Fearfulness Affects Quail Maternal Care and Subsequent Offspring Development

Our study investigated relationships between a precocial bird’s fearfulness and maternal care, and the implication of maternal care as a vector for non-genomic transmission of fearfulness to chicks. We compared care given to chicks between two sets of female Japanese quail selected to present either high (LTI) or low fearfulness (STI). Chicks, from a broiler line, were adopted by these females following a sensitization procedure. Chicks’ fearfulness after separation from their mother was assessed by well-established procedures. LTIs took longer to present maternal responses, pecked chicks more during the first days post-hatch, presented impaired maternal vocal behaviour and were globally less active than STI females. Chicks mothered by LTIs presented more fearful reactions than did chicks mothered by STIs, supporting the hypothesis of a non-genetic maternal transmission of fearfulness. We suggest that the longer latencies required by LTIs to become maternal are a consequence of their greater fear of chicks, and that their lower general and vocal activity could be components of a heightened antipredatory strategy. We discuss the transmission of maternal fearfulness to fostered chicks, taking into account the possible implication of several well-known mechanisms underlying maternal effects.     

Age affects the expression of maternal care and subsequent behavioural development of offspring in a precocial bird

Variations of breeding success with age have been studied largely in iteroparous species and particularly in birds: survival of offspring increases with parental age until senescence. Nevertheless, these results are from observations of free-living individuals and therefore, it remains impossible to determine whether these variations result from parental investment or efficiency or both, and whether these variations occur during the prenatal or the postnatal stage or during both. Our study aimed first, to determine whether age had an impact on the expression of maternal breeding care by comparing inexperienced female birds of two different ages, and second, to define how these potential differences impact chicks' growth and behavioural development. We made 22 2-month-old and 22 8-month-old female Japanese quail foster 1-day-old chicks. We observed their maternal behaviour until the chicks were 11 days old and then tested these chicks after separation from their mothers. Several behavioural tests estimated their fearfulness and their sociality. We observed first that a longer induction was required for young females to express maternal behaviour. Subsequently as many young females as elder females expressed maternal behaviour, but young females warmed chicks less, expressed less covering postures and rejected their chicks more. Chicks brooded by elder females presented higher growth rates and more fearfulness and sociality. Our results reveal that maternal investment increased with age independently of maternal experience, suggesting modification of hormone levels implied in maternal behaviour. Isolated effects of maternal experience should now be assessed in females of the same age. In addition, our results show, for first time in birds, that variations in maternal care directly induce important differences in the behavioural development of chicks. Finally, our results confirm that Japanese quail remains a great laboratory model of avian maternal behaviour and that the way we sample maternal behaviour is highly productive.     

Mothering during early development influences subsequent emotional and social behaviour in Japanese quail

The aim of the present study was to investigate the influence of mothers on the emotional reactivity and social behaviour in young precocial Japanese quail. We used a classical method of maternal deprivation. Ethological tests and observations analysed and compared the behaviour of young artificially raised quail to that of young raised by adoptive maternal quail. After separation from mothers, brooded young were more fearful (frightened easily) in the presence of humans (human-observer tests) and more neophobic in novel environments (open-field and hole-in-the-wall tests) than young raised artificially. As chicks rarely expressed fear during the brooding period, no differences related to mothering could be observed at that time. In separation tests, brooded chicks jumped significantly more frequently than non-brooded chicks and later, observations of groups revealed that brooded chicks remained closer to one another than non-brooded chicks. Social motivation of brooded chicks appeared to be higher. These results indicate that, during their first days of life, mothers influence the emotional and social behaviour of their young.     

Maternal Epigenetic Transmission of Social Motivation in Birds

Behavioural development is a complex phenomenon involving interactions between genetic constraints and environmental influences. One of the most potent environmental influences during the ontogeny of the behavioural characteristics of young is played by mothers. In particular, mammalian mothers modulate the social competences of their young, influencing all their future social life. Here, we investigated the influence of the social characteristics of adoptive mother birds ( Coturnix coturnix japonica ) on the social motivation (origin of every social relationship) of the young they reared. We characterized the social behaviour of standard stock chicks reared by female quail, genetically selected either for low (LSR) or high (HSR) levels of social reinstatement (index of treadmill behaviour which combined the tendency to run towards conspecifics with the tendency to move away from them), to investigate epigenetic transmission of social motivation. Our results show that HSR and LSR adoptive mothers partially transmitted their social characteristics to their young: chicks reared by HSR females presented higher levels of social motivation than chicks reared by LSR females. This maternal influence appeared much clearer in young males than in young females. Our study reveals that, as in mammals, bird mothers influence epigenetically the development of the social behaviour of their young.     

