Maternal Manipulation Research Articles

Maternal manipulation of offspring size can trigger the evolution of eusociality in promiscuous species

Eusocial organisms typically live in colonies with one reproductive queen supported by thousands of sterile workers. It is widely believed that monogamous mating is a precondition for the evolution of eusociality. Here, we present a theoretical model that simulates a realistic scenario for the evolution of eusociality. In the model, mothers can evolve control over resource allocation to offspring, affecting offspring's body size. The offspring can evolve body-size-dependent dispersal, by which they disperse to breed or stay at the nest as helpers. We demonstrate that eusociality can evolve even if mothers are not strictly monogamous, provided that they can constrain their offspring's reproduction through manipulation. We also observe the evolution of social polymorphism with small individuals that help and larger individuals that disperse to breed. Our model unifies the traditional kin selection and maternal manipulation explanations for the evolution of eusociality and demonstrates that-contrary to current consensus belief-eusociality can evolve despite highly promiscuous mating.

Effects on the Lower Limb Anthropometry in Patients with Unilateral Idiopathic Clubfoot Treated Conservatively with Ponseti Technique

Category: Other; Basic Sciences/Biologics Introduction/Purpose: Congenital Talipes Equino-Varus is amongst the most common congenital orthopaedic anomalies. Limb length discrepancy is considered as the commonest complication in patients with unilateral clubfoot despite the best non- operative correction. In the present series of cases we aim to study all the possible effects on the lower limb anthropometry in patients with unilateral idiopathic clubfoot who have been managed conservatively. Methods: This retrospective study included 47 patients with idiopathic, unilateral CTEV who were successfully treated with Ponseti’s technique. Patients with neuromuscular causes, those requiring extensive soft tissue releases or osteotomies and cases with relapsed or recurrent CTEV were excluded. Casting was initiated following maternal manipulation for one to two weeks. Pirani scores were recorded initially and weekly till correction. After deformity correction, anthropometric measurements (femur and tibia length, calf girth and foot length and foot height) were recorded and compared to the opposite normal limb. Results: Only four patients showed a difference in their femur length while difference in tibia length was noted in 10 patients which were statistically not significant. A significant difference was found in the calf girth, foot height and foot length affecting 28, 43 and 42 patients respectively. Conclusion: Patients with unilateral clubfoot treated with Ponseti’s correction tend to have anthropometric difference between the affected and the unaffected extremity. While there is a relative sparing of femur and tibia length, calf girth, foot height and foot length are consistently less compared to the unaffected side.

Can nesting behaviour allow reptiles to adapt to climate change?

A range of abiotic parameters within a reptile nest influence the viability and attributes (including sex, behaviour and body size) of hatchlings that emerge from that nest. As a result of that sensitivity, a reproducing female can manipulate the phenotypic attributes of her offspring by laying her eggs at times and in places that provide specific conditions. Nesting reptiles shift their behaviour in terms of timing of oviposition, nest location and depth of eggs beneath the soil surface across spatial and temporal gradients. Those maternal manipulations affect mean values and variances of both temperature and soil moisture, and may modify the vulnerability of embryos to threats such as predation and parasitism. By altering thermal and hydric conditions in reptile nests, climate change has the potential to dramatically modify the developmental trajectories and survival rates of embryos, and the phenotypes of hatchlings. Reproducing females buffer such effects by modifying the timing, location and structure of nests in ways that enhance offspring viability. Nonetheless, our understanding of nesting behaviours in response to climate change remains limited in reptiles. Priority topics for future studies include documenting climate-induced changes in the nest environment, the degree to which maternal behavioural shifts can mitigate climate-related deleterious impacts on offspring development, and ecological and evolutionary consequences of maternal nesting responses to climate change. This article is part of the theme issue 'The evolutionary ecology of nests: a cross-taxon approach'.

