Post-hatching parental care behaviour and hormonal status in a precocial bird

In birds, the link between parental care behaviour and prolactin release during incubation persists after hatching in altricial birds, but has never been precisely studied during the whole rearing period in precocial species, such as ducks. The present study aims to understand how changes in parental care after hatching are related to circulating prolactin levels in mallard hens rearing ducklings. Blood was sampled in hens over at least 13 post-hatching weeks and the behaviour of the hens and the ducklings was recorded daily until fledging. Contacts between hens and the ducklings, leadership of the ducklings and gathering of them steadily decreased over post-hatching time. Conversely, resting, preening and agonistic behaviour of hens towards ducklings increased. Plasma prolactin concentrations remained at high levels after hatching and then fell after week 6 when body mass and structural size of the young were close to those of the hen. Parental care behaviour declined linearly with brood age, showed a disruption of the hen-brood bond at week 6 post-hatching and was related to prolactin concentration according to a sigmoid function. Our results suggest that a definite threshold in circulating prolactin is necessary to promote and/or to maintain post-hatching parental care in ducks.     

Partitioning of metabolizable energy intake in sucking altricial and precocial rodent pups

We partitioned the metabolizable energy intake (MEI) into energy for maintenance (ME_m) and for growth (ME_g) in sucking precocial and altricial rodent pups. Each of the two components includes energy loss due to the heat increment of feeding. ME_m of precocial pups expressed as average daily energy costs or as a proportion of MEI was greater than in altricial pups of similar size and, therefore, less energy was available for growth. Consequently, the overall energy cost (via total MEI) per unit postnatal growth of precocial pups was greater than for altricial pups of similar size. We used the proportion of calculated ME_m to that predicted by body mass as an index of precociality in rodent pups. The proportion of ME_g to MEI in precocial pups was lower than in altricial pups and was inversely related to the index of precociality.     

Ontogeny of thermoregulation in precocial birds

The aim of this paper is to summarise the results of earlier experiments on thermoregulation and heat balance in birds, to present new results concerning thermoregulation during the perinatal period in precocial embryos and to develop a model of the ontogeny of thermoregulation over the whole lifespan of birds. The ontogeny of thermoregulation in precocial birds is characterised by three phases with different efficiency of the system. In the prenatal phase, all control elements of the thermoregulatory system can function, but the efficiency of the system is low. It is postulated that endothermic reactions during the prenatal period do not have a proximate (immediate), but rather an ultimate influence on the efficiency of thermoregulation. They may support adaptivity to expected environmental conditions and may be involved in epigenetic adaptation processes. During the early postnatal phase, the thermoregulatory system develops and matures. Summit metabolism and resting metabolic rate and their thermoregulatory set points increase. Preferred temperature is significantly different during different behavioural activities. The phase of full-blown homeothermy starts at approximately the 10th day of life. It is characterised by an activation order of thermoregulatory control elements and by secondary chemical thermoregulation. The influence of thermal and non-thermal climatic factors on heat production and heat loss may be described by mathematical models.     

The chemosensory support of feeding behavior in precocial and altricial mammals during ontogeny]

The development of the structure and function of chemosensory apparatus of the tongue of mature and immature mammals was studied by scanning electron microscopy and in behavioral tests. Heterochronic development of receptors structures of dorsal surface of the tongue was established. At birth the chemoreceptors of body and root of the tongue were relatively mature in morphological aspects (the number of taste buds with pore). Taste pores in buds of anterior part of the tongue were found immediately at birth in mature and at the third week of postnatal period in immature animals. Behavioral tests found functional immaturity of chemosensory structures. Immature pups were able to recognize flavoured solutions only on the 7-10 day of age, and preference-aversion reactions of mature pups appeared to the third day of age. The data obtained are discussed in the respect of different ways of adaptation to food contacts with environment of the pups of mature and immature animals.     

