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A short gestation, low birth weight and presence of cutaneous exchange of O2 and CO2 comprise altricial features of newborn marsupials and that collectively implies a highly immature respiratory system. In the present study, we investigated various respiratory characteristics of the neonatal/postnatal tammar wallaby, a species of marsupial in which > 30% of the newborn’s total O2 demands are supported by cutaneous rather than pulmonary gas exchange. The ventilatory response (HVR) to acute hypoxia (10% inspired O2) was absent in the newborn (1 day old) pouch young; a hypoxic hypometabolism contributed entirely to the hyperventilation (increased pulmonary convection requirement). A high (compared to older animals) resting metabolic cost to breathe and an inefficient respiratory system suggest the lack of a HVR might be due to an energetic constraint that impinges on their ability to sustain an increase in ventilation. The latter was supported by the inability of the newborn to tolerate metabolic-ventilatory stimulation following administration of the metabolic uncoupler, 2,4-dinitrophenol (2,4-DNP). At 1 week of age, the cost of breathing was reduced, which coincided with the expression of a significant ventilatory response to hypoxia, a more energetically efficient respiratory system, and tolerance to 2,4-DNP. These data suggest this species of marsupial is born with major respiratory insufficiency, and that their pronounced dependence on the skin for metabolic gas exchange is of critical importance for survival.

Small, non-hibernating endotherms increase their thermogenic capacity to survive seasonal cold, through adult phenotypic flexibility. In mammals, this response is primarily driven by remodeling of brown adipose tissue (BAT), which matures postnatally in altricial species. In many regions, ambient temperatures can vary dramatically throughout the breeding season. We used second-generation lab-born Peromyscus leucopus, cold exposed during two critical developmental windows, to test the hypothesis that adult phenotypic flexibility to cold is influenced by rearing temperature. We found that cold exposure during the postnatal period (14 °C, birth to 30 days) accelerated BAT maturation and permanently remodeled this tissue. As adults, these mice had increased BAT activity and thermogenic capacity relative to controls. However, they also had a blunted acclimation response when subsequently cold exposed as adults (5 °C for 6 weeks). Mice born to cold-exposed mothers (14 °C, entire pregnancy) also showed limited capacity for flexibility as adults, demonstrating that maternal cold stress programs the offspring thermal acclimation response. In contrast, for P. maniculatus adapted to the cold high alpine, BAT maturation rate was unaffected by rearing temperature. However, both postnatal and prenatal cold exposure limited the thermal acclimation response in these cold specialists. Our results suggest a complex interaction between developmental and adult environment, influenced strongly by ancestry, drives thermogenic capacity in the wild.

Encoding of conspecific signals during development can reinforce species barriers as well as set the stage for learning and production of species-typical vocalizations. In altricial songbirds, the development of the auditory system is not complete at hatching, so it is unknown the degree to which recently hatched young can process auditory signals like birdsong. We measured in vivo extracellular responses to song stimuli in a zebra finch (Taeniopygia guttata) secondary auditory forebrain region, the caudomedial nidopallium (NCM). We recorded from three age groups between 13 days post-hatch and adult to identify possible shifts in stimulus encoding that occur before the opening of the sensitive period of song motor learning. We did not find differences in putative cell type composition, firing rate, response strength, and selectivity across ages. Across ages narrow-spiking units had higher firing rates, response strength, accuracy, and trial-by-trial reliability along with lower selectivity than broad-spiking units. In addition, we showed that stimulus-specific adaptation, a characteristic of adult NCM, was also present in nestlings and fledglings. These results indicate that most features of secondary auditory processing are already adult-like shortly after hatching. Furthermore, we showed that selectivity for species-specific stimuli is similar across all ages, with the greatest fidelity in temporal coding in response to conspecific song and domesticated Bengalese finch song, and reduced fidelity in response to owl finch song, a more ecologically relevant heterospecific, and white noise. Our study provides the first evidence that the electrophysiological properties of higher-order auditory neurons are already mature in nestling songbirds.

Key points Maternal shift work increases the risk of pregnancy complications, although its effects on progeny health after birth are not clear. We evaluated the impact of a simulated shift work protocol for one-third, two-thirds or all of pregnancy on the metabolic health of sheep progeny. Simulated shift work had no effect on growth, body size, body composition or glucose tolerance in pre-pubertal or young adult progeny. Glucose-stimulated insulin secretion was reduced in adult female progeny and insulin sensitivity was increased in adult female singleton progeny. The results of the present study do not support the hypothesis that maternal shift work exposure impairs metabolic health of progeny in altricial species. Abstract Disrupted maternal circadian rhythms, such as those experienced during shift work, are associated with impaired progeny metabolism in rodents. The effects of disrupted maternal circadian rhythms on progeny metabolism have not been assessed in altricial, non-litter bearing species. We therefore assessed postnatal growth from birth to adulthood, as well as body composition, glucose tolerance, insulin secretion and insulin sensitivity, in pre-pubertal and young adult progeny of sheep exposed to control conditions (CON: 10 males, 10 females) or to a simulated shift work (SSW) protocol for the first one-third (SSW0-7: 11 males, 9 females), the first two-thirds (SSW0-14: 8 males, 11 females) or all (SSW0-21: 8 males, 13 females) of pregnancy. Progeny growth did not differ between maternal treatments. In pre-pubertal progeny (12-14 weeks of age), adiposity, glucose tolerance and insulin secretion during an i.v. glucose tolerance test and insulin sensitivity did not differ between maternal treatments. Similarly, in young adult progeny (12-14 months of age), food intake, adiposity and glucose tolerance did not differ between maternal treatments. At this age, however, insulin secretion in response to a glucose bolus was 30% lower in female progeny in the combined SSW groups compared to control females (P = 0.031), and insulin sensitivity of SSW0-21 singleton females was 236% compared to that of CON singleton female progeny (P = 0.025). At least in this model, maternal SSW does not impair progeny metabolic health, with some evidence of greater insulin action in female young adult progeny.

KEY POINTS Many mammals are born with immature motor systems that develop through a critical period of postnatal development. In rodents, postnatal maturation of movement occurs from rostral to caudal, correlating with maturation of descending supraspinal and local spinal circuits. We asked whether development of fundamental electrophysiological properties of spinal motoneurons follows the same rostro-caudal sequence. We show that in both regions, repetitive firing parameters increase and excitability decreases with development; however, these characteristics mature earlier in cervical motoneurons. We suggest that in addition to autonomous mechanisms, motoneuron development depends on activity resulting from their circuit milieu. ABSTRACT Altricial mammals are born with immature nervous systems comprised of circuits that do not yet have the neuronal properties and connectivity required to produce future behaviours. During the critical period of postnatal development, neuronal properties are tuned to participate in functional circuits. In rodents, cervical motoneurons are born prior to lumbar motoneurons, and spinal cord development follows a sequential rostro-caudal pattern. Here we asked whether birth order is reflected in the postnatal development of electrophysiological properties. We show that motoneurons of both regions have similar properties at birth and follow the same developmental profile, with maximal firing increasing and excitability decreasing into the third postnatal week. However, these maturative processes occur in cervical motoneurons prior to lumbar motoneurons, correlating with the maturation of premotor descending and local spinal systems. These results suggest that motoneuron properties do not mature by cell autonomous mechanisms alone, but also depend on developing premotor circuits.