Effects of nutritional manipulation on intranidal social behaviour in a small carpenter bee

Understanding how subordinate altruistic roles arise among genetically similar individuals is critical to understanding the evolution of animal societies. In particular, maternal manipulation of offspring resources is an understudied process by which mothers may produce cooperative phenotypes among offspring. Mothers of the small carpenter bee Ceratina calcarata frequently produce a smaller underprovisioned daughter, known as the dwarf eldest daughter (DED) in the first cell of their nests. This DED is more subordinate and more easily coerced into helping siblings, aiding mothers in guarding and supplemental foraging and feeding of adult siblings. Colonies of these bees thus offer a unique opportunity to study the indirect effect of nutritional manipulation on behaviour, in particular mutual tolerance. We hypothesized that aggression would be greatest in the presence of DEDs, as they are actively coerced by their mothers to forage and feed their siblings. Here we experimentally added nutrition to the first cells of C. calcarata observation nests to test the effect of nutritional manipulation on colony intranidal behaviour. Using infrared cameras, control ( N = 13) and nutritionally supplemented nests ( N = 13) were observed for a total of 607.5 h. Behaviours were scored and categorized. Although aggression was unaffected, avoidant behaviour was greater in nutritionally supplemented nests. This study is the first experiment examining the effect of nutritional manipulation on intranidal behaviour in a stem nesting bee. Our results indicate that maternal manipulation and nutritional deprivation by mothers is critical to the production of mutual tolerance and division of labour. • We test effects of nutritional manipulation on mutual tolerance in C. calcarata. • Maternal manipulation of offspring nutrition influenced colony behaviour. • Avoidance behaviour increased in colonies with nutritional supplementation. • Control colonies with naturally underprovisioned daughters were more tolerant. • Mutual tolerance is among the most rudimentary social requirements.

Effect of dietary protein and energy intake on embryonic survival and gene expression in the uterine endometrium of early pregnant gilts

Porcine embryonic loss during early gestation is a serious problem in swine production. Improving embryonic survival can be achieved by maternal manipulation. Protein and energy are two major components of the diet, which play decisive roles in embryonic survival. This study was performed to evaluate the effects of enhancing maternal protein or energy intake on embryonic survival during early gestation in gilts and to explore the underlying mechanism. From day (d) 0 to 30 of gestation, 40 gilts (Landrace × York) were randomly allocated to 5 diets according to daily intake of low (L, National Research Council (NRC) recommendation for gestation gilts), medium (M, 20% higher than NRC) or high (H, 40% higher than NRC) CP or metabolisable energy (ME) (LCPLME, MCPLME, HCPLME, LCPHME, HCPHME). Gilts were sacrificed on d 30 of gestation, and number of foetuses and corpora lutea, embryonic survival rate, uterine weight, and total volume of allantoic fluid were recorded or calculated. Gene expression was determined by Quantitative Real-time PCR (qPCR), western blot or immunohistochemistry. Results showed that increasing protein or ME intake significantly increased embryonic survival rate. Compared with diet LCPLME, plasma progesterone (P4) concentration in diet LCPHME increased at d 14 and d 30 of gestation. Progesterone receptor (PGR) was found not to be expressed in the epithelia but was strongly expressed in the stroma of the endometrium. Increasing protein or ME intake did not alter PGR expression in the endometrium. There was also no change in the amount of P4, hepatocyte growth factor, and fibroblast growth factor-7 in the endometrium. The mRNA abundance of cationic amino acid transporter 1 in the endometrium in diet LCPHME and HCPHME was significantly lower than in diet LCPLME. Diet HCPLME showed a tendency to increase neutral amino acid transporter 1 mRNA expression in the endometrium compared to diet LCPLME (P = 0.087). In conclusion, increasing maternal protein or ME intake had a positive effect on the embryonic survival. Increased protein intake by 20 or 40% did not alter plasma P4 level, but increasing ME intake by 40% improved plasma P4 concentration at d 14 and 30 of gestation. Increasing maternal protein or ME intake did not induce PGR expression in the endometrium. Maternal protein and energy intake likely mediate transportation of cationic and neutral amino acids from mother to foetus to affect embryonic survival and development.

