Variable effects of the Hen Flea Ceratophyllus gallinae on the breeding success of the Great Tit Parus major in relation to weather conditions

We investigated the effect of an ectoparasite, the Hen Flea Ceratophyllus gallinae (Siphonaptera), on the reproductive success of the Great Tit Parus major, a hole-nesting passerine bird. Parasite levels of cleaned nesting boxes were experimentally manipulated during the egg-laying period of the birds either by spraying nests with pyrethrin or by introducing c. 100 adult fleas, while other nests were left as controls. Only in 2 out of 5 years did hen fleas significantly affect reproductive success. Nestlings from parasitized nests showed reduced body-mass, tarsus-length and condition, whereas wing-length did not differ between nestlings reared with or without fleas. The impact of the fleas on the growth performance of host nestlings was more noticeable in years with lower temperatures and higher precipitation.     

Green plants in starling nests: effects on nestlings

European starlings, Sturnus vulgaris, intermingle fresh herbs, especially species rich in volatile compounds, with their otherwise dry nest material. In this field study we investigated whether these herbs reduce ectoparasites and thereby protect nestlings (the nest protection hypothesis). We also considered whether volatile compounds in herbs improve the condition of nestlings (the drug hypothesis). As measures of condition we used body mass, haematocrit levels and immunological parameters. We replaced 148 natural starling nests with artificial ones: half contained herbs and half (controls) contained grass. The ectoparasite loads (mites, lice, fleas) in herb and control nests were indistinguishable. However, nestlings in herb nests weighed more and had higher haematocrit levels at fledging than nestlings in control nests. Fledging success was similar in herb and control nests, but more yearlings from herb nests were identified in the colony the year after hatching. The response of the immune system when challenged with phytohaemagglutinin did not differ in nestlings from herb and control nests. Nestlings from herb nests had more basophils and fewer lymphocytes in their blood than those from control nests, while the eosinophil and heterophil counts did not differ. We conclude that herbs do not reduce the number of ectoparasites, but they improve the condition of nestlings, perhaps by stimulating elements of the immune system that help them to cope better with the harmful activities of ectoparasites. Copyright 2000 The Association for the Study of Animal Behaviour.     

Begging, food provisioning, and nestling competition in great tit broods infested with ectoparasites

Ectoparasites are a ubiquitous environmental component of breedingbirds, and it has repeatedly been shown that hematoph-agousectoparasites such as fleas and mites reduce the quality andnumber of offspring of bird hosts, thereby lowering the valueof a current brood. Selection acting on the hosts will favorphysiological and behavioral responses that will reduce theparasites' impact. However, the results of the few bird studiesthat addressed the question of whether parasitism leads to ahigher rate of food provisioning are equivocal, and the beggingresponse to infestation has rarely been quantified. A changein begging activity and parental rate of food provisioning couldbe predicted in either direction: parents could reduce theirinvestment in the brood in order to invest more in future broods,or they could increase their investment in order to compensatefor the parasites' effect on the current brood. Since the nestlingsare weakened by the ectoparasites they may beg less, but onthe other hand they may beg more in order to obtain more food.In this study we show experimentally that (1) hen fleas (Ceratophyllusgallinae) reduce the body mass and size of great tit (Parusmajor) nestlings, (2) nestlings of parasitized broods more thandouble their begging rate, (3) the male parents increase thefrequency of feeding trips by over 50%, (4) the females do notadjust feeding rate to the lowered nutritional state of nestlings,and (5) food competition among siblings of parasitized broodsis increased. Ultimately the difference in the parental feedingresponse may be understood as the result of a sex-related differencein the trade-off of i0vesting in current versus future broods.     

