their survival and future reproduction. In most studies, daily energy expenditure (DEE) of breeding and non‐breeding birds has been compared, without controlling for the effect of season. In the present study, we examined the energy demands of breeding compared to non‐breeding Palestine sunbirds Nectarinia osea and whether there were sex‐specific differences in DEE within and between different seasons. We predicted that DEE would be elevated when birds rear chicks, especially at cooler ambient temperatures. Time‐energy budgets were constructed for pairs of sunbirds, rearing chicks, or not breeding, in spring and summer. There were significant seasonal differences in estimates of DEE in non‐breeders that were 21% higher in spring than in summer. We attributed these to increases in non‐flight metabolic rate rather than changes in time spent on different activities. Our estimates of DEE for the birds that were rearing chicks were higher than non‐breeding adults. In females the increase in DEE when breeding, compared to when not breeding, was similar in both spring and summer, while males increased their DEE much less when breeding in spring. The differences in estimated DEE, however, were not significant between male and female birds in any season. Between seasons, female breeders had 17.1% higher DEE in spring than in summer, while male breeders showed no difference in DEE when rearing chicks in different seasons. Accordingly, our initial prediction was supported, as DEE in chick‐rearing adults was higher than in non‐breeding adults. In addition, although temperatures are lower in spring, breeding in the spring is only more costly than breeding in summer for females. Apparently, males are more flexible in reallocating their time and energy spent on different activities.     

Sexual dimorphism of egg size in the European Blackbird Turdus merula

Recently, a number of studies have shown that female birds are able to control the sex of their progeny at the stage of the gamete. There is also some evidence that females adjust their investment in offspring depending on the sex of the embryo during egg formation. Differential maternal investment to the eggs depending on their sex is usually interpreted as an adaptive strategy, by which females can increase competitive abilities of the smaller sex, or preferentially invest towards the sex with the potentially higher fitness returns. Here, we studied variation in egg size in relation to embryo sex and laying order in the European Blackbird Turdus merula. We found male and female eggs to differ in size, with larger eggs containing male embryos, as well as a significant interaction between embryo sex and laying order. This interaction resulted from the fact that egg size increased with the laying sequence among eggs bearing females but did not change with laying order among eggs bearing males. There was no relationship between offspring sex and the laying sequence within a clutch. We suggest that sexual dimorphism in egg size recorded in the European Blackbird may reflect favouritism of the sex which may give higher fitness returns.     

Variable energetic responses to clutch size manipulations in white-throated dippers Cinclus cinclus

We used the doubly-labelled water technique to measure daily energy expenditure (DEE) of a free-living uniparental incubator, the white-throated dipper Cinclus cinclus , in Scotland. DEE was 205±8 (s.e.m.) kJ d⁻¹ for 17 females incubating their natural clutch sizes, equivalent to 3.2±0.1×basal metabolic rate (BMR). To investigate the influence of clutch size on the energy budget, we measured the DEE of 14 females with clutches increased or reduced by a single egg. Birds with reduced clutch sizes had an energy expenditure with a mean and variance that did not differ from those of birds with unmanipulated clutches. Enlarging the clutch led to an increase in energy expenditure to over 4×BMR for some individuals but not for others, resulting in greater variance in energy expenditure for birds with enlarged clutches. Individual variation in energy expenditure could not be fully explained by environmental conditions, by patterns of behaviour or clutch size. Incubating females received a maximum of only 4 kJ d⁻¹ (2% of DEE) from provisioning by the male, and mobilised up to 6 kJ d⁻¹ (3% of DEE) from reserves. Females spent 2.9±0.2 h (n=20) away from the nest each day, so a foraging rate of 95 kJ h⁻¹ was required during incubation recesses to balance DEE. This 'required foraging rate' is double previous estimates of the maximum rates of energy acquisition for birds of this size. We suggest that the greater likelihood of a raised energy expenditure associated with larger clutches, combined with the difficulties in maintaining energy supplies, may constitute a constraint on avian clutch size.     