The role of climate and maternal manipulation in determining and maintaining reproductive mode in Liolaemus lizards

AbstractEvolutionary transitions in life‐history strategies, such as the shift from egg‐laying to live birth (viviparity) are of great interest to evolutionary biologists. In squamate reptiles, several hypotheses have been proposed to explain viviparity including the cold climate hypothesis, maternal manipulation hypothesis, hypoxia hypothesis, and several others. We used two approaches: first we studied 45 species of Liolaemus, a genus where nearly 50% of species are viviparous, using a diverse ecophysiological dataset to examine the cold climate and maternal manipulation hypotheses. We collected environmental thermal data (accounting for elevational differences among species), physiological traits including preferred body temperature and its coefficient of variation as an indicator of precision in thermoregulation. Additionally, we collected standard metabolic rates for 23 of the 45 species. In one clade (the darwinii group of species) with both reproductive modes, we ran our second approach. We tested for differences in thermal physiology and metabolic rates between viviparous and oviparous species during pregnancy and non‐pregnancy periods. The cold climate hypothesis received strong support because viviparous species occur in sites with colder air temperatures (including areas at both higher elevations and latitudes) compared with oviparous species. Our detailed analysis showed that the maternal manipulation hypothesis also is supported; pregnant viviparous females show lower variation in their selected temperature. Our evidence suggests that the Andean orogeny is likely to have played a key role in the diversification of Liolaemus lizards and the evolution of viviparity in this clade may have been driven by a variety of physiological advantages accrued at different stages of embryogenesis and over evolutionary time. Thus, historical climate changes may have led to egg retention and may have been accompanied by other adaptations such as thermoregulation precision.

Effects of maternal exposure to a bacterial antigen and altered post-hatching rearing conditions on avian offspring behaviour

The early-life environment plays a crucial role in shaping morphological, physiological, and behavioural traits, with potential long-term consequences for fitness. Indeed, a set of factors experienced by offspring during prenatal and early post-natal development has been recognised to affect behavioural trait expression in later life. Several studies have shown that in birds, nutritional and social rearing conditions and maternal and/or neonatal immunisations may profoundly determine the development and establishment of behaviour in offspring. To our knowledge, no research has examined whether and how the interaction between immune-mediated maternal effects and post-hatching rearing conditions affects offspring behaviour. Here, we studied the effects of maternal exposure to a bacterial antigen and altered brood size on docility, breathing rate, and aggression in the offspring of great tit, Parus major. We used a 2 × 2 design to investigate the interactive effects of maternal immunisation and brood size manipulation on offspring behavioural development. We found no such interactive effect on offspring behaviour, although we observed it regarding to offspring body mass and tarsus length. Maternal immunisation itself did not affect offspring behaviour. However, we demonstrated that the offspring breathing rate and level of aggression were affected by brood size manipulation. Both breathing rate and aggression in offspring reared in enlarged broods were lower than those in offspring reared in non-manipulated broods. Our study did not confirm earlier reports that immune-mediated maternal effects modulate offspring behavioural development, but we showed that brood size during rearing might indeed be a factor that affects offspring behaviour.Significance statementThe early environment experienced by offspring constitutes a significant source of developmental plasticity, which may profoundly affect the establishment of their behavioural traits. Food availability, social conditions, and maternal or offspring infection are crucial factors shaping various behavioural traits in birds. However, there remains a lack of studies emphasising the potential interactive effects of early-life conditions on behavioural trait development in natural bird populations. Here, to our knowledge for the first time, we experimentally examined how maternal immunisation and altered post-hatching rearing conditions interact to determine the behaviour of fledged offspring. We found that maternal treatment and brood size manipulation interactively affected offspring body mass and tarsus length, but this interaction had no effect on offspring behaviour. Our findings suggest that different mechanisms may underlie the development of morphological and behavioural traits.