Hatching asynchrony and brood reduction influence immune response in Common Kestrel Falco tinnunculus nestlings

The onset of incubation before the end of laying imposes asynchrony at hatching and, therefore, a size hierarchy in the brood. It has been argued that hatching asynchrony might be a strategy to improve reproductive output in terms of quality or quantity of offspring. However, little is known about the mediating effect of hatching asynchrony on offspring quality when brood reduction occurs. Here, we investigate the relationship between phenotypic quality and hatching asynchrony in Common Kestrel Falco tinnunculus nestlings in Spain. Hatching asynchrony did not increase breeding success or nestling quality. Furthermore, hatching asynchrony and brood reduction had different effects on nestlings’ phytohaematogglutinin (PHA)‐mediated immune response and nestling growth. In asynchronous and reduced broods (in which at least one nestling died), nestlings showed a stronger PHA‐mediated immune response and tended to have a smaller body size compared with nestlings raised in synchronous and reduced broods. When brood reduction occurred in broods hatched synchronously, there was no effect on nestling size, but nestlings had a relatively poor PHA‐mediated immune response compared with nestlings raised in asynchronous and reduced broods. We suggest that resources for growth can be directed to immune function only in asynchronously hatched broods, resulting in improved nestling quality, as suggested by their immune response. We also found that males produced a greater PHA‐mediated immune response than females only in brood‐reduced nests without any effect on nestling size or condition, suggesting that females may trade off immune activities and body condition, size or weight. Overall, our results suggest that hatching pattern and brood reduction may mediate resource allocation to different fitness traits. They also highlight that the resolution of immune‐related trade‐offs when brood reduction occurs may differ between male and female nestlings.     

Behavioural responses to ectoparasites: time-budget adjustments and what matters to Blue Tits Parus caeruleus infested by fleas

Blue Tit nests are often heavily infested by fleas, which feed on the incubating female and the nestlings. Depending on habitat quality, the drawing of blood by fleas reduces offspring quality, or it is compensated by an increase in food provisioning by the adults and may reduce their future reproduction. Given these fitness costs, tits are expected to have evolved behavioural responses enabling them to remove, destroy or minimize the contact with fleas. To identify these traits, we video-recorded the changes in frequency and duration of the hosts' potential anti-flea behavioural defences in nests experimentally infested with low and high flea densities. We also investigated whether flea load affected the number of male feeds delivered to incubating females, and whether the parents increased their rate of food provisioning to the nestlings equally at high flea density. Flea density significantly affected the nest sanitation and sleeping behaviour of Blue Tit females but had no significant effect on grooming. Female Blue Tits increased the frequency but decreased the duration of bouts of these behavioural traits, and hence their time-budgets, based on per hour duration of behaviour, were not significantly affected by flea density. High flea density reduced nestling weight at the early nestling stage but these costs were fully compensated by an increase in female feeding effort. Males did not increase their frequency of food provisioning to incubating females nor to nestlings in heavily infested nests. The results are discussed in the light of parasite-mediated selection on host behaviour and the reciprocal host selection on flea life-history and behavioural traits.     

Experimental addition of greenery reduces flea loads in nests of a non-greenery using species, the tree swallow Tachycineta bicolor

Several bird species, including cavity-nesters such as European starlings Sturnus vulgaris , add to their nests green sprigs of plants such as yarrow Achillea millefolium that are rich in volatile compounds. In this field study on another cavity-nester, tree swallows Tachycineta bicolor , we tested whether yarrow reduced ectoparasite loads (the nest protection hypothesis), stimulated nestling immune systems (the drug hypothesis), or had other consequences for nestling growth or parental reproductive success (predicted by both preceding hypotheses). Tree swallows do not naturally add greenery to their nests, and thus offer several advantages in testing for effects of greenery independent of other potentially confounding explanations for the behaviour. We placed fresh yarrow in 23 swallow nests on the day the first egg was laid, replenishing every two days until clutch completion (=three times), and at 44 control nests, nesting material was simply touched. At 12 days of age, we measured nestling body size and mass, and took blood smears to do differential white blood cell counts. We subsequently determined the number and proportion of young fledging from nests and the number of fleas remaining after fledging. Higher humidity was associated with higher flea numbers whereas number of feathers in the nest was not. Our most significant finding was that an average of 773 fleas Ceratophyllus idius was found in control nests, versus 419 in yarrow nests. Possibly, parents compensate for blood that nestlings lose to ectoparasites by increasing food delivery, because we detected no differences between treatments in nestling mass, nestling leukocyte profiles, or proportion of young fledging, or relative to flea numbers. Our results provide no support for the drug hypothesis and strong support for the nest protection hypothesis.     