Manipulation of life-history decisions using leptin in a wild passerine

Seasonal timing of reproduction and the number of clutches produced per season are two key avian life-history traits with major fitness consequences. Female condition may play an important role in these decisions. In mammals, body condition and leptin levels are correlated. In birds, the role of leptin remains unclear. We did two experiments where we implanted female great tits with a pellet releasing leptin evenly for 14 days, to manipulate their perceived body condition, or a placebo pellet. In the first experiment where females were implanted when feeding their first brood offspring we found, surprisingly, that placebo treated females were more likely to initiate a second brood compared to leptin treated females. Only one second brood fledged two chicks while five were deserted late in the incubation stage or when the first egg hatched. No difference was found in female or male return rate or in recruitment rate of fledglings of the first brood, possibly due to the desertion of the second broods. In our study population, where there is selection for early egg laying, earlier timing of reproduction might be hampered by food availability and thus nutritional state of the female before egg laying. We therefore implanted similar leptin pellets three weeks before the expected start of egg laying in an attempt to manipulate the laying dates of first clutches. However, leptin treated females did not initiate egg laying earlier compared to placebo treated females, suggesting that other variables than the perceived body condition play a major role in the timing of reproduction. Also, leptin treatment did not affect body mass, basal metabolic rate or feeding rates in captive females. Manipulating life history decisions using experimental protocols which do not alter individuals' energy balance are crucial in understanding the trade-off between costs and benefits of life history decisions.     

Latitudinal variation in parental energy expenditure during brood rearing in the great tit

The variation in time and energy allocation of female great tits, Parus major, was studied in five different European populations across a latitudinal gradient. Daily energy expenditure (DEE) was measured in females tending 12-day-old broods. The number of daylight hours used by the parents to collect food for the brood increased with latitude, while DEE and feeding rate per brood tended to level off with latitude. Individual variation in DEE could be explained by variation in ambient temperature (–), the duration of activity period (+) and area, but not by brood size, female body mass, brood mass or feeding rate. When the effect of ambient temperature and the duration of the activity period on the day of energy expenditure measurements were controlled for, female DEE still tended to level off with latitude. Temperature and activity alone can thus not explain the observed pattern. The present study suggests that parents at southern latitudes may be under a time constraint and do not increase energy expenditure because they have no more daylight hours available for foraging, while birds at northern latitudes may be under an energy constraint because they do not make full use of the long daylight period available. Received: 25 May 1999 / Accepted: 08 September 1999     

Pre-breeding energy requirements: thyroid hormone, metabolism and the timing of reproduction in house sparrows Passer domesticus

We measured thyroid hormone (T3) levels and energy expenditure of pre-breeding house sparrows Passer domesticus in relation to the timing of breeding and reproductive success. The onset of reproduction was synchronised in two waves, separated by a three-week interval. On an annual basis, early breeders (birds that bred for the first time during the first wave) made significantly more breeding attempts, laid significantly more egg and raised 2.3 times more chicks to fledging than late breeders (birds that bred for the first time during the second wave). By the end of March, about one month before the first egg was laid in the population, plasma titres of testosterone in males and estradiol in females were still low and did not differ between early and late breeders. However, birds that subsequently bred early had higher titres of plasma triiodothyronine (T3) than birds that started to breed late. We show for the first time in a free-living bird population that Basal Metabolic Rate (BMR) is positively correlated with the plasma titre of T3. Differences in plasma T3 accounted for 48% of the inter-individual variation in BMR. Elevated T3 levels indicate that energy requirements increase prior to breeding. Although early breeding appears to be advantageous in terms of the number of offspring raised on an annual basis, the increased energy requirements prior to breeding are thought to delay the onset of reproduction in those birds that cannot afford the additional energy expenditure early in the season.     

Hormone and metabolite changes associated with extended breeding fasts in male northern elephant seals (Mirounga angustirostris)