Hyperlipidic diet affects body composition and induces anxiety-like behaviour in intrauterine growth-restricted adult mice.

What is the central question of this study? What is the effect in male and female offspring of a protein-deficient diet producing intrauterine growth restriction (IUGR) in maternal mice on morphometric, metabolic and behavioural parameters before and after a challenge with a fat diet? What is the main finding and its importance? Male and female mice presented different growth trajectories after birth. IUGR favoured increased adiposity in male mice, and high-fat diet-induced anxiety-like behaviour in female mice. As there is sexual dimorphism in the response to maternal manipulations, we aimed to analyse the effects of intrauterine growth restriction (IUGR) in both sexes on morphometric, metabolic and behavioural parameters throughout postnatal development, and after challenge with a hyperlipidic diet. Female Swiss mice (n=59) were distributed into two groups (SD: standard diet, n=26; and PDD: isocaloric protein-deficient diet, n=33), 2weeks before mating and during the gestational period. After birth, offspring from SD and PDD dams were cross-fostered and nurtured by SD dams until postnatal day (PND) 28. At PND 60 all animals were challenged with a hypercaloric diet for 4weeks. Offspring birth weight was significantly reduced in the PDD group compared to the SD group (P=0.0001), but only male offspring presented a rapid catch-up during the first 21days of development. Although no differences in body weight were observed between groups after the challenge with the hyperlipidic diet, an increase in the relative perigonadal white adipose tissue (P=0.009) and a decrease in gross gastrocnemius muscle weight (P=0.010) were observed in the PDD males. In relation to behavioural tests, there was an increase in locomotion in both sexes (P=0.0001), and a decrease in female grooming (P=0.006) in the PDD group. Additionally, females from the PDD group showed increased hyperlipidic food intake. In conclusion, IUGR affected both sexes, with females showing prominent behavioural modifications and males presenting altered body composition elicited by a hyperlipidic diet.

Maternal manipulation during late gestation (GDs 17-20) enhances ethanol consumption and promotes changes and opioid mRNA expression in infant rats

Fetal ethanol experience generates learning and memories capable to increase ethanol consummatory behaviors during infancy. Opioid system seems to be involved in mediating those alcohol-related behaviors. In this work, we proposed to study the impact of prenatal exposure to a moderate ethanol dose, upon ingestion of the drug and possible ethanol-induced molecular changes on opioid precursor peptides (POMC, Pro-enk and Pro-DYN) and receptors (MOR, DOR and KOR) mRNA expression, in hypothalamus. Pregnant rats received during gestational days (GDs) 17-20, a daily intragastric (i.g.) administration with 2g/kg ethanol or water. A third group of dams was left undisturbed during pregnancy (Unmanipulated group). Intake test was conducted at postnatal days (PDs) 14-15. Three groups of pups were performed: control (no intake test), water (vehicle) and 5% ethanol. At the end of intake test blood samples were taken to quantify blood ethanol concentrations (BECs) and hypothalamus sections were obtained to perform qRT-PRC assessment of opioid precursor peptides and receptors. The analysis of the consummatory responses (% of consumption) and pharmacokinetic profiles (BECs) suggested that maternal manipulation induced by i.g. intubations, during the last four days of gestation (whenever ethanol or water), are sufficient to induce infantile ethanol intake during infancy. Gene expression from the hypothalamus of unmanipulated group revealed that infantile ingestive experiences with ethanol can down-regulate expression of mRNA Pro-Dyn and up-regulate mRNA expression of MOR and KOR. Finally, MOR mRNA expression was attenuated by prenatal i.g. manipulation in pups exposed to 5% ethanol.