Locatability of begging calls in nestling altricial birds

Nestling begging calls of altricial species of birds have design features (wide frequency range, abrupt onsets and modulation in amplitude and frequency) that make them easily located by birds and mammals, and so may attract predators to the nest. To be maintained by natural selection, such calls must also be beneficial. It is argued here that sibling competition for food during the early stages of nestling development favours locatability of begging calls, presumably because noisy nestlings attract the attention of parents. In the magpie, Pica pica, begging calls of nestlings have a wider frequency spectrum before the nestlings' eyes have opened, a trait that increases their locatability. A strategy of having locatable calls should spread if favoured by mechanisms that overcome the increased predation risk associated with such calls. Two mechanisms are proposed: increased attenuation of the signals by emphasizing higher frequencies (a feature commonly found in begging calls) and dispersion of energy over a wide time-frequency range, a trait that, because of sound degradation, probably masks the estimate of source distance by predators. This hypothesis agrees with predictions of models of intra-brood conflict: when predation costs are higher, level of solicitation (locatability) should decrease. Hole-nesting species, which have a lower risk of predation, have calls with wider frequency ranges and lower (less attenuable) medium frequencies than those of open-nesting species of a similar weight.     

Timing of incorporation of docosahexaenoic acid into brain and muscle phospholipids during precocial and altricial modes of avian development

We investigated the possibilities that the proportion of docosahexaenoic acid (DHA) in phospholipids of brain and skeletal muscle at hatch, and the ontogenetic timing of the DHA accretion spurt in these tissues, might serve as indices of neonatal functional maturity that discriminate between precocial and altricial avian developmental modes. Comparison of the fatty acid profiles of the initial and residual yolks of two free-living altricial species, the swallow (Hirundo rustica) and the sparrow (Passer domesticus), reveals that, in contrast to precocial birds, there is no preferential uptake of DHA from the yolk during embryonic development. At hatch, the proportions of DHA in brain phospholipid (wt.% of fatty acids) of the swallow and sparrow, at 8.1% and 5.0%, respectively, are far lower than the values (16.9-19.6%) reported for non-altricial species. This reflects a marked difference in the timing of the brain DHA accretion spurt, which occurs during the first half of the embryonic period of precocial birds, but is largely delayed until after hatching in the altricial species. By the time of fledging, the proportion of DHA in the swallow brain phospholipid has increased to 14.3%. For non-altricial birds, the brain DHA concentration at hatch shows little interspecies variation, despite major differences in yolk DHA content. The proportions of DHA in leg muscle phospholipid of the newly hatched swallow and sparrow, at 2.9% and 2.5%, respectively, are far lower than the value (6.7%) for the precocial chicken. Again, this relates to differences in developmental timing, with muscle DHA accretion occurring in the first half of the chicken's embryonic period, whereas, in the swallow, this increase is delayed until after hatching. By the time of fledging in the swallow, DHA forms 9.3% of muscle phospholipid fatty acids, equivalent to the level attained in chicken muscle at the mid-embryo stage. The results indicate a clear distinction between altricial and non-altricial avian species in the timing of tissue DHA accretion during development, presumably reflecting differences in neonatal functional maturity.     

Changes in lipid content during the post-hatching development of the brain of altricial birds

Summary The post-hatching development of the brain was studied in three altricial birds (those whose development is completed after hatching), namely, the domestic pigeon, house swift and house sparrow, to assess the state of maturity of the brain at the time that the fledglings leave the nest. Malurity is related to the process of myelination and, therefore, Sudan Black was chosen as it is a sensitive indicator of myelin lipids. On hatching, it was found that sudanophilia was very low in all three birds, indicating a lack of maturation. However, between 15 and 25 days in the pigeon and swift and between 8 and 14 days in the sparrow, there was a steady increase in Sudan Black staining. By the time the birds left their nests, the staining was similar to that of adults, thus indicating that the process of myelination was complete and that the brains were nature.     

A morphometric study of postnatal synaptogenesis in the neocortex of precocial and altricial rodents]

Quantitative electronmicroscopic studies have been made on the development of synapses in two modally different areas of the brain (V-VI layers of the visual and auditory cortex) in the rat and mouse Acomys cahirinus within first two weeks of their postnatal life. The density of synapses as well the relative amount of different types of synapses (symmetrical, asymmetrical, axo-spinal and synapses with large amounts of synaptic vesicles) were measured. It was shown that only in rats the development of synapses in the visual area usually is faster than in the auditory one.     