Sympatric and allopatric combinations of hen fleas and great tits: a test of the local adaptation hypothesis

To test whether allopatric nest parasites differ from sympatric ones in their effect on various life history traits of their avian host, I designed a cross-transfer experiment in which hen fleas (Ceratophyllus gallinae) were exchanged between great tit (Parus major) nests in two geographically widely separated areas. In neither of the areas did allopatric fleas influence body mass, tarsus length, wing length, duration of nestling period or mortality in great tit nestlings more severely than did sympatric fleas. Duration of incubation was also similar among females independent of experimental treatment. This lack of difference between allopatric and sympatric fleas is hypothesized to reflect the comparative harmlessness of hen fleas for their hosts.     

Nest predation and nestling growth rate of two lark species in the Negev Desert, Israel

Clutch-size, nestling growth and predation rates on eggs and nestlings in two sympatric alaudids, the Crested Lark Galerida cristata and the Desert Lark Ammomanes deserti, were studied in the Negev desert of Israel. Logistic growth constants were similar for both species, and experimental brood reduction did not accelerate the growth rate of their nestlings. It is suggested that the predation rate on nests of both species has been sufficiently high to select for maximal growth rate of nestlings. Overall probability of predation was 0.85 on a Desert Lark nest and 0.76 on a Crested Lark nest. The smaller clutch-size of the Desert Lark (3.7) may be an adaptation to a higher predation risk, as compared to that of the Crested Lark (4.6) which is more vigilant and nests in more concealed sites.     

Interacting effects of aromatic plants and female age on nest-dwelling ectoparasites and blood-sucking flies in avian nests

Some avian species incorporate aromatic plants to their nests. The "nest protection hypothesis", which posits that volatile secondary compounds contained in these plants may have antiparasite properties, has not received full support. All previous tests of this hypothesis have only considered effects on nest-dwelling ectoparasites, but not on blood-sucking flies. The "drug hypothesis" posits that aromatic plants may stimulate nestling immune system, development, or condition. We tested these hypotheses experimentally in wild blue tits, Cyanistes caeruleus, a species that adds aromatic plants to their nests. We supplemented aromatic plants to half of a sample of nests, while adding grass to the other half of nests. We quantified abundance of two groups of blood-sucking flies (blackflies and biting midges) at two different stages of the reproductive period, and abundance of three nest-dwelling ectoparasites (fleas, mites, and blowflies). Experimental supplementation of aromatic plants reduced abundance of fleas only in nests of yearling females and not in nests of older females. Blackflies and biting midges were both more numerous in nests of yearling females than in nests of older females. Mass of aromatic plants added by females was negatively related with abundance of fleas in control nests but not in experimental nests supplied with aromatic plants. Mass of plants added by females was also positively related with abundance of blackflies during the nestling stage. Finally, aromatic plants did not affect nestling growth or immune responses. We conclude that several factors such as female experience and their ability to add plants to the nest interact to explain effects of aromatic plants on different parasites.     