Offspring of aged mothers of rotifer Brachionus calyciflorus shows lower sexual propensity than their elder siblings under crowded conditions

Maternal effects are one of the most interesting topics in evolutionary ecology as they can affect the rate of evolution and population dynamics by phenotypic manipulation of offspring related to fitness. At present, studies examining the interaction between maternal environment effects and maternal age (birth order) effects are scarce. We designed an experiment to reveal whether environmentally induced maternal manipulation works equally on offspring across birth orders in rotifer Brachionus calyciflorus. In the experiment, newborn amictic mothers (F0) were cultured parallelly in low (LD) and high (HD) population density. Offspring (F1) of young (YF0) and old (OF0) mothers in both LD and HD groups were cultured under a crowded condition and the mixis ratios of both F1 and F2 were examined. Results show that F1 from HD-YF0 had a significantly higher mixis ratio and those amictic ones produced daughters (F2) with higher mixis ratio in response to crowding than those from HD-OF0. In contrast, no such differences among F1 siblings were found in the LD-F0 group. Therefore, the effect of maternal age on the sexual propensity of the offspring can be affected independently by the maternal population density.

An Adlerian Analysis of the Problematic Mother-Daughter Relationship in Eileen Chang's The Golden Cangue and Elfriede Jelinek's Die Klavierspielerin

The Chinese writer Eileen Chang's novella Jinsuo Ji (The Golden Cangue, 1943) and Elfriede Jelinek's novel Die Klavierspielerin (1983) depict a problematic mother-daughter relationship. Despite their disparate dynastic and national backgrounds, these two works bear many similarities in terms of extreme maternal manipulation and despair the daughters undergo. Unlike the idealized stereotype of a mother who selflessly nurtures and protects her daughter, the mothers in these two works are not caregivers; rather, they destroy their daughters' lives. Drawing upon Alfred Adler's theory of individual psychology, this paper investigates the mothers' mentality in their respective contexts and probes unconventional mother-daughter relationships.

The influence of sociality, caste, and size on behavior in a facultatively eusocial bee

Social cooperation requires increased tolerance of other individuals. We used social and solitary individuals of the facultatively eusocial bee Megalopta genalis to compare interactions with non-nestmate individuals in a standardized behavioral assay, a circle tube. We set up interactions between bees from different nests matched for caste (solitary, social: queen or worker). We found more tolerance in social than solitary pairs, but found no difference in aggression. We also found that workers continued expressing caste-typical behavior even when matched against other workers from different nests. However, there was no difference in expression of queen-typical behaviors between the three groups. Our data on social caste show that outside of the queen–worker social context, both queens and workers express similar levels of queen-like behavior. However, workers still express higher levels of worker-like behavior than do queens. We found no effect of variation in ovary size on behavior. We found that body size correlated positively with queen-like behaviors, and negatively with worker-like behaviors. Our body size data suggest that the worker phenotype may result from naturally occurring size-correlated variation in behavior, combined with maternal manipulation of both body size through nutrition and behavior and ovary development through social aggression.