Genetic selection on a behavioural fear trait is associated with changes in heart rate variability in quail

This study investigated whether genetic selection on a divergent behavioural trait of fearfulness (tonic immobility duration) was related to changes in the nervous control of the heart. Quail selected for either long or short tonic immobility (LTI or STI, respectively) duration was compared with an unselected control line (CTI). The autonomic control of the heart was assessed by heart rate variability analysis and pharmacological blockades. Quail were surgically fitted with a telemetric device. Heart rate before injection did not differ between the three lines. The vagal-sympathetic effect (VSE) at rest differed significantly from 1 in CTI and STI quail, suggesting that parasympathetic activity was dominant. In LTI quail, VSE did not differ from 1, suggesting a balance between parasympathetic and sympathetic activities. The intrinsic heart rate reached after the successive injections of propranolol and atropine did not differ between lines and was higher than the heart rate at rest in STI, which was in line with results of VSE at rest. After atropine injection, the sympathetic activity indicated by the low-frequency power was lower in CTI than in the two selected quail. After propranolol injection, the parasympathetic activity indicated by the root of the mean squares of successive differences and the high-frequency power was higher in STI than in CTI and LTI quail. Selection on tonic immobility duration thus appears to be associated with changes in the sympathovagal control of the heart, which may influence behavioural responses to stressful situations.     

Social instability in laying quail: consequences on yolk steroids and offspring's phenotype

Individual phenotypic characteristics of many species are influenced by non-genetic maternal effects. Female birds can influence the development of their offspring before birth via the yolk steroid content of their eggs. We investigated this prenatal maternal effect by analysing the influence of laying females' social environment on their eggs' hormonal content and on their offspring's development. Social instability was applied to groups of laying Japanese quail females. We evaluated the impact of this procedure on laying females, on yolk steroid levels and on the general development of chicks. Agonistic interactions were more frequent between females kept in an unstable social environment (unstable females) than between females kept in a stable social environment (stable females). Testosterone concentrations were higher in unstable females' eggs than in those of stable females. Unstable females' chicks hatched later and developed more slowly during their first weeks of life than those of stable females. The emotional reactivity of unstable females' chicks was higher than that of stable females' chicks. In conclusion, our study showed that social instability applied to laying females affected, in a non-genetic way, their offspring's development, thus stressing the fact that females' living conditions during laying can have transgenerational effects.     

Genetic differences in coping strategies in response to prolonged and repeated restraint in Japanese quail divergently selected for long or short tonic immobility

Exposure to fearful situations elicits behavioral and Hypothalamic–Pituitary–Adrenal (HPA) axis responses characteristic of the coping response of individual animals to counteract environmental challenges. The aim of this study was to investigate behavioral and corticotropic responses concomitantly following prolonged or repeated restraint stress by placing two genotypes of Japanese quail divergently selected for long (LTI) or short (STI) duration of tonic immobility (TI) in a crush cage. In our study, STI quail exhibited higher corticosterone (CORT) levels than LTI quail in response to prolonged restraint. STI quail struggled sooner and much more than LTI quail, and struggling behavior in STI quail progressively decreased during the course of restraint whereas LTI quail displayed very little struggling behavior in the crush cage. LTI quail are thus more likely to adopt a passive behavior coping strategy upon exposure to threat whereas STI quail behave more as active copers. The corticosterone responses shown by LTI and STI quail under restraint stress suggest that adrenocortical correlates of coping behavior in these genotypes of quail may be different from the coping styles previously described in other species. Repeated restraint slightly decreased CORT responses to stress in all experimental groups, but more markedly in male STI quail, whereas adrenal sensitivity and maximum adrenal corticosterone response capacity did not change in any group. On the other hand, neither behavioral habituation nor sensitization processes occurred in the context of repeated restraint in female and male LTI quail and female STI quail, whereas the decreases observed in some behavioral responses were interpreted to be the result of a habituation process in male STI quail.     

Adoption in Eastern Grey Kangaroos: A Consequence of Misdirected Care?