The fore‐ and hindlimbs of birds are specialized to perform different functions. The growth patterns of limb bones and their relationship with the ontogeny of locomotion are critical to our understanding of variation in morphological, physiological and life‐history traits within and among species. Unfortunately, the ontogenetic development of limb bones has not been well explored, especially in altricial birds. In this study, we sampled the entire measurements of the pigeon (Columba livia) of individual skeletons, to investigate the ontogenetic allometry of limb bones by reduced major axis regression. The ulna and humerus were found to be positively allometric in relation to body mass, with the ulna growing more rapidly than the humerus. Together with previous data, this suggests that strong positive allometric growth in forelimb bones could be a common trend among diverse Carinatae groups. Hindlimb was dominated by positive allometry, but was variable in the growth of the tarsometatarsus which included three allometric patterns. A greater dorsoventral diameter in the midsection of the humerus and ulna confers superior bending resistance and is ideal for flapping/gliding flight. Shape variation in the midsection of different hindlimb components reflects different mechanical loading, and the markedly inverse trend between the tibiotarsus and tarsometatarsus before 28 days of age also suggests loading change before fledging. Before fledging, the growth of the leg bones was prior to that of the wing bones. This kind of asynchronous development of the fore‐ and hindlimbs was associated with the establishment and improvement of different functions, and with shifts in the importance of different functions over time., Post‐natal growth of limb bones was less explored, especially for altricial birds. We use detailed measurements, including width and depth of the proximal and distal ends, rather than the traditional focus on the length and midshaft width to investigate the ontogenetic allometry of the pigeon (Columba livia). The results of this study provide valuable information in poultry management, and are necessary for kinetics and kinematics studies.

1. In vertebrates, thyroid hormones (THs) play an important role in the regulation of growth, development, metabolism, photoperiodic responses and migration. Maternally transferred THs are important for normal early-phase embryonic development when embryos are not able to produce endogenous THs. Previous studies have shown that variation in maternal THs within the physiological range can influence offspring phenotype. 2. Given the essential functions of maternal THs in development and metabolism, THs may be a mediator of life-history variation across species. 3. We tested the hypothesis that differences in life histories are associated with differences in maternal TH transfer across species. Using birds as a model, we specifically tested whether maternally transferred yolk THs co-vary with migratory status, developmental mode, and traits related to pace-of-life (e.g. basal metabolic rate, maximum lifespan). 4. We collected un-incubated eggs (n = 1-21 eggs per species, median = 7) from 34 wild and captive bird species across 17 families and 6 orders to measure yolk THs (both triiodothyronine, T3 and thyroxine, T4), compiled life-history trait data from the literature, and used Bayesian phylogenetic mixed models to test our hypotheses. 5. Our models indicated that both concentrations and total amounts of the two main forms of THs (T3 and T4) were higher in the eggs of migratory species compared to resident species, and total amounts were higher in the eggs of precocial species, which have longer prenatal developmental periods, than in those of altricial species. However, maternal yolk THs did not show clear associations with pace-of-life related traits, such as fecundity, basal metabolic rate, or maximum lifespan. 6. We quantified interspecific variation in maternal yolk THs in birds and our findings suggest higher maternal TH transfer is associated with the precocial mode of development and migratory status. Whether maternal THs represent a part of the mechanism underlying the evolution of precocial development and migration or a consequence of such life histories is currently unclear. We therefore encourage further studies to explore the physiological mechanisms and evolutionary processes underlying these patterns.

Background In animals with altricial offspring, most growth occurs after birth and may be optimized by post-natal maternal care. Maternal effects on growth may be influenced by individual characteristics of the mothers, such as social status, individual investment strategies and the length of association with offspring. The prolonged juvenile dependence seen in humans is a distinctive life history adaptation, which may have evolved to facilitate sustained somatic and brain growth. In chimpanzees, offspring are typically weaned at approximately 4 years old, yet immature individuals continue to associate with their mothers for up to 10 years beyond weaning. Whether this lengthy association or the individual characteristics of mothers influences growth patterns in this species is not clear. The relationship between urinary creatinine and specific gravity is an established non-invasive measure of muscle mass in humans and chimpanzees. We analysed the urinary creatinine and specific gravity of 1318 urine samples from 70 wild chimpanzees from the Taï Forest, Ivory Coast aged 4 to 15 years. Results We showed a clear increase in urinary creatinine levels with age in both males and females, replicating established growth curves in this species and reaffirming this measure as a reliable proxy for lean body mass. Comparing those who experience maternal loss (orphans) with non-orphan chimpanzees, maternal presence beyond weaning age and into late juvenility positively influenced offspring muscle mass throughout ontogeny such that orphans had significantly less muscle mass than age-matched non-orphans. In age-matched offspring with mothers, those with high-ranking mothers had greater muscle mass. Accounting for variation in muscle mass attributable to maternal presence, we found no effect of maternal investment (length of inter birth interval, from own birth to birth of following sibling) on offspring muscle mass. Conclusion Chimpanzee mothers have an extended and multi-faceted influence on offspring phenotypes. Our results suggest that maternal investment extends beyond lactation and into early adulthood and has clear benefits to offspring physical development. Therefore, prolonged juvenile dependence, although unique in its form in human societies, may be a trait with deeper evolutionary origins.

In marine mammals, muscular development has been identified as a rate-limiting factor in achieving adult dive capacities. This study investigates the rate that myosin heavy chain (MHC) composition matures in a postural and locomotor skeletal muscle for four pinniped species with different lactation lengths: hooded seals, Cystophora cristata; harp seals, Pagophilus groenlandicus; northern fur seals, Callorhinus ursinus, and Steller sea lions, Eumetopias jubatus. The ontogeny of MHC isoform expression was compared with developmental rates of myoglobin concentrations, and aerobic (citrate synthase, β-hydroxyacyl-CoA dehydrogenase) and anaerobic (lactate dehydrogenase) enzyme activities. Within taxonomic families, species with shorter lactation periods had more mature muscles biochemically at birth, and fiber types differentiated earlier during ontogeny (Phocidae: hooded > harp seals, Otariidae: northern fur seals > Steller sea lions). Northern fur seal neonates had the most phenotypically-mature muscles in this study, with no immature MHC isoforms. The relationship between muscle biochemistry and MHC composition became more pronounced with age, and developed to reflect swimming mode and activity levels. In adults, phocids had more slow-twitch oxidative protein in their primary locomotor muscle, the Longissimus dorsi (LD), than otariids which likely reflects oxygen-sparing strategies for the phocids’ longer dives. Conversely, northern fur seal muscles had higher proportions of fast-twitch MHCs in the Pectoralis and LD, likely indicative of this species’ smaller size and higher mass-specific metabolic rates. Thus, muscle phenotype is linked with species life history, and a mismatch between muscle biochemistry and MHC composition at weaning has important implications for the first year of independent foraging in pinniped pups.

Life history theory predicts that physiological and behavioral responsiveness to stress should be delayed in development until the benefits of heightened reactivity outweigh the costs of potentially chronic glucocorticoid levels. Birds often acquire stress-responsiveness at locomotor independence, however, both stress-responsiveness and locomotor ability are delayed in birds with altricial developmental strategies. Parrots (Psittacidae) are extremely altricial, but it is not known whether they also postpone physiological responsiveness to stress until locomotor independence. We quantified individual variation in baseline and stress-induced plasma corticosterone (CORT) concentrations, the main avian glucocorticoid, in wild green-rumped parrotlets (Forpus passerinus) of Venezuela at four stages of nestling development. Parrotlet neonates are very underdeveloped and compete for parental care among extreme sibling size hierarchies, a competitive scenario that might benefit from early hypothalamic-pituitary-adrenal (HPA) functionality. Nestlings that underwent a standardized restraint stress-treatment showed higher average CORT concentrations compared to baseline in all age groups sampled, and exhibited no evidence of age-related changes in the stress response. This is 2 weeks before locomotor independence and earlier than previously documented for altricial species. Results suggest that precocity of HPA function may be advantageous to growth and survivorship in extremely altricial birds.

Milk provides mammalian neonates with nutritional support and passive immunity. This is particularly true in marsupials where young are born highly altricial and lacking many components of a fully functional adaptive immune system. Here we investigated the T cell populations in the mammaries of a lactating marsupial, the gray short-tailed opossum Monodelphis domestica. Immunohistochemistry confirmed the presence of T cells within the opossum mammaries throughout lactation. Results of quantifying transcript abundance for lymphocyte markers are consistent with γδ T cells being the most common T cell type within lactating mammaries. Numbers of γδ T cells appear to peak early during the first postnatal week, and then decline throughout lactation until weaning. In contrast, numbers of αβ T cells and γμ T cells appear to be low to non-existent in the lactating mammaries. The results support an ancient and conserved role of immune cells in the evolution and function of mammalian mammary tissue.