Impacts of parasites in early life: contrasting effects on juvenile growth for different family members

Parasitism experienced early in ontogeny can have a major impact on host growth, development and future fitness, but whether siblings are affected equally by parasitism is poorly understood. In birds, hatching asynchrony induced by hormonal or behavioural mechanisms largely under parental control might predispose young to respond to infection in different ways. Here we show that parasites can have different consequences for offspring depending on their position in the family hierarchy. We experimentally treated European Shag (Phalacrocorax aristoteli) nestlings with the broad-spectrum anti-parasite drug ivermectin and compared their growth rates with nestlings from control broods. Average growth rates measured over the period of linear growth (10 days to 30 days of age) and survival did not differ for nestlings from treated and control broods. However, when considering individuals within broods, parasite treatment reversed the patterns of growth for individual family members: last-hatched nestlings grew significantly slower than their siblings in control nests but grew faster in treated nests. This was at the expense of their earlier-hatched brood-mates, who showed an overall growth rate reduction relative to last-hatched nestlings in treated nests. These results highlight the importance of exploring individual variation in the costs of infection and suggest that parasites could be a key factor modulating within-family dynamics, sibling competition and developmental trajectories from an early age.     

Hatching asynchrony, nestling competition, and the cost of interspecific brood parasitism

All parental hosts of heterospecific brood parasites must paythe cost of rearing non-kin. Previous research on nest parasitismby brown-headed cowbirds (Molothrus ater) concluded that competitivesuperiority of the typically more intensively begging and largercowbird chick leads to preferential feeding by foster parentsand causes a reduction in the hosts' own brood. The larger sizeof cowbird nestlings can be the result of at least two causes:(1) cowbirds preferentially parasitize species with smallernestlings and lower growth rates; and/or (2) cowbirds hatchearlier than hosts. I estimated the cost of cowbird parasitismfor each of 29 species by calculating the difference betweenhosts' published brood sizes in nonparasitized and parasitizednests and using clutch size to standardize values. In this analysis,greater incubation length and lower adult mass, surrogate measuresof the hatching asynchrony and size difference between parasiteand hosts, were both related to greater costs of cowbird parasitismwithout bias owing to phylogeny. To establish causality, I manipulatedclutch contents of eastern phoebes (Sayornis phoebe) and examinedwhether earlier hatching by a single cowbird or phoebe egg reducesthe size of the rest of the original host brood. As predicted,greater hatching asynchrony increased the proportion of theoriginal phoebe brood that was lost. This measure of the costof parasitism was partially owing to increased hatching failureof the original eggs in asynchronous broods but was not at allrelated to the size differences of older and younger conspecificnestmates. However, proportional brood loss owing to an earlierhatching conspecific was consistently smaller than brood lossowing to asynchronous cowbirds in both naturally and experimentallyparasitized phoebe nests. These results imply that althoughhatching asynchrony is an important cause of the reduction ofhost broods in parasitized clutches, competitive features ofcowbird nestlings remain necessary to explain the full extentof hosts' reproductive costs caused by interspecific brood parasitism.     

Influence of hatching order on growth rate and resting metabolism of kestrel nestlings

Hatching asynchrony in altricial birds may result in a competitive disadvantage for the youngest nestlings compared to older siblings. We studied the effects of a size hierarchy on the growth rate of Eurasian kestrels Falco tinnunculus chicks in nests with and without access to supplemented food in western Finland. Body mass stopped increasing on the 19th day after hatching while body size, estimated by a combination of bone and feather lengths continued to increase at least until fledging at 26 days. Body condition, reflecting muscle and fat, did not change markedly during the growth period from the 12th day to fledging. Body temperature and resting metabolism were usually lower in nestlings 12 days old than in nestlings at fledging. Growth of body mass, size and condition, and resting metabolism were delayed in last-hatched nestlings aged 19 days. Just before fledging, last-hatched nestlings attained a similar body mass and size, and had a similar resting metabolism to those of older siblings. At fledging, only in nests without access to supplemented food was the body condition of last-hatched chicks lower than that of its siblings, but in nests with access to supplemented food no such difference was detected. Our results highlight that the level of lipids in the last-hatched nestling can be affected by the food restriction imposed by hatching order.     