Adoption is rare in animals and is usually attributed to kin selection. In a 6-year study of eastern grey kangaroos (Macropus giganteus), 11 of 326 juveniles were adopted. We detected eight adoptions by observing behavioural associations and nursing between marked mothers and young and three more by analysing the relatedness of mothers and young using microsatellite DNA. Four adoptions involved reciprocal switches and three were by mothers whose own pouch young were known to subsequently disappear. Adoptive mothers were not closely related to each other or to adoptees but adoptive mothers and young associated as closely as did biological pairs, as measured by half-weight indices. Switch mothers did not associate closely. Maternal age and body condition did not influence the likelihood of adoption but females were more likely to adopt in years with high densities of females with large pouch young. Adoption did not improve juvenile survival. We conclude that adoptions in this wild population were potentially costly and likely caused by misdirected care, suggesting that eastern grey kangaroos may have poorly developed mother-offspring recognition mechanisms.     

Food Availability and Maternal Immunization Affect Transfer and Persistence of Maternal Antibodies in Nestling Pigeons

The ability of mothers to transfer antibodies (Abs) to their young and the temporal persistence of maternal Abs in offspring constitute important life-history traits that can impact the evolution of host-parasite interactions. Here, we examined the effects of food availability and parental immunization on the transfer and persistence of maternal antibodies in nestling pigeons (Columba livia). This species can transmit maternal Abs to offspring before hatching through the egg yolk and potentially after hatching through crop milk. However, the role of this postnatal substance in immunity remains elusive. We used a full cross-fostering design to disentangle the effects of food limitation and parental immunization both before and after hatching on the levels and persistence of maternal Abs in chicks. Parents were immunized via injection with keyhole limpet hemocyanin antigens. Using an immunoassay that specifically detected the IgY antibodies that are known to be transmitted via the yolk, we found that the levels of anti-KLH Abs in newly hatched chicks were positively correlated with the levels of anti-KLH Abs in the blood of their biological mothers. However, this correlation was not present between chicks and their foster parents, suggesting limited IgY transfer via crop milk to the chick’s bloodstream. Interestingly, biological mothers subjected to food limitation during egg laying transferred significantly fewer specific maternal Abs, which suggests that the transfer of antibodies might be costly for them. In addition, the persistence of maternal Abs in a chick’s bloodstream was not affected by food limitation or the foster parents’ anti-KLH Ab levels; it was only affected by the initial level of maternal anti-KLH Abs that were present in newly hatched chicks. These results suggest that the maternal transfer of Abs could be costly but that their persistence in an offspring’s bloodstream may not necessarily be affected by environmental conditions.     

Genetic differences in coping strategies in response to prolonged and repeated restraint in Japanese quail divergently selected for long or short tonic immobility

Exposure to fearful situations elicits behavioral and Hypothalamic–Pituitary–Adrenal (HPA) axis responses characteristic of the coping response of individual animals to counteract environmental challenges. The aim of this study was to investigate behavioral and corticotropic responses concomitantly following prolonged or repeated restraint stress by placing two genotypes of Japanese quail divergently selected for long (LTI) or short (STI) duration of tonic immobility (TI) in a crush cage. In our study, STI quail exhibited higher corticosterone (CORT) levels than LTI quail in response to prolonged restraint. STI quail struggled sooner and much more than LTI quail, and struggling behavior in STI quail progressively decreased during the course of restraint whereas LTI quail displayed very little struggling behavior in the crush cage. LTI quail are thus more likely to adopt a passive behavior coping strategy upon exposure to threat whereas STI quail behave more as active copers. The corticosterone responses shown by LTI and STI quail under restraint stress suggest that adrenocortical correlates of coping behavior in these genotypes of quail may be different from the coping styles previously described in other species. Repeated restraint slightly decreased CORT responses to stress in all experimental groups, but more markedly in male STI quail, whereas adrenal sensitivity and maximum adrenal corticosterone response capacity did not change in any group. On the other hand, neither behavioral habituation nor sensitization processes occurred in the context of repeated restraint in female and male LTI quail and female STI quail, whereas the decreases observed in some behavioral responses were interpreted to be the result of a habituation process in male STI quail.     