The altricial-precocial spectrum describes the degree of morphological maturation of offspring at the moment of hatching. In fishes, precocial species develop all their structures at early stages of embryogenesis and larvae hatch at an advanced stage of development, while altricial species hatch at a less developed stage. The timing of retinal development also varies significantly between precocial and altricial fish species. Thus, retinal development is completed before hatching in precocial species. In contrast, a relatively simple retina is observed in altricial newborns and the acquisition of the adult retinal features extends until late in larval life. Therefore, retinal maturation at hatching could be considered as a morphological character to describe the developmental mode of fish newborns. Syngnathids fishes hatch with well-developed sensory systems, jaws, and feeding structures and, therefore, they are considered as precocial fish species. Using as a model the retina of two species of syngnathids (Syngnathus typhle and Hippocampus guttulatus), we describe the retinal maturity during different embryological stages and compare the results with previous studies in the retina of other altricial and precocial fish species. This will be done through a review of the pertinent literature, as well as by drawing on our own experience gathered through recent studies on fish retinogenesis. These differences in the maturity of the visual system have implications for the vision-based survival skills during the early life stages after hatching and for the overall ecology and fitness of the species. Fundacao para a Ciencia e a Tecnologia, PortugalPortuguese Foundation for Science and Technology [PTDC/MAR/122616/2010] Spanish Ministerio de Ciencia y TecnologiaSpanish Government [BFU2007-67540] Junta de Extremadura [PRI06A195, GR15158]

Within the substantially different time scales characterizing human and rodent brain development, key developmental processes are remarkably preserved. Shared processes include neurogenesis, myelination, synaptogenesis, and neuronal and synaptic pruning. In general, altricial rodents experience greater central nervous system (CNS) immaturity at birth and accelerated postnatal development compared to humans, in which protracted development of certain processes such as neocortical myelination and synaptic maturation extend into adulthood. Within this generalization, differences in developmental rates of various structures must be understood to accurately model human neurodevelopmental toxicity in rodents. Examples include greater postnatal neurogenesis in rodents, particularly within the dentate gyrus of rats, ongoing generation of neurons in the rodent olfactory bulb, differing time lines of neurotransmitter maturation, and differing time lines of cerebellar development. Comparisons are made to the precocial guinea pig and the long-lived naked mole rat, which, like primates, experiences more advanced CNS development at birth, with more protracted postnatal development. Methods to study various developmental processes are summarized using examples of comparative postnatal injury in humans and rodents.

Epidemiologically, metabolic disorders have garnered much attention, perhaps due to the predominance of obesity. The early postnatal life represents a critical period for programming multifactorial metabolic disorders of adult life. Though altricial rodents are prime subjects for investigating neonatal programming, there is still no sufficiently generalised literature on their usage and methodology. This review focuses on establishing five approach-based models of neonatal rodents adopted for studying metabolic phenotypes. Here, some modelled interventions that currently exist to avoid or prevent metabolic disorders are also highlighted. We also bring forth recommendations, guidelines and considerations to aid research on neonatal programming. It is hoped that this provides a background to researchers focused on the aetiology, mechanisms, prevention and treatment of metabolic disorders.

Made available in DSpace on 2022-04-28T19:42:14Z (GMT). No. of bitstreams: 0 Previous issue date: 2021-06-01 Body mass variation is an effective way to evaluate the effect of seasonality on the physiological status of individuals. The breeding period is critical for altricial birds in terms of energy balance and leads to sex-specific demands in species with asymmetrical division of parental care. However, seasonal body mass variation is still poorly known in Neotropical birds, which share a suit of life-history traits, and sex-related seasonal body mass variation in birds is poorly studied overall. Here, we investigate body mass variation between the sexes in both breeding and non-breeding periods in two suburban populations of a Neotropical bird with asymmetric division of parental tasks, the Pale-breasted thrush (Turdus leucomelas). Using wing length as a covariate in a linear mixed-effects model, we found that thrushes were heavier in the breeding period, and that sexual difference in body mass was only marginally significant (males: 67.62 ± 3.75 g, females: 71.30 ± 4.24 g, n = 218 captures of 152 individuals). However, the interaction between sex and period was significant, and a post hoc test revealed that males were lighter than females only during the breeding period. We suggest that the sex-specific seasonal variation in body mass results from distinct parental care activities performed by each sex during the breeding period. Instituto de Biociências Departamento de Biodiversidade Universidade Estadual Paulista Programa de Pós-Graduação em Ecologia e Recursos Naturais Universidade Federal de São Carlos Instituto de Biociências Departamento de Biodiversidade Universidade Estadual Paulista

In altricial avian species, nutrition can significantly impact nestling fitness by increasing their survival and recruitment chances after fledging. Therefore, the effort invested by parents towards provisioning nestlings is crucial and represents a critical link between habitat resources and reproductive success. Recent studies suggest that the provisioning rate has little or no effect on the nestling growth rate. However, these studies do not consider prey quality, which may force breeding pairs to adjust provisioning rates to account for variation in prey nutritional value. In this 8-year study using black-capped (Poecile atricapillus) and boreal (Poecile hudsonicus) chickadees, we hypothesized that provisioning rates would negatively correlate with prey quality (i.e., energy content) across years if parents adjust their effort to maintain nestling growth rates. The mean daily growth rate was consistent across years in both species. However, prey energy content differed among years, and our results showed that parents brought more food to the nest and fed at a higher rate in years of low prey quality. This compensatory effect likely explains the lack of relationship between provisioning rate and growth rate reported in this and other studies. Therefore, our data support the hypothesis that parents increase provisioning efforts to compensate for poor prey quality and maintain offspring growth rates.

At high altitude (HA), unremitting low oxygen and persistent cold push small mammals close to their metabolic ceilings, leaving limited scope for aerobically demanding activities. However, HA breeding seasons are relatively short and endemic rodents compensate with larger litters than low altitude (LA) conspecifics. Rodent mothers are the sole source of heat and nutrition for altricial offspring and lactation is energetically costly. Thus, it is unclear how HA females balance energy allocation during the nursing period. We hypothesized that HA female rodents invest heavily in each litter to ensure postnatal survival. We measured maternal energetic output and behaviour in nursing deer mice (Peromyscus maniculatus) native to LA (400 m a.s.l.) and HA (4350 m a.s.l.) under control (24°C, 760 mmHg) and cold hypoxia conditions, simulating HA (5°C, 430 mmHg). Strikingly, resting metabolic rates of lactating HA and LA females under cold hypoxia were 70–85% of their maximum aerobic capacity. In cold hypoxia, LA mothers increased both nursing time and milk fat content, however their pups were leaner and severely growth restricted at weaning. HA mothers also increased nursing in cold hypoxia but for far less time than LA mothers. Despite receiving less care, HA pups in cold hypoxia only experienced small growth restrictions at weaning and maintained body composition. As adults, HA mice raised in cold hypoxia had increased aerobic capacity compared to controls. These data suggest that HA mothers prioritize their own maintenance costs over investing heavily in their offspring. Pups compensate for this lack of care, likely by reducing their own metabolic costs during development.

High-altitude environments, characterized by low oxygen levels and low ambient temperatures, have been repeatedly colonized by small altricial mammals. These species inhabit mountainous regions year-round, enduring chronic cold and hypoxia. The adaptations that allow small mammals to thrive at altitude have been well studied in non-reproducing adults; however, our knowledge of adaptations specific to earlier life stages and reproductive females is extremely limited. In lowland natives, chronic hypoxia during gestation affects maternal physiology and placental function, ultimately limiting fetal growth. During post-natal development, hypoxia and cold further limit growth both directly by acting on neonatal physiology and indirectly via impacts on maternal milk production and care. Although lowland natives can survive brief sojourns to even extreme high altitude as adults, reproductive success in these environments is very low, and lowland young rarely survive to sexual maturity in chronic cold and hypoxia. Here, we review the limits to maternal and offspring physiology – both pre-natal and post-natal – that highland-adapted species have overcome, with a focus on recent studies on high-altitude populations of the North American deer mouse (Peromyscus maniculatus). We conclude that a combination of maternal and developmental adaptations were likely to have been critical steps in the evolutionary history of high-altitude native mammals.