A parasite-induced maternal effect can reduce survival times of fleas feeding on great tit nestlings

Parents can increase their reproductive success by assisting their neonate offspring in parasite defence. In birds, parental tactics include post-hatching parental responses such as increased parental care and pre-hatching maternal effects such as the transfer of maternal antibodies via the egg. These parasite-induced parental responses are known to reduce the effects of parasites on offspring, but their costs for the parasite are largely unknown. In two separate experiments on great tits Parus major we assessed these costs for hen fleas Ceratophyllus gallinae . Half of the parents where exposed to fleas during egg-laying to induce the parental response, while control nests were left flea-free. In experiment 1 parents raised their own young and we measured the effect of combined pre- and post-hatching parental effects, while in experiment 2 a cross-foster design allowed us to assess the effects of pre-hatching maternal effects alone. In both experiments we let fleas take a blood meal on nestlings from either flea-exposed or unexposed parents. We then measured flea-feeding duration, the quantity of extracted blood, and the fleas' subsequent survival time. We found in both experiments that on the largest nestlings of a brood flea survival was significantly reduced by the parental effects, whereas on the smaller nestlings it was independent of parental effects. The pre- and post-hatching parental responses did neither affect duration nor size of a flea blood meal. These results suggest first that the pre-hatching maternal effects, i.e. the substances transferred to the nestling via the egg, have the potential to harm fleas without reducing flea feeding capacity, and second that the strength of the maternal response varies between the nestlings, either because maternal products are unequally distributed among eggs within a clutch, or because large nestlings can build up a response that enhances the effect of the maternal products.     

Reduced immunocompetence of nestlings in replacement clutches of the European magpie (Pica pica)

Laying date is one of the most important determinants of reproductive success and recruitment probability in birds. Late breeders usually fledge fewer chicks than individuals with earlier breeding dates, and fledglings produced late in the season have high mortality rates. Food availability and nestling mass have been evoked as the principal mechanistic links between laying date and offspring survival. Here we suggest that another factor may actually account for the difference in survival rate between early and late offspring: immunocompetence. We predicted that nestlings produced later in the season or in replacement clutches should have lower immune responses when challenged with an antigen, than early nestlings or nestlings produced in first clutches. This hypothesis was tested in a population of magpies (Pica pica), in which we experimentally induced breeding failure in a group of nests and compared the immune response of nestlings in replacement clutches with the immune response of first clutch nestlings. Cellular immune response, as measured by wing web swelling (a correlate of T-lymphocyte production after injection of phytohaemagglutinin-P), significantly decreased with hatching date and was significantly lower in nestlings of replacement clutches. Furthermore, coefficients of intraclutch variation in immune response were higher in nestlings of replacement clutches. This experiment demonstrates an inverse relationship between immune responsiveness and breeding date, and reduced recruitment probability of late nestlings may be a direct consequence of their inability to cope with parasites.     

How selfish is a cowbird nestling?

Brood-parasitic young are reared in the nests of different species and can derive no genetic benefit from the survival of host offspring. However, although the nestlings of many parasitic cuckoo and honeyguide species routinely kill host young soon after hatching, nestling brown-headed cowbirds, Molothrus ater, tolerate host offspring and are commonly reared alongside them for at least part of the nestling period. I used comparative analyses of data from the literature to investigate whether brown-headed cowbird nestlings gain direct benefits by allowing host young to live. The brown-headed cowbird (44 g) parasitizes many passerines (adult mass range about 5-90 g) and the likelihood that host young survive to fledge from parasitized nests varies between species. In common with previous work, I found that host offspring mortality was highest in species whose offspring were relatively small compared with the cowbird nestling. Furthermore, cowbird nestlings were most likely to fledge when reared alongside host young of intermediate size. In these nests, one or two host young typically fledged as well. I suggest that cowbirds, and other host-tolerant brood parasites, could benefit from the presence of host nestlings through the assistance that host chicks offer in soliciting a higher provisioning rate, and that such benefits might outweigh the costs of having competition for food at each nest visit. Variation in this cost-benefit ratio could explain differences between brood parasite species in their tolerance of host young.     