Ease of capture in lines of Japanese quail (Coturnix japonica) subjected to contrasting selection for fear or sociability

Japanese quail (Coturnix japonica) of lines, which have been subjected to contrasting selection for duration of the tonic immobility (TI) reaction or social reinstatement (SR) behaviour over many generations show corresponding differences in underlying fearfulness and sociality. As fearfulness and sociality are particularly influential traits in domesticated species, the finding that such traits respond to artificial selection may have important implications for poultry welfare and performance. However, it is not known if or how such selection has influenced human-animal interactions. The present experiment investigated the influence of fearfulness and SR behaviour on the ease with which birds could be caught and handled. Birds of lines selected for duration of the TI response or SR behaviour were reared in mixed line groups (LTI and STI or HSR and LSR) of 491 and 346 birds, respectively, until 6 weeks of age. When the birds were 2, 4, and 6 weeks of age, they were caught one by one and their individual capture ranks noted. In the group of birds selected for duration of the TI response, birds selected of the line selected for short duration of TI were caught before those selected for long duration of the response. In the group of birds selected for SR motivation, birds of the high line were caught before their low lines counterparts. Coefficients of concordance between capture ranks were significant and capture ranks did not differ significantly across ages. These results imply that selection for low levels of fear or high levels of sociality produces animals that are less disturbed by human interventions than animals selected for the opposite traits. The greater ease of capture of low fear line birds than high fear line birds may be explained by reduced fear of humans. The fact that the birds selected for high levels of SR behaviour are easier to catch than birds selected for low levels of sociality is less readily explicable. One hypothesis is that HSR line chicks tend to be more strongly imprinted on each other and the human caretaker. However, SR behaviour is highly species specific in both lines, existing evidence for line differences in social discrimination is limited and birds of the two lines show similar duration of the TI response. Despite this, whatever their underlying causation, these results demonstrate that genetic selection can be used to reduce negative reactions to human beings and may be of value in the improvement of both animal welfare and productivity.     

Habituation to humans affects yolk steroid levels and offspring phenotype in quail

In the field as well as in the laboratory, human-generated stress responses are reduced in adult animals previously habituated to humans in comparison to non-habituated individuals. In birds, yolk steroid levels vary with maternal environment and condition. We tested the hypothesis that the experience of female birds with humans could affect yolk steroids levels and offspring phenotype. Two groups of Japanese quail, one habituated to humans (H) and a second non-habituated (NH), were exposed daily to brief human disturbances. We analysed egg quality, offspring growth, and offspring emotional reactivity. NH females produced eggs with less androgens (testosterone and androstenedione) and more immunoreactive progesterone compared to birds habituated to humans. NH females produced eggs with less yolk, heavier shell and chicks hatching later and being smaller as compared to habituated individuals. A lower emotional reactivity was found in young of NH females compared to young of H females. Thus, human disturbance of the mother triggered different effects on chick phenotype depending on previous experience of mother birds with humans. In addition, we describe for the first time the influence of environmental stimuli on yolk immunoreactive progesterone levels. Our results show that a relatively minor difference in behavioral habituation may have substantial effects on eggs and offspring. This has obvious implications for keeping and handling laboratory animals, for conservation biology and for animal welfare.     

Social Effects of Maternal Age and Experience in Bison: Pre- and Post-weaning Contact Maintenance with Daughters

Effects of maternal age and experience on long-term contact maintenance interactions with female offspring were examined in bison (Bison bison), as part of a study of factors influencing dyadic behavioural interaction. During the first five months, patterns of change in interactions varied with maternal age and parity. Young (largely primiparous) mothers and their calves steadily increased the frequency with which they maintained contact, while older mothers and their calves became increasingly independent (e.g. calves walked away from mothers more and followed them less often). Filial independence was most extreme in the (last) calves of the two oldest mothers. These age-specific patterns appeared to result largely from contrasting shifts in maternal behaviour from Month 1 to 2; differences between groups before and after this shift were opposite in direction. In the first month, older mothers maintained more frequent contact and were behaviourally more synchronized with their calves. After the first month, however, the efforts of older mothers to maintain dyadic proximity abruptly decreased, while those of young mothers and their calves gradually increased. Subsequent differences between groups were more consistent. Young mothers and their daughters maintained closer contact and continued to do so well past weaning, at least as late as the third year, when daughters reached sexual maturity. Filial behaviour played an important role in determining patterns of dyadic spatial relations; the close proximity maintained by young mothers and their offspring appeared due largely to the frequency with which these calves followed their mothers' movements. The relatively independent calves of older mothers took greater initiative in contact interactions with mothers (e.g. nosing, licking). Young mothers appeared to strengthen long-term bonds with daughters by initiating contact frequently; the closeness of post-weaning dyadic association increased with the frequency of pre-weaning maternal contact.     