Polar bear maternal behavior and neonatal development are difficult to study because maternal dens are secluded and generally inaccessible. As a result, much of what we know comes from polar bears in zoos. Altricial polar bear cubs (litter size 1–3) have limited sensory capacity, sparse fur, and are completely dependent on the female for nourishment, warmth, and protection. Energy for the female’s metabolic requirements and milk production comes from body fat during the 12-week denning period. Because females fast and lactate while caring for neonatal cubs, they lose ~44% of their body mass during the denning period. Maternal attentiveness to cubs accounts for

Female sea otters become sexually mature at 2–4 years of age. Birth may occur throughout the year with a normal litter size of one. The reproductive cycle (period of gestation and dependency) is ~12 months, so female sea otters are normally either pregnant or caring for a pup. After birth, the female provides the pup with high-fat milk from two mammary glands on the lower abdomen. Female sea otters are income breeders and must feed daily to maintain milk production, although small fat reserves, which develop during gestation, may reduce the need to forage during the initial 3 weeks post partum. Maternal behavior, which is influenced by hormones and sensory cues, manifests as vigilance against abiotic or biotic threats to neonates, allogrooming (pup grooming provided by the female), and nursing. Sea otters often give birth on the surface of the water, so there is an immediate threat to the pup from drowning, hypothermia, and predation. Pups are altricial and dependent on the female for care (nursing and grooming) and protection, although maternal behavior changes in synchrony with the pup’s physical and behavioral development. Neonates sleep and nurse most of the day, but new behaviors, such as transiting (i.e., surface swimming), self-grooming, foraging, and interacting, develop over the first 3 months of life. In response, the female exhibits reduced vigilance and enhanced foraging to support the growing pup’s nutritional needs with both milk and prey. At some point, this energetic demand may exceed the female’s foraging ability and cause a decrease in body condition, which could lead to pup abandonment or early weaning in severe cases. This behavior enables the female to survive and raise future offspring and enhance fitness. As a result, pup survival will vary geographically and annually in synchrony with favorable conditions. After females wean or abandon their pups, mating occurs within a few days to weeks, and the cycle repeats.

Despite the importance of ecotourism in species conservation, little is known about the industry’s effects on wildlife. In South Africa, some African penguin (Spheniscus demersus) colonies have become tourist attractions. The species is globally endangered, with population sizes decreasing over the past 40 years. As African penguin chicks are altricial and unable to move away from anthropogenic stressors, it is important to evaluate the effect of tourist activities on baseline glucocorticoid levels as a measure of potential disturbance. Chicks at three study sites within two breeding colonies (Robben Island, Stony Point), with varying levels of exposure to tourism (low/moderate/high) were monitored. Urofaecal samples were collected to determine urofaecal glucocorticoid metabolite (ufGCM) concentrations as an indication of baseline stress physiology. Morphometric measurements were taken to compare body condition between sites. Penguin chicks experiencing low, infrequent human presence had significantly higher mean (± standard deviation) ufGCM levels [1.34 ± 1.70 μg/g dry weight (DW)] compared to chicks experiencing both medium (0.50 ± 0.40 μg/g DW, P = 0.001) and high levels of human presence (0.57 ± 0.47 μg/g DW, P = 0.003). There was no difference in chick body condition across sites. These results suggest that exposure to frequent human activity may induce habituation/desensitization in African penguin chicks. Acute, infrequent human presence was likely an important driver for comparatively higher ufGCM levels in chicks, though several other environmental stressors may also play an important role in driving adrenocortical activity. Nevertheless, as unhabituated chicks experiencing infrequent anthropogenic presence showed significantly higher ufGCM levels, managers and legislation should attempt to minimize all forms of activity around important breeding colonies that are not already exposed to regular tourism. Although the results of this study are crucial for developing enhanced conservation and management protocols, additional research on the long-term effect of anthropogenic activities on African penguin physiology is required.

Maternal odor is known to play an important role in mother-infant-interaction in many altricial species such as rodents. However, we only know very little about its role in early human development. The present study therefore investigated the impact of maternal odor on infant brain responses to emotional expression. We recorded the electroencephalographic (EEG) signal of seven-month-old infants watching happy and fearful faces. Infants in two control groups exposed to no specific odor (control 1) or the odor of a different infant’s mother (control 2) showed the expected EEG fear response. Crucially, this response was markedly absent in the experimental group exposed to their mother’s odor. Thus, infants respond differently to fear signals in the presence of maternal odor. Our data therefore suggest that maternal odor can be a strong modulator of social perception in human infants.

Mercury exposure can disrupt development of the cerebellum, part of the brain essential for coordination of movement through a complex environment, including flight. In precocial birds, such as fowl, the cerebellum develops embryonically, and the chick is capable of leaving the nest within hours of hatching. However, most birds, including all songbirds, are altricial, and spend weeks in the nest between hatching and fledging. The objective of this study was to describe the normal development of the cerebellum in a model altricial songbird so as to determine the effect of exposure to mercury on cerebellar maturation. Adult zebra finch (Taeniopygia guttata) pairs were fed either a control diet, or a diet augmented with one of four treatment-levels of methylmercury (0.3–2.4 μg/g wet weight), and their offspring, the subjects of this study, were fed the same diet by parents. We documented, for the first time, the schedule of cerebellar development in an altricial bird, and compared stages of development among methylmercury-exposed groups. For all treatments of methylmercury, the age of completion of cellular migration was later than for control zebra finches, indicating a delay in cerebellar maturation. Displaced (heterotopic) Purkinje neurons, a pathology typical of methylmercury exposure in developing vertebrate brains, were more numerous in methylmercury-exposed birds, and persisted at least until the age of independence. Delays in maturation of the cerebellum could delay fledging in altricial bird species, with potential serious implications for the fitness of exposed individuals, as predation rates in the nest are often very high.

The tarsometatarsus conformation and foot types in birds are unique traits within vertebrates. We investigate how the tarsometatarsus and the zygodactyl foot are formed during development in the monk parakeet (Myiopsitta monachus). Using bones, whole mount specimens stained for cartilage and bone, and histological sections, we focus on the osteology and morphogenesis of the tarsometatarsus. We also compare the tarsometatarsus development between the altricial monk parakeet with the precocial chicken. The results and conclusions we reached are: (1) the hypotarsus, a character of phylogenetic significance, is monocanaliculate in the adult; (2) digit I retroversion is consequence of the displacement of the articulation site of the metatarsal 1 and its torsion; (3) digit IV retroversion is linked to the development of the trochlea accesoria; (4) in ovo, the ossification and fusion of the metatarsals 2-4 begin in their mid-diaphysis and extends cylindrically to both proximal and distal directions; and (5) the differences in the development of the tarsometatarsus between the monk parakeet and the chicken evidence heterochronies, probably related with their different types of development.

In chicken, corticotropin-releasing hormone (CRH) acts as a thyrotropin (TSH)-releasing factor, mediated by the type 2 CRH receptor (CRHR2) on the thyrotropes of the pituitary gland. It is not known whether CRH also controls TSH release in non-precocial avian species that have a different pattern of thyroidal activity during their life cycle. Therefore, we investigated the TSH-releasing capacity of CRH in an altricial species, the zebra finch ( Taeniopygia guttata ). Cellular localisation of type 1 CRH receptor ( CRHR1 ) and CRHR2 mRNA in the pituitary was determined by in situ hybridisation, combined with immunohistochemical staining of pituitary thyrotropes. In addition, isolated pituitary glands were stimulated with CRH to determine the effect on TSH release. Lastly, the mRNA levels of hormones and receptors involved in the control of thyroidal and adrenal function were measured by qPCR in zebra finch chicks between hatching and fledging, and in adults. Most of the hypophyseal CRHR2 mRNA co-localised with thyrotropes, whereas CRHR1 mRNA was found inbetween thyrotropes. Pituitary glands stimulated in vitro with CRH showed increased secretion of TSH-like activity. Pituitary CRHR2 mRNA expression decreased while pituitary TSHB mRNA and brain CRH mRNA levels increased towards fledging, similar as seen in chicken hatching. These results suggest that CRHR2 expressed on thyrotropes is likely mediating CRH-induced TSH release in altricial avian species like it does in precocial species, and that the increased thyroid hormone levels towards fledging in altricial birds are the result of increased hypothalamic stimulation, in which the thyrotropic activity of CRH may initially play a role.