Maternal Dietary Manipulation Programs Offspring’s Metabolism

There is emerging evidence supporting fetal metabolism is determined in utero, and the programmed metabolism persists into adulthood. Understanding how metabolic dysfunction is transmitted through generations will help identify the biological mechanisms underlying obesity and cardiometabolic disease, and provide insight regarding rational strategies for treatment and prevention. Metabolic parameters were assessed in offspring from dams fed different diets for 4 weeks before pregnancy and during pregnancy and lactation; open standard (OSD), isocaloric low protein (LPD, 8% protein) and high-fat diets (HFD, 60% fat). After weaning, offspring had OSD. Offspring from dams fed LPD showed 31% reduced birthweights compared to offspring from dams fed OSD, and the difference persisted into 6 months of age. At 3 and 6 months, they exhibited significantly smaller perigonadal fat pad than offspring from dams fed OSD. In addition, they showed significantly higher blood glucose concentration than offspring from dams fed OSD at birth (52.35mg/dl vs. 63.71 mg/dl). In case of offspring from dams fed HFD, they exhibited significantly higher body weights (5.99g vs. 6.56g) and blood glucose level at birth (52.35mg/dl vs. 64.64 mg/dl) compared to offspring from dams fed OSD. The difference of body weights continued to 3week-old at weaning, and they showed significantly impaired glucose tolerance compared to offspring from dams fed OSD at the same age. Maternal dietary manipulation strongly influenced metabolic phenotypes such as birthweight, blood glucose concentration at birth and glucose tolerance, and the programed phenotypes persisted into adulthood. Therefore, maternal nutrition programs a glucose intolerant phenotype in offspring indicating that in utero stress augments diabetes risk not only childhood but also adulthood. We expect our findings in this study to help clarify the mechanism how the utero environment affects offspring’s metabolism, which would explain in part the intergenerational transmission of obesity. Disclosure M. Kang: None. J.P. Antipenko: None. X. Liu: None. K.M. Habegger: Consultant; Self; Glyscend, Inc.. Research Support; Self; Glyscend, Inc.. Consultant; Self; Intarcia Therapeutics, Inc. W. Garvey: Advisory Panel; Self; Novo Nordisk Inc., Merck & Co., Inc.. Research Support; Self; Sanofi, Pfizer Inc., Novo Nordisk Inc., AstraZeneca, Merck & Co., Inc., Elcelyx Therapeutics, Inc., Lexicon Pharmaceuticals, Inc., Eisai Inc..

The effect of pre-laying maternal immunization on offspring growth and immunity differs across experimentally altered postnatal rearing conditions in a wild songbird

BackgroundPrenatal antibody transfer is an immune-mediated maternal effect by which females can shape postnatal offspring resistance to pathogens and parasites. Maternal antibodies passed on to offspring provide primary protection to neonates against diverse pathogenic antigens, but they may also affect offspring growth and influence the development of an offspring’s own immune response. The effects of maternal antibodies on offspring performance commonly require that the disease environment experienced by a mother prior to breeding matches the environment encountered by her offspring after hatching/birth. However, other circumstances, like postnatal rearing conditions that affect offspring food availability, may also determine the effects of maternal antibodies on offspring growth and immunity. To date, knowledge about how prenatal immune-mediated maternal effects interact with various postnatal rearing conditions to affect offspring development and phenotype in wild bird population remains elusive. Here we experimentally studied the interactive effects of pre-laying maternal immunization with a bacterial antigen (lipopolysaccharide) and post-hatching rearing conditions, altered by brood size manipulation, on offspring growth and humoral immunity of wild great tits (Parus major).ResultsWe found that maternal immunization and brood size manipulation interactively affected the growth and specific humoral immune response of avian offspring. Among nestlings reared in enlarged broods, only those that originated from immunized mothers grew better and were heavier at fledging stage compared to those that originated from non-immunized mothers. In contrast, no such effects were observed among nestlings reared in non-manipulated (control) broods. Moreover, offspring of immunized females had a stronger humoral immune response to lipopolysaccharide during postnatal development than offspring of non-immunized females, but only when the nestling was reared in control broods.ConclusionsThis study demonstrates that offspring development and their ability to cope with pathogens after hatching are driven by mutual influences of pathogen-induced prenatal maternal effects and post-hatching rearing conditions. Our findings suggest that immune-mediated maternal effects may have context-dependent influences on offspring growth and immune function, related to the postnatal environmental conditions experienced by the progeny.

Bonds Forged on the Double-Sided Anvil of Guilt and Praise: Maternal Manipulation in After the Fall and Suddenly Last Summer

Abstract Drawing on their own experiences, both Arthur Miller and Tennessee Williams explored the boundaries of mother-son relationships within their respective plays: After the Fall and Suddenly Last Summer. What Miller and Williams portrayed in these works, however, is far removed from pure, selfless, and loving examples of maternal care. Rose and Violet Venable are both, in turn, guilty of manipulating their sons to suit their own needs—resulting in devastating psychological damages inflicted on both of their sons.

Chicken or egg? Outcomes of experimental manipulations of maternally transmitted hormones depend on administration method - a meta-analysis.