We measured metabolic hormones and several key metabolites in breeding adult male northern elephant seals to examine the regulation of fuel metabolism during extended natural fasts of over 3 months associated with high levels of energy expenditure. Males were sampled twice, early and late in the fast, losing an average of 23% of body mass and 47% of adipose stores between measurements. Males exhibited metabolic homeostasis over the breeding fast with no changes in glucose, non-esterified fatty acids, or blood urea nitrogen. Ketoacids increased over the fast but were very low when compared to other fasting species. Changes within individuals in total triiodothyronine (tT₃) were positively related to daily energy expenditure (DEE) and protein catabolism. Differences in levels of thyroid hormones relative to that observed in weaned pups and females suggest a greater deiodination of T₄ to support the high DEE of breeding males. Relative levels of leptin and ghrelin were consistent with the suppression of appetite but a significant reduction in growth hormone across the fast was contrary to expectation in fasting mammals. The lack of the increase in cortisol during fasting found in conspecific weaned pups and lactating females may contribute to the ability of breeding males to spare protein despite high levels of energy expenditure. Together these findings reveal significant differences with conspecifics under varying nutrient demands, suggesting metabolic adaptation to extended high energy fasts.     

Plasticity of brood patch development and its influence on incubation periods in the yellow-eyed penguin Megadyptes antipodes: an experimental approach

In many bird species, eggs laid late in the laying period hatch after a shorter incubation period than early-laid eggs. However, the mechanisms that explain these seasonal declines in incubation periods among clutches remain poorly understood. In this study we investigated the plasticity of brood patch development during incubation in yellow-eyed penguins Megadyptes antipodes and established whether differences exist in brood patch formation among early, mean and late-breeding penguins. We also examined whether brood patch development was influenced by sex and age of birds. We then placed an artificial egg in nests a few days prior to egg laying to investigate whether the presence of an egg influences brood patch development and whether an advanced brood patch development at the time of egg laying causes declines in incubation periods. Initial brood patch width on the day the first egg was laid was dependent on sex and age, while the development of brood patch width after first egg laying was slower in early-laying birds than in mean- and late-laying birds. Initial brood patch temperature as well as temperature throughout incubation was largely dependent upon sex, whereby males had higher brood patch surface temperatures than females. Placement of an artificial egg in nests stimulated successfully brood patch development in manipulated birds, so that by the time they laid their own first egg, their brood patches were wider and had higher temperatures than those of control birds. Moreover, incubation periods of first eggs from manipulated nests were significantly shorter (43.5 days) than were those from control nests (47.3 days). Thus, variation in brood patch development and related differences in incubation temperature during early incubation could contribute to seasonal declines in incubation periods.     

Energetics of the annual cycle of Dippers Cinclus cinclus

Time-activity budgets and energy expenditure of Dippers Cinclus cinclus were studied in all months of the year and for every stage of the annual cycle. The commonest daytime activity was feeding (54%), then resting (43%) and flying (4%). On a 24-hr day basis the most marked changes in activity followed from changing daylengths. DEE (Daily Energy Expenditure), derived from time-activity budgets through the year and laboratory estimates of metabolism, averaged 201 kj d ^-1 in females and 228 kj d ^-1 in the larger males. Over a more restricted range of circumstances, direct estimates of DEE obtained from 77 Dippers using the doubly labelled water technique averaged 205 ± 43 kj d^-1 and 251 ± 55 kj d^-1 in females and males, respectively. Overall, the correspondence between these largely independent estimates of energy expenditure was reasonably close. DEE was highest during breeding (laying-females; rearing-males) and in late winter for both sexes. The lowest energy expenditures occurred during moult, amongst juveniles and in early winter. Incubating females and mate-guarding males also had low energy costs. Across all stages of the annual cycle body size, activity patterns, ambient temperature and river flow had significant effects on energy expenditure. The rate at which food was gathered to meet these changing energy demands varied widely. While some of this variation was imposed by a seasonal environment, it was also likely to reflect adaptive shifts in rates of food gathering, in some cases consequent upon the changing fitness benefits of various non-feeding activities.     

Selection for synchronous breeding in the European starling

Colonial birds often demonstrate considerable breeding synchrony. In southern Sweden the semi-colonial European starling initiated the vast majority of clutches within one week. Laying dates were positively skewed so that many birds initiated clutches at similar dates early in the season. Breeding was further synchronised by a particularly strong clutch-size reduction equivalent to one third of an egg per day during the first part of the breeding season. The decline in clutch size with season also held true for separate age-classes of females, for individual females laying at different times at different years and for individual females laying at different times the same year. Trends in breeding success during nestling rearing were unlikely to explain the high degree of breeding synchrony or the seasonal decline in clutch size; nestling survival and growth were weakly related or unrelated to reproductive timing. In contrast recruitment success of fledged offspring declined sharply with season. Even within the synchronous laying period, defined as clutches initiated during the first week each year, local recruitment success declined. It is suggested that the early seasonal decline is caused by selection for synchronous fledging permitting the immediate formation of flocks after fledging, whereas the late seasonal trends may be caused by either population differences in female quality or deteriorating conditions for raising young.     