Avian species show variation in longevity, habitat, physiologic characteristics, and lifetime endocrine patterns. Lifetime reproductive and metabolic function vary. Much is known about the neurobiology of the song system in many altricial birds. Little is known about aging in neural systems in birds. Captive birds often survive beyond the age they would in the wild, providing an opportunity to gain an understanding of the physiologic and neural changes. This paper reviews the available information with the goal of capturing areas of potential investigation into gaps in our understanding of neural aging as reflected in physiologic, endocrine, and cognitive aging.

Bone histology of crown-group birds is a research topic of great interest, permitting insight into the evolution of remarkably high growth rates in this clade and variation across the altricial-precocial spectrum. In this study, we describe microanatomical characteristics of the humerus and femur in partial growth series from 14 crown group birds representing ten major clades (Struthioniformes, Galliformes, Apodiformes, Columbiformes, Charadriiformes, Accipitriformes, Strigiformes, Psittaciformes, Falconiformes, and Passeriformes). Our goals were to: (1) describe the microanatomy of each individual; (2) make inter-and intra-taxonomic comparisons; (3) assess patterns that correspond with developmental mode; and (4) to further parse out phylogenetic, developmental, and functional constraints on avian osteological development. Across taxa, the femoral and humeral tissue of neonates can be broadly characterized as highly-vascularized, disorganized woven bone with great variation in cortical thickness (inter-and intrataxonomically, within an individual specimen, and within a single section). The tissue of precocial chicks is relatively more mature at hatching than in altricial, but other categories along the developmental spectrum were less easy to distinguish, thus we were unable to identify a definitive histological proxy for developmental mode. We did not find evidence to support hypotheses that precocial chicks exclusively have thicker cortices and more mature bone in the femur than the humerus at time of hatching; instead, this is a characteristic of nearly all taxa (regardless of developmental mode), suggesting deep evolutionary origins and the effects of developmental channeling. Bone tissue in adults exhibited unexpected variation, corresponding to differences in body size. Large-bodied birds have cortices of fibrolamellar bone, but organization of tissue increases and vascularity decreases with diminishing body size. The outer circumferential layer (OCL) also appears at earlier growth stages in small-bodied taxa. Thus, while the OCL is indicative of a cessation of appositional growth it is not always indicative of cortical maturity (that is, maximum organization of bony tissue for a given taxon). Small size is achieved by truncating the period of fast growth; manipulation of the timing of offset of bone growth is therefore an important factor in changing growth trajectories to alter adult body size.

Partial mortality in broods of altricial passerines results from various reasons, including deficiency of resources, inadequate parental care, diseases and other external factors. In this study we test two hypotheses concerning the phenomenon of partial mortality in nestling altricial passerines: (1) non-adaptive starvation hypothesis based on the assumption that partial brood losses are caused by starvation and undernourishment and has no adaptive value, and (2) adaptive starvation hypothesis assuming that losses from starvation might be elements of an adaptive strategy of reducing surplus nestlings in broods that are over-large. To test the above hypotheses we divided broods of two tit species Great Tit Parus major and Blue Tit Cyanistes caeruleus into two categories: broods with incomplete, less than 100% fledging success and broods with complete 100% fledging success. Subsequently, we compared physiological performance of nestlings between brood categories using blood hemoglobin concentratio...

What an animal needs to learn to survive is altered dramatically as they change from dependence on the parent for protection to independence and reliance on self-defense. This transition occurs in most altricial animals, but our understanding of the behavioral neurobiology has mostly relied on the infant rat. The transformation from dependence to independence occurs over three weeks in pups and is accompanied by complex changes in responses to both natural and learned threats and the supporting neural circuitry. Overall, in early life, the threat system is quiescent and learning is biased towards acquiring attachment related behaviors to support attachment to the caregiver and proximity seeking. Caregiver-associated cues learned in infancy have the ability to provide a sense of safety throughout lifetime. This attachment/safety system is activated by learning involving presumably pleasurable stimuli (food, warmth) but also painful stimuli (tailpinch, moderate shock). At about the midway point to independence, pups begin to have access to the adult-like amygdala-dependent threat system and amygdala-dependent responses to natural dangers such as predator odors. However, pups have the ability to switch between the infant and adult-like system, which is controlled by maternal presence and modification of stress hormones. Specifically, if the pup is alone, it will learn fear but if with the mother it will learn attachment (10–15 days of age). As pups begin to approach weaning, pups lose access to the attachment system and rely only on the amygdala-dependent threat system. However, pups learning system is complex and exhibits flexibility that enables the mother to override the control of the attachment circuit, since newborn pups may acquire threat responses from the mother expressing fear in their presence. Together, these data suggest that the development of pups’ threat learning system is not only dependent upon maturation of the amygdala, but it is also exquisitely controlled by the environment. Most notably the mother can switch pup learning between attachment to threat learning in a moment’s notice. This enables the mother to navigate pup’s learning about the world and what is threatening and what is safe.

Adult barn owl hearing is acute, but development of this sense is not well understood. We, therefore, measured auditory brainstem responses in barn owls from before the onset of hearing (posthatch day 2, or P2) to adulthood (P69). The first consistent responses were detected at P4 for 1 and 2 kHz, followed by responses to 0.5 and 4 kHz at P9, and 5 kHz at P13. Sensitivity to higher frequencies increased with age, with responses to 12 kHz appearing about 2 months after hatching, once the facial ruff was mature. Therefore, these altricial birds achieve their sensitivity to sound during a prolonged period of development, which coincides with maturation of the skull and facial ruff (Haresign and Moiseff in Auk 105:699-705, 1988).

SYNOPSIS Flight is the defining characteristic of birds, yet the mechanisms through which flight ability develops are only beginning to be understood. Wing-assisted incline running (WAIR) and controlled flapping descent (CFD) are behaviors that may offer significant adaptive benefits to developing birds. Recent research into these forms of locomotion has focused on species with precocial development, with a particularly rich data set from chukar partridge (Alectoris chukar). Here we briefly review the kinematics and aerodynamics of flight development in this species. We then present novel measurements of the development of pectoralis contractile behavior during the ontogenetic transition toward powered flight. To obtain these new empirical data, we used indwelling electromyography (EMG) and sonomicrometry and tested WAIR and CFD in seven age classes of chukar (n = 2-4 birds per age) from 5 days post hatching (dph) to adult (300+ dph). For each age class, we measured muscle activity during maximal performance, which was WAIR at 65° in birds 5 dph, CFD in birds 9 dph, WAIR at 80° in birds 14 dph, level flight in birds 25-61 dph, and ascending flight in adults. We also measured muscle activity during sub-maximal performance in all age classes. Flapping chukar chicks use near-continuous activation of their pectoralis at relatively low electromyography amplitudes for the first 8 days and progress to stereotypic higher-amplitude activation bursts by Day 12. The pectoralis undergoes increasing strain at higher strain rates with age, and length trajectory becomes more asymmetrical with greater variation in contractile velocity within the shortening phase of individual contractions. At 20-25 days (12-15% adult chukar mass), pectoralis activity and locomotor performance approaches that of adults, although strain rate exhibits a temporary decrease at 61 dph concurrent with using newly-replaced primary feathers. To better understand how these patterns relate to the evolution of life-history strategy and locomotion, we encourage future efforts to explore these behaviors in altricial and semi-altricial bird species.

Summary Sleep deprivation during pregnancy is an emerging concern, as it can adversely affect the development of the offspring brain. This study was conducted to evaluate the effects of deprivation of rapid eye movement (REM) sleep during the third term of pregnancy on the sleep–wake profiles of neonates in the Wistar rat model. Sleep–wake patterns were assessed through electrophysiological measures and behavioural observations during postnatal days 1–21 on pups born to REM sleep-deprived dams and control rats. Pups of REM sleep-deprived dams had active sleep that was not only markedly higher in percentage during all the days studied, but also had reduced latency during later postnatal days 15–21. Quiet sleep and wake periods were lower. These factors, along with less frequent but longer sleep–wake cycles, indicated maturational delay in the sleep–wake neural networks. The disruption of time-bound growth of sleep–wake neural networks was substantiated further by the decreased slope of survival plots in the sleep bouts. Examination of altered sleep–wake patterns during early development may provide crucial information concerning deranged neural development in the offspring. This is the first report, to our knowledge, to show that maternal sleep deprivation during pregnancy can delay and impair the development of sleep–wake profile in the offspring.