On the origin of brood parasitism in altricial birds

The probability that obligate interspecific brood parasitism(OP), among altricial birds evolved directly from the normalbreeding (no parasitism, NP) mode or indirectly through intraspecificnest parasitism (INP) was examined by using maximum-likelihoodand parsimony approaches. We examined the probability of ancestralstates at 24 key nodes in order to test our hypotheses. Thestate of the most basal node in a tree of 565 genera of altricialbirds is equivocal; however, the state probability of NP atthis node is about 5.5-fold more likely than the state of obligateparasite. A similar trend was observed for basal nodes of mostfamilies examined. The INP state was supported only in the Hirundinidae.The high incidence of INP among martins and swallows explainsthis finding. Contrary to our predictions, even in other groupswhere there is a high incidence of INP and OP, such as in thetribe Icteri and the Old World finches, the probability of NPbeing ancestral was very high. We conclude that in all casesbut one (Hirundinidae) obligate, and probably facultative, broodparasitism evolved directly from normal breeding mode ratherthan indirectly through some other form of parasitism.     

Perceptual consequences of early social experience in precocial birds

This article reviews recent work with precocial avian hatchlings demonstrating the important contribution that social experience with conspecifics can make to the development of species-typical perceptual preferences. In particular, experiments on the role of sibling social interaction in the development of early auditory and visually directed maternal preferences are surveyed. Results reveal that young hatchlings denied the opportunity for direct social experience with siblings consistently display auditory and visual preferences different from those shown by hatchlings allowed ongoing experience with their broodmates during the period immediately following hatching. Taken together, the studies reviewed here demonstrate that the perceptual preferences underlying the process of filial imprinting, long thought to be simply ‘innate’ or ‘instinctive’, are sensitive to an array of social factors present in the young bird's posthatching environment. The findings also provide support for the view that the minimum unit for the developmental analysis of species-typical behavior must be the developmental system, comprised of both the developing organism and its specific stimulative environment.     

Early appearance of catecholaminergic neurons in the central nervous system of precocial and altricial avian species

Summary The early appearance of catecholaminergic neurons, as revealed by fluorescence histochemistry, has been determined in the central nervous system of quail, pheasant, and pigeon embryos. The first neuronal assemblies displaying specific fluorescence are the locus coeruleus and the nucleus subcoeruleus ventralis. Taking into account the differences in the length of the prehatching period of these three avian species, the first catecholamine-containing neurons appear earlier in the precocial quail and pheasant than in the altricial pigeon.Investigation supported by grants from the Italian National Research Council (CNR) No 83.02058.04 (R.G.) and No 83.00492.04 (G.C.P.).     

Glucose-6-phosphate dehydrogenase and NADP-linked malate dehydrogenase during posthatching development of brain of altricial birds

Glucose-6-phosphate dehydrogenase and NADP-linked malate dehydrogenase were studied in different areas of the brain of three altricial birds during posthatching development. The birds were pigeon and swift, having a posthatching nestling period of 30 days; and sparrow, having a posthatching nestling period of 14 days. The activity of the two enzymes was high during development. G-6-PD activity may be high because of the need for pentoses in the early part of development and the need for reducing equivalents (NADPH₂) for synthesis of lipids and other compounds in the later stages of development. Malic enzyme activity also seems to be high because of the need for reducing equivalents. The activity of malic enzyme was found to be higher than that of G-6-PD.     

Characteristics of body mass growth in semialtricial and altricial bird species during the nestling period

The dynamics of body mass growth were studied in nestlings of 22 semialtricial and altricial bird species based on materials collected in seven regions of Russia in the years 1976–2013. The bird species belong to four orders and 13 families. The results of the study indicate the nonuniform growth of nestlings in different bird species. Of the species investigated, only seven were found to reach or exceed the mass of adult birds. Over the nestling period, the nestlings of open-nesting species, such as the hooded crow (Corvus cornix), rook (Corvus frugilegus), magpie (Pica pica), fieldfare (Turdus pilaris), song thrush (Turdus philomelos), and goldfinch (Carduelis carduelis), do not reach the weight of adult birds and their growth continues after they leave the nest. In closed-nesting species, only the nestlings of the barn swallow (Hirundo rustica) reach or exceed the definitive mass, whereas the nestlings of the jackdaw (Corvus monedula), starling (Sturnus vulgaris), wryneck (Jynx torquilla), tree sparrow (Passer montanus), and great tit (Parus major) continue to grow after leaving the nest. The body mass of birds on the day of their hatching and before their departure from the nests and the mass of adult birds depend on the nesting type, duration of the nestling period, size groups of species, and their definitive size. The average specific growth rate of body mass and its maximum values for different species are also associated with these factors. The maximum specific growth rate in small-sized and medium-sized bird species was observed on the 0–1st days of life; in large bird species, on the 2nd–4th days. The specific growth rate did not depend on the type of nesting, but it was inversely related to the duration of the nestling period and the definitive sizes of birds.     