Steroid hormones are important mediators of prenatal maternal effects in animals. Despite a growing number of studies involving experimental manipulation of these hormones, little is known about the impact of methodological differences among experiments on the final results expressed as offspring traits. Using a meta-analytical approach and a representative sample of experimental studies performed on birds, we tested the effect of two types of direct hormonal manipulations: manipulation of females (either by implantation of hormone pellets or injection of hormonal solutions) and manipulation of eggs by injection. In both types of manipulation we looked at the effects of two groups of hormones: corticosterone and androgens in the form of testosterone and androstenedione. We found that the average effect on offspring traits differed between the manipulation types, with a well-supported positive effect of egg manipulation and lack of a significant effect of maternal manipulation. The observed average positive effect for egg manipulation was driven mainly by androgen manipulations, while corticosterone manipulations exerted no overall effect, regardless of manipulation type. Detailed analyses revealed effects of varying size and direction depending on the specific offspring traits; e.g., egg manipulation positively affected physiology and behaviour (androgens), and negatively affected future reproduction (corticosterone). Effect size was negatively related to the dose of androgen injected into the eggs, but unrelated to timing of manipulation, offspring developmental stage at the time of measuring their traits, solvent type, the site of egg injection and maternal hormone delivery method. Despite the generally acknowledged importance of maternal hormones for offspring development in birds, the overall effect of their experimental elevation is rather weak, significantly heterogeneous and dependent on the hormone and type of manipulation. We conclude by providing general recommendations as to how hormonal manipulations should be performed in order to standardize their impact and the results achieved. We also emphasize the need for research on free-living birds with a focus on fitness-related and other long-term effects of maternal hormones.

Sibling competition and not maternal allocation drives differential offspring feeding in a sexually size-dimorphic bird

Sex allocation models still fail to predict the complex sex ratio patterns in broods of vertebrates. A major problem when studying mother–brood interactions is the difficulty in disentangling hypotheses involving maternal preferences from processes that do not imply maternal manipulation. We studied maternal resource allocation in mixed-sex, same-sex and single-chick broods in the great bustard, Otis tarda. Females normally rear a single chick, and previous work has shown that maternal investment influences male more than female breeding success. Therefore, mothers of two-chick broods were assumed to be in good condition and candidates to show a preference for sons. Results showed that male chicks of mixed-sex broods remained close to the mother for twice as long as their sisters, and received double the number of maternal feedings. However, sex differences in maternal feeding rate disappeared when considering only simultaneous begging approaches from both siblings. Proximity to the mother and its interaction with begging approach intensity were the factors determining the higher begging success of male chicks. In single-chick broods, females did not receive fewer maternal feedings than males. Overall, our results suggest that female chicks of mixed-sex broods become outcompeted by their larger brothers, which remain close to the mother much longer, preventing their sisters from taking a larger share of maternal feedings. We conclude that mothers do not show a preference for feeding male over female chicks, and that the sex differences in feeding rate are determined by the higher food requirements of male chicks due to their sexually selected, much faster growth rates. The higher mortality of females in mixed-sex broods contrasts with the pattern of male-biased mortality typical in this species, and supports our interpretation of an asymmetric competitive ability of male offspring as the mechanism responsible for the sex bias in maternal expenditure.

Maternal effects and parent-offspring conflict.

Maternal effects can provide offspring with reliable information about the environment they are likely to experience, but also offer scope for maternal manipulation of young when interests diverge between parents and offspring. To predict the impact of parent-offspring conflict, we model the evolution of maternal effects on local adaptation of young. We find that parent-offspring conflict strongly influences the stability of maternal effects; moreover, the nature of the disagreement between parents and young predicts how conflict is resolved: when mothers favor less extreme mixtures of phenotypes relative to offspring (i.e., when mothers stand to gain by hedging their bets), mothers win the conflict by providing offspring with limited amounts of information. When offspring favor overproduction of one and the same phenotype across all environments compared to mothers (e.g., when offspring favor a larger body size), neither side wins the conflict and signaling breaks down. Only when offspring favor less extreme mixtures relative to their mothers (something no current model predicts), offspring win the conflict and obtain full information about the environment. We conclude that a partial or complete breakdown of informative maternal effects will be the norm rather than the exception in the presence of parent-offspring conflict.