Maternal androgens in the pied flycatcher: timing of breeding and within-female consistency

Maternal hormones can have substantial phenotypic effects in the progeny of many vertebrates. It has been proposed that mothers adaptively adjust hormone levels experienced by particular young to optimize their reproductive output. In birds, systematic variation in egg hormone levels has been related to different female reproductive strategies. Because in many bird species prospects of the offspring change seasonally and with brood number, strategic adjustment of yolk androgen levels would be expected. To test this idea, we induced pied flycatcher (Ficedula hypoleuca) females to nest twice during the same season by removing their first clutches shortly after clutch completion. We collected eggs of first and replacement clutches to measure yolk concentrations of androstenedione (A4) and testosterone (T) and captured the females that laid these clutches for phenotypic measurements. Although average egg androgen levels were remarkably consistent within females, hormone patterns differed considerably between first and replacement clutches. Eggs of replacement clutches were heavier with larger yolks compared to first clutches, but they contained on average lower levels of androgens. Within clutches, androgen concentration increased over the laying sequence in the first clutch, but decreased or remained more constant over the laying sequence in the replacement clutch. Mean yolk T, but not A4 levels, were negatively associated with laying date for both breeding attempts. Moreover, females in good body condition produced eggs containing lower levels of androgens than females in poor condition. Our results are consistent with the idea that differences in yolk androgen levels may be one mechanism underlying seasonal variation in reproductive success and it is possible that changes in egg androgen patterns may reflect a change in female reproductive strategy. High within-female consistency also highlights the possibility that there may be some underlying genetic variation in yolk androgen levels.     

Mate provisioning, nutritional requirements for egg production, and primary reproductive effort of female Common Terns Sterna hirundo

We assessed the nutritional importance of mate provisioning to females during egg production and its effects on clutch parameters (egg size, length of the laying period) in Common Terns Sterna hirundo: (1) we estimated the costs of egg production by modeling the daily protein, lipid, and energy requirements of laying females, and (2) compared these costs to both the amount, and the timing, of the male's contribution via mate provisioning. Net lipid, net protein, and gross energy requirements for a three‐egg clutch were estimated to be 5.4 g, 8.6 g, and 569 kJ respectively. Peak protein and lipid requirements occurred one (day ?1) and two (day ?2) days before laying, respectively. Peak energy requirement occurred on day ?1; a cost of 127% to 157% above maintenance. Variation in male provisioning effort (in terms of energy and nutrients delivered) paralleled variation in predicted female requirements for egg production at the level of individual pairs. Males delivered protein in excess of the female's requirements on all days investigated. Male lipid delivery accounted for 45% of female net requirement on the day when demand was greatest (day ?2), but exceeded requirements on all other days. However, the proportion of the female's total energy budget (egg production, maintenance, and activity costs) that was supplied by her mate rose from an average of 29% on day ?2 to 76% during the interval between second and third eggs. Paradoxically, females that were fed at higher rates during the interval between first and second eggs produced clutches with lower total volumes, smaller last‐laid eggs, and clutches with a greater egg‐size hierarchy than conspecifics receiving less food from their mates. Also, females fed at higher rates during the interval between second and third eggs took longer to produce their clutch. These negative relationships between mate provisioning and clutch parameters contrast with previous studies in this and other species.     