Experiential and hormonal correlates of maternal behavior in teen and adult mothers

This study explores the role of cortisol and early life experiences in the regulation of maternal behavior and mood in teen and adult mothers. Primiparous mothers (n=119) (teen mothers < 19 years, n=42), young mothers (19-25 years, n= 4), and mature mothers, (>25 years, n=43) were assessed for their maternal behavior, mood, and hormonal profile at approximately 6 weeks postpartum. Outcome measures were analyzed as a function of age and early life experience. Results showed an interaction between age and type of maternal behavior, where teen mothers engaged in more instrumental (e.g. changing diapers, adjusting clothes) less affectionate (e.g., stroking, kissing, patting) behavior, and mature mothers engaged in more affectionate and less instrumental behavior. When groups were reassessed based on early life experience (consistency of care during the first 12 years of life: consistent care; having at least one consistent caregiver, inconsistent care; having multiple and changing caregivers), an interaction was also found between consistency of care and type of behavior shown, where mothers who received inconsistent care engaged in more instrumental and less affectionate behavior. Compared to mature mothers, teen mothers who were breast feeding also had higher salivary cortisol levels, and high cortisol in teen mothers related to decreased fatigue and increased energy. These results suggest that early life experiences are linked to mothering behavior and are consistent with the emerging human and animal literature on intergenerational effects of mothering style.     

Mothers determine offspring size in response to own juvenile growth conditions

Through non-genetic maternal effects, mothers can tailor offspring phenotype to the environment in which young will grow up. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult environment of mothers. In this case maternal decisions about investment in offspring quality should already be determined during the juvenile phase of mothers. I tested this hypothesis by manipulating juvenile and adult maternal environments independently in a cichlid fish. Females raised in a poor environment produced larger young than females raised without food limitations, irrespective of the feeding conditions experienced during adulthood. This maternal boost was due to a higher investment in eggs and to faster larval growth. Apparently, mothers prepare their offspring for similar environmental conditions to those they encountered as juveniles. This explanation is supported by the distribution of these fishes under natural conditions. Juveniles live in a different and much narrower range of habitats than adults. Therefore, the habitat mothers experienced as juveniles will allow them to predict their offspring's environment better than the conditions in the adult home range.     

Prenatal light exposure affects early feather-pecking behaviour in the domestic chick

Recently we proposed that early feather pecking is a form of social exploration. Social recognition, important for exploration, is a lateralized function in the domestic chick. Lateralization of functions can be influenced by light exposure late in embryonic development. Therefore, we investigated whether this light exposure affected early posthatching feather-pecking behaviour in domestic chicks, Gallus gallus domesticus. White leghorn embryos either were exposed to light or remained in darkness in the last week of incubation. After hatching, they were housed in groups of two light-exposed and two dark-incubated chicks. Light-exposed chicks showed more feather pecking than did their dark-incubated cagemates. Dark-incubated chicks preferred to direct feather pecks to unfamiliar peers than to familiar peers; light-exposed chicks showed no preference. These effects were present in the first week after hatching and remained at least another 3 weeks. These results support the hypothesis that early gentle feather pecking is part of the normal behavioural repertoire of young chicks and influences social exploration. We discuss a possible mechanism underlying these results. We also suggest that it may be worthwhile not to expose embryos to light during the last week of incubation when housing hatchlings in commercial conditions, where feather pecking is a serious problem.     

Food availability modulates the effects of maternal antibodies on growth and immunity in young feral pigeons

It is now widely acknowledged that mothers can transfer their own immune experience to their progeny through the allocation of specific maternal antibodies (hereafter referred as MatAb) that can shape offspring phenotype and affect their fitness. However, the importance of environmental variability in modulating the effects of MatAb on offspring traits is still elusive. Using an experimental approach, we investigated how food availability interacted with MatAb to solve the trade‐off between humoral immunity and growth in young feral pigeons Columba livia. Results show that the inhibitory effect of MatAb on the humoral response of chicks was detected regardless of the food treatment. In addition, body mass growth was higher in chicks receiving lower amounts of maternal antibodies but only in chicks of the ad libitum food treatment. This contradicts previous studies and suggests that the transfer of MatAb could entail some costs for chicks and reduce their growth. Taken together these results reinforce the idea that the maternal antibodies play a central role in shaping offspring life‐history traits but that their adaptive value is highly dependent on the environmental context in which they are transmitted by the mother.     