Development of object manipulation in wild chimpanzees

Chimpanzees' natural propensity to explore and play with objects is likely to be an important precursor of tool use. Manipulating objects provides individuals with pivotal perceptual-motor experience when interacting with the material world, which may then pave the way for subsequent tool use. In this study, we were interested in the influence of social models on the developmental patterns of object manipulation in young chimpanzees, Pan troglodytes schweinfurthii , of the Sonso community of Budongo Forest, Uganda. This community is interesting because of its limited tool repertoire, with no records of stick-based foraging in over 20 years of continuous observations. Using cross-sectional data, we found evidence for social learning in that young individuals preferentially played with and explored materials manipulated by their mothers. We also found that object manipulation rates decreased with age, whereas the goal directedness of these manipulations increased. Specifically, stick manipulations gradually decreased with age, which culminated in complete disregard of sticks around the age of 10 years, a pattern not found for other tool materials, which were all used throughout adulthood. Overall, young chimpanzees initially explored and played unselectively with any object found in the environment before becoming increasingly influenced by their mothers' goal-directed object manipulations. • Sonso chimpanzees of Budongo Forest, Uganda, are atypical in lacking stick tools. • We studied this cultural phenomenon with cross-sectional ontogenetic data. • Object manipulation rates decreased with age while goal directedness increased. • Loss of stick use occurred during the transition to adulthood, around the age of 10. • Infant choice and interest in tool materials was driven by maternal manipulations.

Sociodemographic variation in foraging behavior and the adaptive significance of worker production in the facultatively social small carpenter bee, Ceratina calcarata

Provisioning for young offspring is an archetypical form of parental investment. Ceratina calcarata bees provide extended maternal care to their young and demonstrate an unusual strategy of dual-phase pollen provisioning. Most bees first gather provisions as they establish nests in spring. However, C. calcarata mothers will also feed their newly eclosed young a second time, perhaps ensuring their survival during a long winter diapause. Some mothers rear a small, worker-like daughter to assist them during this second provisioning phase. We studied provisioning behavior in C. calcarata to examine patterns of maternal investment and foraging dynamics throughout the breeding season. Mothers typically made a high number of short-duration foraging trips each day, whereas late-season females tended to make fewer and longer trips. This difference in foraging duration may indicate a lower risk of brood loss in those nests where mature offspring are present. Nest demographic data revealed that an offspring laid in the first brood cell position is typically female and usually smaller than her siblings. In 29% of the nests, this small daughter was observed to adopt a forager role at maturity and provisioned for her siblings. Dwarf daughters had a higher number of active days and foraging trips per day in orphaned nests than in nests where a mother was present. The foraging behaviors of worker-like daughters were similar in length of foraging trip and handling time to mothers during this second provisioning period. We hypothesize that incipiently social foraging by this smallest daughter may act as a form of insurance against brood loss during occasions when a mother is unable to sufficiently provision for her eclosed offspring during the second phase. Parental investment in the size and sex of offspring is under strong selection for assured fitness returns. For example, many social insect mothers make an initial investment in small offspring to take on risky foraging behavior while they specialize on future reproduction. Solitary and facultatively social species provide an important baseline to understand the evolution of social complexity from natural variation in maternal care and foraging behavior. Here, we characterize the parental investment strategies of a subsocial small carpenter bee and reveal the potential adaptive significance of prolonged maternal care and worker production in this species. Mothers provide an initial investment that is extended by workers providing alloparental care to siblings. Maternal manipulation of dwarf eldest daughters may serve as an insurance mechanism in the event of maternal mortality to assure the survival of siblings.