Reproductive performance changes with age and laying experience in the Red-legged Partridge Alectoris rufa

Most studies of birds have reported positive effects of ageing on reproductive traits during the first years of life followed by a decline. We examined the effects of age and laying experience, controlling for confounding factors such as food availability/quality and timing of pair formation in a captive population of Red-legged Partridge Alectoris rufa . We found that all reproductive traits, except for the period between eggs, were highly repeatable within females both within and between years. The repeatability of egg mass was lower than that of clutch size, laying onset or chick traits other than the mass and size increase rates, perhaps suggesting a lower heritability value. As females aged, they showed a tendency to produce marginally lighter eggs (1.3%) and larger clutches, although this latter trend was not significant. We suggest that the senescence process implies the involvement of hormonal changes that could affect the reduction of egg mass (e.g. low levels of oestradiol). Younger females tended to start laying later than older females, which could be due to a low responsiveness to photoperiodic stimulation of the reproductive system, which would indicate a delay in gametogenesis and steroidogenesis. The mass of last-laid eggs decreased with age of the female. We suggest that physiological changes through the laying sequence could vary with age. Experienced breeders laid larger clutches linked with the advancement of laying onset. For a given age, females with more experience fared better than those with less experience.     

Female variation in allocation of steroid hormones,antioxidants and fatty acids: a multilevel analysis in a wild passerine bird

The environment where an embryo develops can be influenced by components of maternal origin, which can shape offspring phenotypes and therefore maternal fitness. In birds that produce more than one egg per clutch, females differ in the concentration of components they allocate into the yolk along the laying sequence. However, identification of processes that shape female yolk allocation and thus offspring phenotype still remains a major challenge within evolutionary ecology. A way to increase our understanding is by acknowledging that allocation patterns can differ depending on the level of analysis, such as the population versus the among‐female (within‐population) level. We employed mixed models to analyze at both levels the variation in allocation along the laying sequence of four steroid hormones, three antioxidants, and four groups of fatty acids present in the egg yolks of wild great tits Parus major. We also quantified repeatabilities for each component to study female consistency. At a population level, the concentrations/proportions of five yolk components varied along the laying sequence, implying that the developmental environment is different for offspring developing in first versus last eggs. Females varied substantially in the mean allocation of components and in their plasticity along the laying sequence. For most components, these two parameters were negatively correlated. Females were also remarkably repeatable in their allocation. Overall, our data emphasize the need to account for female variation in yolk allocation along the laying sequence at multiple levels, as variation at a population level is underpinned by different individual patterns. Our findings also highlight the importance of considering both levels of analysis in future studies investigating the causes and fitness consequences of yolk compounds. Finally, our results on female repeatability confirm that analyzing one egg per nest is a suitable way to address the consequences of yolk resource deposition for the offspring.     

Two eggs,two different constraints: a potential explanation for the puzzling intraclutch egg size dimorphism in Eudyptes penguins

Phenotypic plasticity and phenotypic stability are major components of the adaptive evolution of organisms to environmental variation. The invariant two-egg clutch size of Eudyptes penguins has recently been proposed to be a unique example of a maladaptive phenotypic stability, while their egg mass is a plastic trait. We tested whether this phenotypic plasticity during reproduction might result from constraints imposed by migration (migratory carry-over effect) and breeding (due to the depletion of female body reserves). For the first time, we examined whether these constraints differ between eggs within clutches and between egg components (yolk and albumen). The interval between colony return and clutch initiation positively influenced the yolk mass, the albumen mass, and the subsequent total egg mass of first-laid eggs. This time interval had only a slight negative influence on the yolk mass of second-laid eggs and no influence on their albumen and subsequent total masses. For both eggs, female body mass at laying positively influenced albumen and total egg masses. Female investment into the entire clutch was not related to the time in the colony before laying but increased with female body mass. These novel results suggest that the unique intraclutch egg size dimorphism exhibited in Eudyptes penguins, with first-laid eggs being consistently smaller than second-laid eggs, might be due to a combination of constraints: a migratory carry-over effect on the first-laid egg and a body reserve depletion effect on the second-laid egg. Both these constraints might explain why the timing of reproduction, especially egg formation, is narrow in migratory capital breeders.     