Reproductive traits provide information about the ways by which available resources are allocated during breeding. We tested for environmental drivers of large scale geographical patterns in assemblage mean clutch size, number of broods and overall reproductive investment per breeding season in European birds. We combined data about geographical distribution with published information about reproductive traits, and calculated mean trait values for avian assemblages occurring in 50x50 km grid cells. In total, we employed data from 499 species and 2059 assemblages. As the time available for breeding and the amount of food limit the reproductive effort, we related the geographical variation in reproductive traits to the length of breeding season, normalized difference vegetation index (NDVI) as a surrogate of resource availability, and its seasonality. Geographical patterns in traits may differ between reproductive modes, thus we performed the analyses separately for altricial Passerines (N=203) and precocial non-passerine species (N=164) and controlled for the effect of taxonomy. Large clutches dominated in areas with high NDVI and, in precocial birds, also in areas with high annual seasonality and a long breeding season. High number of broods and high overall reproductive investment dominated in areas with a long breeding season, and high number of broods was found also in areas with low annual seasonality, but only in precocial species. High overall reproductive investment dominated in highly productive areas and also in areas with low annual seasonality in both groups. The increase in reproductive investment is caused mostly by an increase in the number of broods related to the length of season and partly by increase in clutch size related to NDVI. We found a negative correlation between clutch size and the number of broods in Passerines, which might suggest a trade-off between these traits. Processes behind trait patterns differ between altricial and precocial species. This article is protected by copyright. All rights reserved.

We have shown that female swamp wallabies ovulate, mate, and form a new embryo prepartum thereby continuously supporting conceptuses and young at different development stages before and after birth. This system is unique compared to the normal staged system of reproduction in mammals so that swamp wallabies are normally pregnant and lactating throughout their reproductive life.Reproduction in mammals requires distinct cycles of ovulation, fertilization, pregnancy, and lactation often interspersed with periods of anoestrus when breeding does not occur. Macropodids, the largest extant species of marsupials, the kangaroos and wallabies, have a very different reproductive strategy to most eutherian mammals whereby young are born at a highly altricial stage of development with the majority of development occurring over a lengthy lactation period. Furthermore, the timings of ovulation and birth in some species occurs within a very short interval of each other (sometimes hours). Female swamp wallabies have an oestrous cycle shorter than their pregnancy length and were, therefore, speculated to mate and form a new embryo before birth thereby supporting two conceptuses at different stages of pregnancy. To confirm this, we used high-resolution ultrasound to monitor reproduction in swamp wallabies during pregnancy. Here, we show that females ovulate, mate, and form a new embryo prepartum while still carrying a full-term fetus in the contralateral uterus. This embryo enters embryonic diapause until the newborn leaves the pouch 9 mo later. Thus, combined with embryonic diapause, females are continuously pregnant and lactating at the same time throughout their reproductive life, a unique reproductive strategy that completely blurs the normal staged system of reproduction in mammals.

BackgroundWithin-species skull shape variation of marsupial mammals is widely considered low and strongly size-dependent (allometric), possibly due to developmental constraints arising from the altricial birth of marsupials. However, species whose skulls are impacted by strong muscular stresses – particularly those produced through mastication of tough food items – may not display such intrinsic patterns very clearly because of the known plastic response of bone to muscle activity of the individual. In such cases, allometry may not dominate within-species shape variation, even if it is a driver of evolutionary shape divergence; ordination of shape in a geometric morphometric context through principal component analysis (PCA) should reveal main variation in areas under masticatory stress (incisor region/zygomatic arches/mandibular ramus); but this main variation should emerge from high individual variability and thus have low eigenvalues.ResultsWe assessed the evidence for high individual variation through 3D geometric morphometric shape analysis of crania and mandibles of three species of grazing-specialized wombats, whose diet of tough grasses puts considerable strain on their masticatory system. As expected, we found little allometry and low Principal Component 1 (PC1) eigenvalues within crania and mandibles of all three species. Also as expected, the main variation was in the muzzle, zygomatic arches, and masticatory muscle attachments of the mandibular ramus. We then implemented a new test to ask if the landmark variation reflected on PC1 was reflected in individuals with opposite PC1 scores and with opposite shapes in Procrustes space. This showed that correspondence between individual and ordinated shape variation was limited, indicating high levels of individual variability in the masticatory apparatus.DiscussionOur results are inconsistent with hypotheses that skull shape variation within marsupial species reflects a constraint pattern. Rather, they support suggestions that individual plasticity can be an important determinant of within-species shape variation in marsupials (and possibly other mammals) with high masticatory stresses, making it difficult to understand the degree to which intrinsic constraints act on shape variation at the within-species level. We conclude that studies that link micro- and macroevolutionary patterns of shape variation might benefit from a focus on species with low-impact mastication, such as carnivorous or frugivorous species.

Many endotherms native to cold and hypoxic high altitude (HA) environments have evolved a highly vascularized and aerobic skeletal muscle. This specialized muscle phenotype contributes via shivering to an enhanced capacity for aerobic thermogenesis (cold-induced VO2max). However, it is unclear how selection at HA for shivering thermogenesis acts early in the development of small altricial mammals, which are born with immature skeletal muscles and without the capacity for homeothermic endothermy. We have previously shown that postnatal maturation of brown adipose tissue and non-shivering thermogenesis is delayed in HA native deer mouse pups (Peromyscus maniculatus). To assess if HA adaptation has also altered the developmental program of skeletal muscle and shivering thermogenesis, we used laboratory-reared descendants of deer mice native to low (430 m a.s.l) and high (4350 m a.s.l.) altitude and a low altitude (LA) congeneric outgroup (P. leucopus). We found that LA juveniles were able to shiver robustly at 2 weeks after birth. However, HA juveniles were unable to shiver at this point, resulting in a 30% lower capacity for thermoregulation compared to lowlanders. It was only at 27 days after birth that HA juveniles had established the aerobic muscle phenotype characteristic of HA adults and a superior cold-induced VO2max compared to LA mice of the same age. The capacity for shivering may be delayed in HA mice to allow energy to be allocated to other important processes such as growth.

In birds, the magnitude of the adrenocortical stress response can be down‐regulated during specific life‐history stages. Such modulation likely occurs when the effects of mounting robust corticosterone (Cort) elevations interfere with the normal progression of critical lifecycle activities (e.g. development, molt, migration, reproduction). The Developmental Hypothesis posits that altricial birds should display a “Stress Hyporesponsive Period” during the early post‐natal life stages, characterized by reduced adrenocortical stress responses compared to adult birds and a gradual age‐related increase. Such modulation would allow avoiding the potential deleterious effects that long‐term elevations of circulating Cort might exert on growth and development, when the physiological and behavioral abilities to cope with disturbance are limited. Two proximate hypotheses have been proposed to explain this age‐dependent pattern of Cort secretion. The Maturation Hypothesis proposes a progressive age‐related growth, maturation and enhanced sensitivity to sensory input of the Hypothalamic‐Pituitary‐Adrenal (HPA) axis tissues, whereas the Negative Feedback Attenuation Hypothesis proposes a gradual attenuation in the intensity of the negative feedback in the HPA axis. Here we tested these hypotheses by experimentally inducing negative feedback on the HPA axis via dexamethasone (DEX) treatment in nestling white storks Ciconia ciconia. Nestling age positively affected stress‐induced plasma Cort (STRESS‐Cort) levels during experimental handling and restraint, thus supporting the Developmental Hypothesis. DEX treatment significantly reduced STRESS‐Cort levels compared to saline (SAL) treatment, thus eliciting the expected negative feedback on the HPA axis. However, inter‐ and intra‐individual comparisons indicated no age effects on the intensity of the negative feedback exerted by DEX. Our results do not support the Negative Feedback Attenuation Hypothesis and suggest that progressive Maturation of the HPA axis tissues is the proximate mechanism responsible for age‐related changes in the stress response during avian post‐natal development. We encourage further tests of the proposed proximate mechanisms during migration, breeding and molt., This work was supported by the Spanish Ministry of Economy and Competitiveness and European funds (project CGL2012-32544 to J.B.) and the Spanish Ministry of Agriculture, Food and the Environment (Organismo Autónomo de Parques Nacionales, grant 511/2012 to J.B.), the University of Saskatchewan (Isabel Maria Lopez Martinez Memorial Scholarship to F.T-M), and the Spanish Council of Science CSIC (CSIC Projects 201730I011 and i-link 0564 to J.B.).