Nestling colouration is adjusted to parent visual performance in altricial birds

Hitherto, most of the investigation on the perceptual efficacy of begging signals has dwelled on how patterns of nestling colouration adjust to predominant nest luminosity. However, visual sensitivity of birds varies across species, which raises the question of whether colouration of traits involved in begging displays is adjusted to parent visual capacities. Here, by comparing nestling colouration and visual sensitivity across 22 altricial bird species, we provide a first test of this hypothesis. Firstly, we assessed differences in performance of typical UV‐tuned and violet‐tuned bird eyes when looking at the nestling traits under the light regimes prevailing at their nests. Secondly, while controlling for common ancestry in a comparative approach, we explored variation in colouration of nestlings in relation to parent visual system. The colour discrimination model indicated a general higher performance of the ultraviolet over the violet eye at detecting gape and body skin traits in either open‐ or hole‐nest light conditions. Gape colouration was associated with parental visual system as the nestlings of UVS species displayed more yellow and less pure ultraviolet mouths than the nestlings of VS species. Thus, our results agree with an adaptive parent–offspring communication scenario where the nestlings’ colours tuned the perception capacities of their parents.     

Dark nests and conspicuousness in color patterns of nestlings of altricial birds

Nests of altricial birds exhibit variable spectral properties that may affect the efficacy (conspicuousness) of the colored begging traits that a nestling displays to its parents. Here we explored whether selection for efficient perception has favored the evolution of nestling color designs that maximizes nestling detectability in variable light environments. Visual models were used to estimate how parents perceive the coloration of mouths, flanges, heads, and breasts of nestlings within their nest in 21 species of European birds. We show that the largest chromatic and achromatic contrasts against the nest background appeared for nestling mouths and flanges, respectively. Nestlings of open-nesting species showed a larger general achromatic contrast with the nest than did nestlings of hole-nesting species. However, nestlings of hole nesters showed a more evident achromatic contrast between flanges and other traits than did nestlings of open nesters. In addition, species with larger clutch sizes showed larger general achromatic contrasts with the nest. Gaping traits of open-nesting species contrasting with the nest background were better perceived under rich light regimes than under poor ones. These findings are consistent with a scenario in which selection for nestling detectability in dark environments has favored the evolution of particular achromatic components of gape coloration but also nestling traits that enhance signal efficacy by maximizing color contrasts within a nestling.     

Fledgling sex ratios in relation to brood size in size-dimorphic altricial birds

In six species of dimorphic raptors (females larger than males)and one passerine (males larger than females), the sex ratioat fledging varied systematically with brood size at fledging.In all species the strongest bias toward the smaller sex wasestablished in the largest as well as the smallest broods; amore even distribution of males and females was observed inbroods of intermediate size. We explored a specific differentialmortality explanation for this sex ratio variation. Our hypothesispostulates that variation in mortality is caused by differencesin food demand between broods of the same size, due to theirsex composition. Data from the marsh harrier Circus aeruginosuson gender-related food demand and overall nestling mortalitywere used to predict the frequency of surviving males and femalesat fledging, assuming an even sex ratio at hatching and randommortality with respect to both sexes within broods. The modelquantitatively fits the marsh harrier data well, especiallyin broods originating from large dutches. Although we anticipatethat other mechanisms are also involved, the results supportthe hypothesis of sex-ratio-dependent mortality, differentialbetween broods, as the process generating the observed brood-sizedependence of fledgling sex ratios in sexually dimorphic birds.