Geographic variation in reproductive traits and trade-offs between size and number of eggs of the Chinese cobra (Naja atra)

We collected gravid Chinese cobras (Naja atra) from one island (Dinghai) and three mainland (Yiwu, Lishui and Quanzhou) populations in south‐eastern China to study geographical variation in female reproductive traits and the trade‐off between the size and number of eggs. We then conducted an common experiment on cobras from two of the four populations to further identify factors contributing to the observed trade‐offs. The mean size (snout–vent length) of the smallest five reproductive females increased with increasing latitude. Oviposition occurred between late June and early August, with females from the warmer localities laying eggs earlier than those from the colder localities. Maternal size was a major determinant of the reproductive investment in all populations, with larger females producing not only more but also larger eggs. Clutch size was more variable than egg size within and among populations. The observed geographical variation in clutch size, egg size, clutch mass and post‐oviposition body condition was not a simple consequence of variation in maternal size among populations, because interpopulation differences in these traits were still evident when the influence of maternal size was removed. The upper limit to reproductive investment was more likely to be set by the space availability in the island population, but by the resource availability in the three mainland populations. Trade‐offs between size and number of eggs were detected in all populations, with females that had larger clutches for their size having smaller eggs. Egg size at any given level of relative fecundity differed among populations, primarily because of interpopulation differences in the resource availability rather than the space availability. Except for the timing date of oviposition and the mean size of the smallest five reproductive females, all other examined traits did not vary in a geographically continuous trend. The common garden experiment, which standardized environmental factors, synchronized the timing date of oviposition, but it did not modify the conclusion drawn from the gravid females collected from the field. The observed geographical variation in the female reproductive traits could be attributed to the consequence of the effects of either proximate or ultimate factors. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 27–40.     

Causes and consequences of egg mass variation between and within blue tit clutches

Causes of egg-size variation between and within clutches were studied in clutches of the blue tit ( Parus caeruleus L.). We measured the mass of each egg in the laying sequence in unmanipulated clutches, in clutches of parents experimentally supplied with extra food before egg-laying, and in clutches of parents supplemented with extra food after the start of egg-laying. Hatchlings were weighed at an age of two days and their mass was found to be positively related to egg mass. No general trend of decreasing or increasing egg mass was found within the laying sequence. Females provided with extra food before egg-laying laid clutches with significantly less variation in egg mass than did control females. The reason for this was that the first-laid egg of unmanipulated females was lighter than the rest of the eggs in the clutch. This pattern disappeared in clutches of females receiving extra food. Thus, the reduction in egg mass variation among clutches of foodsupplemented females depended on an ability of these females, in contrast to control females, to lay a first egg of the same mass as the rest of the clutch. Eggs laid after the initiation of incubation were significantly heavier than equivalent eggs in those clutches where incubation started after clutch completion. The difference was small, however, and the adaptive significance of the finding is questionable. We argue that intra-clutch variation in egg mass is connected with greater fitness consequences than in inter-clutch variation. Furthermore, our results indicate that energetical constraints on the laying female are more important as a cause of the observed intra-clutch variation in egg mass than are adaptive responses to the environment.     

A model of clutch size determination in birds

Summary This paper tests the hypothesis that the timing and the size of bird clutches are determined by (1) a tendency to delay reproduction so that the female can store enough internal resources for laying a large clutch; (2) a tendency to breed early because survival of offspring late in the breeding season might be lower than early. These factors are approximated by linear functions in a model. The model predicts that there will be a correlation between early breeding females and those that have large clutches. This agrees with published data for several bird species. Predictions from the model are also made of clutch size and the impact on final clutch-size if one egg is removed or added during the laying. These predictions were compared with data from a crow (Corvus cornix) population and were supported if it is assumed that the formation of an egg is a relatively costly process. More precisely, the model is supported if the formation of an extra egg requires a delay of about four or five days before laying can begin.     

Maternal effects influence the sexual behavior of sons and daughters in the zebra finch

Individual differences in sexual behavior have received much attention by evolutionary biologists, but relatively little is known about the proximate causes of this variation. We studied the quantitative genetics of male and female sexual behavior of captive zebra finches and found surprisingly strong maternal effects (differing between individual mothers) on the aggressiveness and song rate of sons and on the daughters' mating preferences for these male traits. We also found that daughters differed in their choosiness during mate-choice experiments depending on whether they originated from eggs produced early or late within the laying sequence of a clutch. Because this effect of laying order occurred independently of hatching order in cross-fostered broods, it must have been caused by consistent within-mother variation in maternal effects transmitted through the egg. Our findings raise the question whether these maternal effects might represent strategic programming of offspring behavior in response to the environment experienced by mothers or whether they are merely nonadaptive byproducts of developmental processes.