Male attractiveness regulates daughter fecundity non-genetically via maternal investment

Mothers can non-genetically influence offspring phenotype in response to environmental conditions, including mate attractiveness. If such 'maternal effects' influence the offspring's reproduction and F2 generation, there is a mechanism for non-genetic trans-generational effects on phenotype, including epigenetic phenomena, with implications for evolution and population dynamics. We demonstrate in the zebra finch Taeniopygia guttata such non-genetic effects on offspring fecundity and the size of early stage F2 (eggs) in response to experimentally manipulated father's attractiveness. Our experimental design allowed us to deduce that the mechanism for this non-genetic paternal effect was via maternal investment in eggs. This affected female offspring size and, consequently, fecundity and F2 (egg) size. This demonstrates that female perception of mate attractiveness can have non-genetic, trans-generational fitness consequences and this may have important implications for the evolution of sexually selected traits and population dynamics.     

Long-term exposure to unpredictable and uncontrollable aversive events alters fearfulness in sheep

Numerous studies have investigated the emotional effects of various acute, potentially alarming events in animals, but little is known about how an accumulation of emotional experiences affects fearfulness. Fearfulness is a temperament trait that characterizes the propensity of an individual to be frightened by a variety of alarming events. The aim of this study was to investigate a putative alteration of fearfulness in sheep repeatedly exposed to various aversive events. Forty-eight 5-month-old female lambs were used. Over a period of 6 weeks, 24 of them (treated group) were exposed daily to various unpredictable and uncontrollable aversive events related to predatory cues, social context and negative handling that can occur under farming conditions. The other 24 lambs (control group) were housed in standard farming conditions (predictable food distribution and group handling). Fearfulness (behavioural and physiological responses) was assessed before and after the treatment period by subjecting the lambs to three standardized tests: individual exposure to suddenness and then to novelty in a test arena, and group exposure to a motionless human in the home pen. As biomarkers of stress, leukocyte counts, heart rate and cortisol concentrations were measured in the lambs in their home pens. Before the treatment, the emotional responses of the groups did not differ. After the treatment, treated lambs approached the human less often, had less contact with the novel object and vocalized more than controls in individual tests, suggesting that long-term exposure to unpredictable and uncontrollable aversive events increases subsequent fearfulness in sheep. In addition, treated lambs had lower leukocyte counts, heart rate and cortisol levels, pointing to a chronic stress state. These findings suggest that increased fearfulness may be used as a sign of chronic stress in farm animals.     

Moderate Heat Challenge Increased Yolk Steroid Hormones and Shaped Offspring Growth and Behavior in Chickens

Background

Environmental challenges might affect the maternal organism and indirectly affect the later ontogeny of the progeny. We investigated the cross-generation impact of a moderate heat challenge in chickens. We hypothesized that a warm temperature–within the thermotolerance range- would affect the hormonal environment provided to embryos by mothers, and in turn, affect the morphology and behavioral phenotype of offspring.

Methodology/Principal Findings

Laying hens were raised under a standard thermal condition at 21°C (controls) or 30°C (experimental) for 5 consecutive weeks. A significant increase was observed in the internal temperature of hens exposed to the warm treatment; however plasma corticosterone levels remained unaffected. The laying rate was not affected, but experimental hens laid lighter eggs than the controls during the treatment. As expected, the maternal thermal environment affected yolk hormone contents. Eggs laid by the experimental hens showed significantly higher concentrations of yolk progesterone, testosterone, and estradiol. All chicks were raised under standard thermal conditions. The quality of hatchlings, growth, feeding behavior and emotional reactivity of chicks were analyzed. Offspring of experimental hens (C30 chicks) were lighter but obtained better morphological quality scores at hatching than the controls (C21 chicks). C30 chicks expressed lesser distress calls when exposed to a novel food. Unlike C21 chicks, C30 chicks expressed no preference for energetic food.

Conclusion/Significance

Our findings suggest that moderate heat challenge triggers maternal effects and modulate the developmental trajectory of offspring in a way that may be adaptive. This suggests that the impact of heat challenges on captive or wild populations might have a cross-generation effect.