Sleep-wake development in postnatal rodent life could reflect the brain maturational stages. As the altricial rodents, rats are born in a very undeveloped state. Continuous sleep recording is necessary to study the sleep-wake cycle profiles. However, it is difficult to realize in infant rats since they rely on periodic feeding before weaning and constant warming and appropriate EEG electrodes. We developed a new approach including two types of EEG electrodes and milk-feeding system and temperature-controlled incubator to make continuously polysomnographic (PSG) recording possible. The results showed that there was no evident difference in weight gaining and behaviors between pups fed through the milk-feeding system and warmed with temperature-controlled incubator and those kept with their dam. Evolutional profiles of EEG and electromyogram (EMG) activities across sleep-wake states were achieved perfectly during dark and light period from postnatal day (P) 11 to P75 rats. The ontogenetic features of sleep-wake states displayed that the proportion of rapid eye movement (REM) was 57.0 ± 2.4% and 59.7 ± 1.7% and non-REM (NREM) sleep was 5.2 ± 0.8% and 4.9 ± 0.5% respectively, in dark and light phase at P11, and then REM sleep progressively decreased and NREM sleep increased with age. At P75, REM sleep in dark and light phase respectively, reduced to 6.3 ± 0.6% and 6.9 ± 0.5%, while NREM correspondingly increased to 37.5 ± 2.1% and 58.4 ± 1.7%. Wakefulness from P11 to P75 in dark phase increased from 37.8 ± 2.2% to 56.2 ± 2.6%, but the change in light phase was not obvious. P20 pups began to sleep more in light phase than in dark phase. The episode number of vigilance states progressively decreased with age, while the mean duration of that significantly increased. EEG power spectra in 0.5–4 Hz increased with age accompanied with prolonged duration of cortical slow wave activity. Results also indicated that the dramatic changes of sleep-wake cycle mainly occurred in the first month after birth. The novel approaches used in our study are reliable and valid for continuous PSG recording for infant rats and unravel the ontogenetic features of sleep-wake cycle.

Studying the timing of the main events of embryonic and fetal development may clarify the strategies adopted by species to maximize neonatal survival and the consequences of these events for their life history. This study describes bone development during the fetal phase of the lowland paca (Cuniculus paca), comparing it with other precocial or altricial species, and its relationship with the species' adaptive strategies. A total of 102 embryos/fetuses obtained over the course of 17 years through collaboration with local subsistence hunters in the Amazon were analyzed. Measurements of mineralization of the axial and appendicular skeletons were performed by ultrasonography using a 10-18-MHz linear transducer. The chronological order of occurrence of mineralization in relation to the total dorsal length (TDL) was: skull (TDL = 4.1 cm); vertebral bodies (TDL = 4.6 cm); scapula, humerus, radius, ulna, ilium, ischium, femur, tibia, and fibula (TDL = 6.7 cm); ribs (TDL = 7.8 cm); clavicle (TDL = 8.5 cm); metacarpi/metatarsi (TDL = 11 cm); phalanges (TDL = 15 cm); tarsus (TDL = 18 cm); patella (TDL = 23 cm); and carpus (TDL = 27.2 cm). Secondary ossification centers first appeared in the femoral distal epiphysis (TDL = 16.6 cm) and tibial proximal epiphysis (TDL = 18.4 cm). Advanced fetuses (TDL > 30 cm, 97% gestational period) presented mineralization in all primary and most secondary centers. Compared to other species, paca neonates have a well-developed skeletal system at birth, which is important for their independent postnatal locomotion. Our results may contribute to the monitoring of bone development in other wild species, helping us to understand their life history, and serving as parameters for comparisons between precocial and altricial mammals.

We describe the process of retinal development in mandarinfish Siniperca chuatsi from larvae to young fish. The developmental characteristics of the retinal structure and related cells were identified. Siniperca chuatsi were found to exhibit an altricial mode of retinal development that required considerable time to be completed after hatching. The retina was classed as a pure cone type during the early developmental stage. In the subsequent developmental stages, however, double cones gradually occupied the majority of the cone cells, while rod cells represented the majority of the photoreceptor cells. The outer segment (OS) of the rod cells were significantly longer compared with other morphological features, the OS of the two kinds of cone cells were significantly elongated and the diameters of the inner segment (IS) and OS of the double cone cells were significantly narrower in the later developmental stages. Combined with the scattered arrangement of cone cells at the different stages, the retina was found to have sacrificed a considerable part of visual acuity in the developmental process. The distribution of cone cells was observed to have gradually become regionalised during development. The findings of the present study also indicated that S. chuatsi have a high photosensitivity under dim light conditions as a result of specialised structures of the OS of photoreceptor cells and an increased number of rod cells. The loose arrangement of the cone mosaic presumably resulted in a poor imaging quality and, to some extent, the regionalisation of the cone-cell distribution compensated for the above shortcomings, which would enhance the ability of S. chuatsi to perceive targets in important directions for effective predation behaviour.

Sarco(endo)plasmic reticulum Ca2+-ATPase (SERCA) is a transmembrane pump critical to muscle calcium cycling during contraction, and SERCA has also been proposed as the basis for a non-shivering thermogenesis mechanism in birds. Despite its potential importance to both shivering and non-shivering thermogenesis, the activity of this transporter has rarely been studied in altricial birds, and never during the developmental transition from ectothermy to endothermy. Here, we describe SERCA activity in the pectoralis muscle and heart ventricle of red-winged blackbird (Agelaius phoeniceus) nestlings, fledglings and adults. Additionally, using a diet manipulation, we tested the hypothesis that muscle SERCA activity is affected by dietary fatty acid composition, as has been shown in some previous studies. In blackbird hearts, SERCA activity increased throughout development and into adulthood, conspicuously jumping higher just prior to fledging. In pectoralis muscle, SERCA activity increased throughout the nestling period, but then declined after fledging, an effect we attribute to remodeling of the muscle from a primarily heat-generating organ to a primarily force-generating organ. SERCA activity of the pectoralis muscle was correlated with the proportion of linoleic acid in muscle phospholipids when including all ages in the control group. However, in diet-manipulated birds, there was no consistent relationship between SERCA activity and muscle membrane fatty acid composition at any tested age (5–9 days old). It is unclear whether SERCA might be affected by developmental changes in fatty acid composition at younger ages.

The bird retina offers an excellent model to investigate the mechanisms that coordinate the morphogenesis, histogenesis, and differentiation of neuron and glial cells. Although these developmental features have been intensively studied in the chicken (Gallus gallus, Linnaeus 1758), a precocial bird species, little is known about retinogenesis in altricial birds. The purpose of this study was to examine the differentiation of retinal cells in the altricial zebra finch (Taeniopygia guttata, Vieillot, 1817) and compare the results with those from previous studies in G. gallus. By using immunohistochemical techniques, the first differentiated TUJ1-/Isl1-positive neuroblasts were detected in the vitreal surface of the neuroblastic layer at later incubation times in T. guttata than in G. gallus (108 h vs 55 h). The immunoreactivity of these early differentiation markers coincided temporo-spatially with the appearance of the first PCNA-negative nuclei. Furthermore, the first visinin-positive photoreceptors (132 h vs 120 h) and the first Prox-1-immunoreactive neuroblasts (embryonic day 7.25 (E7.25) vs E6.5) were also detected at later embryonic stages in the retina of T. guttata than in the retina of G. gallus. At E13, one day before hatching, abundant PCNA- and pHisH3-immunoreactivities were detected in the T. guttata retina, while proliferation was almost absent in the G. gallus retina at perinatal stages. Therefore, these results suggest that cell differentiation in the retina is delayed in the altricial bird compared to precocial birds. Furthermore, the T. guttata retina was not completely developed at hatching, and abundant mitotically active precursor cells of retinal neurons were found, suggesting that retinal neurogenesis was intense at perinatal stages.