The Process and Causes of Fledging in a Cavity-Nesting Passerine Bird, the House Wren (Troglodytes aedon)

Little is known about the process or causes of fledging or nest‐leaving in passerine birds because researchers can rarely predict when fledging will occur in a given nest. We used continuous videotaping of nests to both document the process of fledging in the house wren, Troglodytes aedon, a small, cavity‐nesting songbird, and test hypotheses as to what might cause fledging to begin. Fledging began any time from 14 to 19 d after hatching commenced. Slower‐developing broods fledged later than faster‐developing broods. Fledging typically began within 5 h of sunrise and over 80% of all nestlings fledged before noon. All nestlings fledged on the same day at 65% of nests and over two consecutive days in most other nests. We found no evidence that fledging was triggered by changes in parental behaviour. Parental rate of food delivery to nestlings did not decline during a 3‐h period leading up to the first fledging, nor was the rate of feeding just prior to the first fledging lower than the rate at the same time the day before. Moreover, parents did not slow the rate of food delivery to nests after part of the brood had fledged. Hatching is asynchronous in our study population which creates a marked age/size hierarchy within broods. At most nests, the first nestling to fledge was the most well‐developed nestling in the brood or nearly so (as measured by feather length). This suggests that fledging typically begins when the most well‐developed nestlings in the brood reach some threshold size. However, at about one‐fifth of nests, the first nestling to fledge was only moderate in size. At these nests, severe competition for food may have caused smaller, less competitive nestlings to fledge first to increase their access to food. We found no strong support for the suggestion that the oldest nestlings delay fledging until their least‐developed nestmate reaches some minimum size, although further experimental work on this question is warranted.     

Nest-site preference and maternal effects on offspring growth

Maternal preferences for oviposition sites are assumed to beadaptive, but offspring fitness is not always higher at preferredsites and, thus, further study of the selection pressures thatinfluence oviposition behavior is warranted. Among birds, predationis regarded as the primary agent of selection on nest-site microhabitatpreferences, but alternatives are rarely considered. We testedthe hypothesis that avian nest-site preferences are an adaptiveresponse to fitness costs imposed by variation in nest-sitemicroclimate. We documented that Chestnut-collared Longspurs(Calcarius ornatus) strongly preferred to orient nests towardsthe southeast and showed that this preference influenced microclimate:nests facing southeast had the highest midday temperatures.Yet, preferences were not adaptive because nestlings in nestswith the preferred orientation gained mass at a slower rate,had retarded skeletal growth, and reached a smaller final size.We experimentally tested this result by altering orientationof nests and confirmed, for the first time, that variation innestling growth was causally linked to variation in nest microclimatearising from nest-orientation preferences. Adults respondedto the high temperatures at preferred southeast-facing nestsby spending more time shading young from the sun, apparentlyattempting to ameliorate heat costs. This response, however,resulted in parents spending less time feeding young, potentiallyexplaining slower growth in these nests. Direct effects of highertemperatures may also play a role in slower growth. Althoughwe lack an explanation for this apparently maladaptive preference,these results demonstrate that nest-site choices of birds canyield fitness costs imposed by variation in nest microclimate.     

Nest size predicts the effect of food supplementation to magpie nestlings on their immunocompetence: an experimental test of nest size indicating parental ability

Post-mating sexually selected signals are expected to indicateparental quality. The good parent model assumes that expressionof the sexual character positively reflects parental ability,resulting in a potential link between the exaggeration of thecharacter and nestling-fitness traits. We tested this predictionin a population of a monogamous passerine, the magpie (Picapica), for which nest size is known to act as a post-matingsexually selected signal. We provided a food supplement to halfof the magpie nestlings in each nest, keeping the other halfas control nestlings. We found that food-supplemented nestlingsexperienced a significantly higher T-cell-mediated immune responseand a tendency to an increased condition index. In accordancewith the good parent model, we found that nest size was positivelyrelated to T-cell mediated immune response for control magpie,whereas this relationship was nonexistent in food-supplementednestlings. In addition, the difference in T-cell mediated immuneresponse between food-supplemented and control nestlings ofthe same nest was principally explained by nest size. Basedon our results, we discuss that magpie pairs with large nestsprovided their nestlings with higher quality food as comparedto pairs with smaller nests, nest size thereby being an indicatorof parental ability. To our knowledge, this is the first studyshowing a link between a post-mating sexually selected signaland nestling immunocompetence, a trait closely related to fitnessin birds.     

Behavioural responses to ectoparasites in pied flycatchers Ficedula hypoleuca: an experimental study

Nests of cavity‐nesting birds usually harbor some species of haematophagous ectoparasites that feed on the incubating adults and nestlings. Given the negative impact of ectoparasites on nestlings there will be selection on hosts to reduce parasite infestations through behavioural means. We have experimentally reduced the abundance of all ectoparasites in nests of pied flycatchers Ficedula hypoleuca to explore both whether there are changes in the frequency and duration of putative anti‐parasite behaviours by tending adults, as well as whether such anti‐parasite behaviours are able to compensate for the deleterious effects that parasites may have on nestlings. Heat treatment of nests substantially decreased the density of ectoparasites, and thereby positively affected nestling growth. The frequency and intensity of female grooming and nest sanitation behaviours during the incubation and nestling periods decreased as a consequence of the experimental reduction of ectoparasite infestation. Although nestlings begged more intensely in infested nests, the experiment had no significant effect on parental provisioning effort. Reduction of parasites resulted in larger nestlings shortly before fledging and increased fledging success. This study shows a clear effect of a complete natural nest ectoparasite fauna on parental behaviour at the nest and nestling growth in a cavity‐nesting bird. Although ectoparasites induce anti‐parasite behaviours in females, these behaviours are not able to fully remove parasite's deleterious effects on nestling growth and survival.     

Effects of food shortage and oversupply on energy utilization, histology, and function of the gut in nestling song thrushes (Turdus philomelos)

We measured food intake, digestive efficiency, body mass increments, resting metabolic rate (RMR), carcass fat content, size and histological structure of the gut, and the rate of intestinal brush border uptake of l-proline in song thrush (Turdus philomelos) nestlings subjected to food shortage or food surplus under laboratory conditions. We assigned nestlings between 3 and 7 d of age to one of the following treatments: (1) food restriction, which resulted in a slowed growth at the rate found in undernourished, wild nestlings; (2) overfeeding, which totally suppressed begging; and (3) intermediate feeding. Threefold differences in energy consumption caused fivefold differences in body mass increments of the nestlings. Despite this, body mass-corrected RMR and intestinal mass were not affected by the feeding regime. The energy content of fecal output was highest in food-restricted birds, while their carcass fat content was lowest among treatment groups. Intestinal uptake rates of l-proline were low in the overfed and intermediate-fed young but significantly increased in the food-restricted birds, who attempted to maximize their rates of growth and development within the restrictive limits set by feeding regime. We noted a marked decrease of intestinal villi height in overfed birds as compared to intermediate-fed and food-restricted nestlings. We conclude that song thrush nestlings are characterized by a limited plasticity of their developmental program, which prohibits overfed nestlings from significantly up-regulating their gut function to accommodate increased food intake. This suggests that they already grew at a rate close to their physiological maximum. We suggest two interpretations: (1) under natural conditions, song thrush nestlings do not face frequent, unpredictable fluctuations in food abundance that could select for developmental plasticity, or (2) strong selection for uniform adult phenotypes prevents flexible developmental trajectories, which would result in a diversity of adult phenotypes.