Human placentation differs from that of other mammals. A suite of characteristics is shared with haplorrhine primates, including early development of the embryonic membranes and placental hormones such as chorionic gonadotrophin and placental lactogen. A comparable architecture of the intervillous space is found only in Old World monkeys and apes. The routes of trophoblast invasion and the precise role of extravillous trophoblast in uterine artery transformation is similar in chimpanzee and gorilla. Extended parental care is shared with the great apes, and though human babies are rather helpless at birth, they are well developed (precocial) in other respects. Primates and rodents last shared a common ancestor in the Cretaceous period, and their placentation has evolved independently for some 80 million years. This is reflected in many aspects of their placentation. Some apparent resemblances such as interstitial implantation and placental lactogens are the result of convergent evolution. For rodent models such as the mouse, the differences are compounded by short gestations leading to the delivery of poorly developed (altricial) young. Human placentation differs from that of other mammals. A suite of characteristics is shared with haplorrhine primates, including early development of the embryonic membranes and placental hormones such as chorionic gonadotrophin and placental lactogen. A comparable archi-tecture of the intervillous space is found only in Old World monkeys and apes. The routes of trophoblast invasion and the precise role of extravillous trophoblast in uterine artery transformation is similar in chimpanzee and gorilla. Extended parental care is shared with the great apes, and though human babies are rather helpless at birth, they are well developed (precocial) in other respects. Primates and rodents last shared a common ancestor in the Cretaceous period, and their placentation has evolved independently for some 80 million years. This is reflected in many aspects of their plac-entation. Some apparent resemblances such as interstitial implantation and placental lactogens are the result of convergent evolution. For rodent models such as the mouse, the differences are com-pounded by short gestations leading to the delivery of poorly developed (altricial) young.

Conditions experienced early in life can shape brain development and later cognition. Altricial songbirds are particularly vulnerable to early environmental perturbations. Research on "Developmental Stress" in songbirds has addressed how early-life conditions may impair song learning and has been extended to consider other components of adult phenotype. Early-life challenges ranging from ectoparasites to competition with siblings have been shown to compromise song learning and other measures of cognition, as well as behavioral strategy. Here, we examined both the effects of hatching asynchrony and early-life immune system challenge with lipopolysaccharide (LPS) on neophobia, song learning, motoric learning, and spatial cognition in male zebra finches (Taeniopygia guttata). We found that hatch order had a significant impact on motoric and spatial learning, such that later hatched males performed better than first and second hatched birds. In contrast, LPS treatment only impacted motoric learning and neither hatch order nor immune system challenge impacted song quality, song learning accuracy, or neophobia. These results are consistent with a growing body of evidence that conditions early in life can improve cognitive performance at adulthood. Moreover, these findings indicate that hatch order is an important factor to consider in developmental studies in asynchronously hatching birds.

In juvenile and adult animals, including mammals, birds, fishes and a crustacean, competition for food becomes increasingly aggressive as its spatial concentration increases. This ecological relationship has not been investigated in infant animals, although it is thought that broods of precocial chicks of some avian species compete aggressively for food or status in a brood hierarchy. When pairs of common quail broodmates were offered the same amount of ground corn in four spatial concentrations between ages 15 and 59 days, aggression increased progressively with concentration, culminating in an overall 16-fold increase when corn was in a single clump. These results suggest that aggressive defence may increase with spatial concentration of food in precocial chicks generally, and raise the possibility that a similar pattern could occur in some altricial chicks during the transition to independence and in infants of other animal species.

In vertebrates, exposure to stressful stimuli early in development may alter the activity of the hypothalamo-pituitary-adrenal (HPA) axis, with the potential for fitness consequences later in life. For altricial species, whose young rely on their parents for food, warmth, and protection from predators, adult behavior can modify the impact of some stressors on their offspring after birth or hatching. We have shown that single bouts of cooling that normally occur when brooding females leave the nest elevate corticosterone secretion in very young free-living eastern bluebird (Sialia sialis) chicks. Thus, natural variation in maternal brooding patterns can result in differential exposure of offspring to cooling, and also to glucocorticoids, very early in development. We tested the hypothesis that exposure to repeated bouts of cooling (mimicking those that occur normally when females leave the nest) would alter the activity of the chicks' HPA axis later in life. We exposed free-living chicks to either four 18-min bouts of cooling or brooding temperatures (control) during the first week after hatching. Then, just before fledging (i.e., at least 7 d after the cooling treatments had ceased), we assessed their corticosterone responses to restraint. Repeatedly cooled chicks had a significantly lower corticosterone response to restraint than did control chicks but did not differ from controls in other measures of growth and development. Our data suggest that natural variation in maternal brooding patterns, and hence natural variation in the chicks' body temperature, can alter the activity of the HPA axis well beyond the brooding period.

In humans and other altricial species, parental care in early life has profound influences on brain development (Tang, Reeb-Sutherland, Romeo, & McEwen, 2014), gene transcription (Champagne, 2012),...

The ossification sequence of Myiopsitta monachus was determined. Myiopsitta has a similar sequence to other altricial birds, with delayed skeletons compared to precocial species. The hindlimbs ossify before the forelimbs, a condition that could be linked to altriciality. To determine the stability of the sequences of ossification across birds, we selected species of different groups of Aves and used event-pairing method and character mapping on a phylogeny. Our results show that the homogeneity in the development of birds was supported by 56.77% of the character states. Event-pair cracking phylogenetic method was applied to identify sequence heterochronies. Results reveal a high number of heterochronies and show that the long bones in limbs may behave as modules. In Myiopsitta, the ossa ectethmoidale and mesethmoidale ossify early. These bones provide the origin site of the Psittaciformes' novel adductor m. ethmomandibularis, associated with strong bite forces, and its acceleration in the sequence may correspond to the functional hypothesis. Also, the early appearance of some hyoid apparatus elements occurs, and could be related to the development of tongue in Psittaciformes and its role in handling food, and is in concordance with the functional and size hypothesis.

Altricial nestlings in structured families show a diverse array of behavioural mechanisms to compete for food, ranging from signalling scrambles to aggressive interference. Rates of filial infanticide are moderately high in white storks. It has been hypothesized that this unusual behaviour is an adaptive parental response to the absence of efficient mechanisms of brood reduction (aggression or direct physical interference) by nestlings. To test this latter assumption, we analyzed video recordings of 41 complete feeding episodes at 32 broods during the first half of the nestling period, when nestlings complete 90% of growth and chick mortality and size asymmetries are highest. Parents delivered food to all nestlings simultaneously by regurgitating on the nest floor. No direct (bill to bill) feeding was recorded. Senior nestlings were never observed to limit their junior nestlings from eating food, either by aggression or physical interference. Experimental feeding tests revealed that heavier nestlings handled prey items more efficiently and ate food at a higher speed. The high degree of tolerance shown by senior nestlings is unusual among birds with similar ecological and phylogenetic affinities, such as herons. Tolerance by seniors cannot be easily explained by absence of parental favouritism or proximate factors known to affect the occurrence of sibling aggression in other species (rate of food transfer, brood size, hatching asynchrony or length of nestling period).

Lactation is a critical and energetically expensive period of reproduction, especially for female sea otters (Enhydra lutris) and their pups, both of which have resting metabolic rates that are ca. 2.5-fold higher than terrestrial mammals of similar size. The simultaneous energy budgets for female Alaskan sea otters and their pups during the first three months postpartum were calculated based on published activity budgets for wild animals and metabolic rates for specific behaviors of captive sea otters. Pups were classified into three behavioral/size categories: Category 1 (C1) 0–