Ontogeny of heart rate in embryonic and nestling crows (Corvus corone and Corvus macrorhynchos)

The developmental patterns of mean heart rate (MHR) and instantaneous heart rate (IHR) were investigated in embryos and chicks of altricial Corvuscorone and Corvus macrorhynchos. The MHR of embryos increased linearly with time from 250 beats · min⁻¹ at mid-incubation to 290 beats · min⁻¹ in hatchlings. MHR during the pipping period was maximal, but only marginally higher than in hatchlings. MHR was stable at about 290–300 beats · min⁻¹ during the 1st week after hatching. Spontaneous heart rate (HR) decelerations and accelerations were found in embryos and chicks, disturbing the baseline HR with increasing frequency during development. However, the IHR accelerations developed later and were less frequent than in precocial species. IHR and body temperature decreased during mild cold exposure (23–25 °C) and IHR accelerations were reduced in nestlings during the 1st week. We suggest that the development of parasympathetic control of HR in crows occurs at 60% of incubation, similar to precocial embryos, but sympathetic control may be delayed and suppressed in contrast to precocial embryos. Accepted: 3 March 1999     

Differential calls by Carrion and Hooded Crows (Corvus corone corone and C. c. cornix) in the Alpine hybrid zone

Spectrograms of calls given by Carrion and Hooded Crows in the Italian Alpine hybrid zone have been tested for differences between the two subspecies by univariate and multivariate techniques. Vocalizations of the two Crows are similar and composed of 2–15 caws, each usually made up of two harmonics, separated by short intervals. However, one-way ANOVA and discriminant analysis showed that they were differentiated with respect both to temporal and frequency parameters. We suggest that differences in vocalizations reflect mostly the phylogenetic divergence between Hooded and Carrion Crows. The results are in keeping with previous ecological, morphological and molecular data and would confirm that the present designation of the two Crows as distinct subspecies is correct. Differences in vocalizations, together with others regarding ecology and behaviour, may result in assortative mating and homotypic flocking and, therefore, may contribute to reinforcement of the reproductive isolation between the Carrion and the Hooded Crow in the Alpine hybrid zone.     

Partial albinism and natural selection in the hooded crow Corvus corone cornix

We studied the frequency of partial albinism amongst hooded crows in Norway, mostly in a population at Trondheim. A total of 3461 birds were collected from the field. About 5% of fledged young showed albinoid markings on their third right-hand primary; the frequency decreased to 2% among yearling birds and to only 1% among birds that had acquired their adult plumage. No significant difference was found in relation to sex. The partially albinoid birds were typically small-sized, both as regards their bone structure (ulna, tibia, tarsus) and feathers (wing, tail). Their feathers bore more defective markings than those of birds of normal plumage coloration. A study of the plumage of preserved specimens of hooded crows in the collections of Natural History museums in Norway also indicated that a differential mortality takes place in the wild, such albinoid birds disappearing rapidly from the population.
Experimental interchanges of eggs and hatchlings between nests indicated that the occurrence of partial albinism may be related to the feeding conditions during the nestling stage, rather than to any genetical differences. It is not known, however, whether the former was due to starvation, to an unbalanced diet, or to eating poisonous food.     

Conditioning of gray crows (Corvus cornix L.) in a "giant" eight-arm radial maze

Twelve adult hooded crows (Corvus cornix) were trained to perform a standard radial-maze task in a giant eight-arm outdoor radial maze constructed at the "Chistyi les" Biological Station (Tver oblast) for comparative investigations of spatial memory in birds and mammals. The maze consists of a central part of 250 cm diameter, and has arms of 650 cm length, 170 cm height and 80 cm width. The examined hooded crows showed good task performance. Errors (repeated visits to empty arms) were few in number and were predominantly connected with the tendency to avoid arms close to an observer.     

Comparative phylogeography of two crow species: jungle crow Corvus macrorhynchos and carrion crow Corvus corone

The jungle crow Corvus macrorhynchos Wagler, 1827, and the carrion crow Corvus corone L., 1758, are two closely related species with similar ecological requirements that occupy wide distribution ranges in the Palearctic. We studied patterns of their genetic variation by using sequences of the mitochondrial cytochrome b gene. Corvus macrorhynchos demonstrates a low level of variation and differentiation throughout its range, except for a highly diverged population of Cheju Island (Korea). The haplotype network shows two haplogroups. The island group comprises populations of Sakhalin, Hokkaido, Honshu, and Kyushu, while the haplotypes of Taiwan and Ryukyu Islands proved to be closer to the mainland group, which also includes populations from the Primorye, Khabarovsk, Amur, and Magadan regions in the Russian Far East. This pattern allowed us to develop a phylogeographic hypothesis regarding the two modes of settling of the island populations. Concerning C. corone, the presence of two distinct haplogroups was confirmed within the range of C. c. orientalis. Both haplogroups are found within the same populations in Kamchatka and North Sakhalin, which implies secondary contacts there. Populations of C. corone are found to be rather stable in the western parts of its range, while in the Far East populations experienced recent growth, as was observed for C. macrorhynchos in general. The two species appear to have passed through different evolutionary scenarios.     

Use of sensory cues by fish crows Corvus ossifragus preying on artificial bird nests

How predators locate avian nests is poorly understood and has been subjected to little experimental inquiry. We examined which sensory stimuli were important in the nest-finding behavior of fish crows Corvus ossifragus , a common nest predator in the southeastern United States. Using an array of potted trees in a large enclosure, we presented artificial nests to captive crows and quantified responses to visual, auditory, and olfactory nest cues, and nest position. Partial ranks of nest-treatment preferences were analyzed using log-linear models. Nest visibility significantly increased the likelihood of predation by fish crows and increasing nest height was a marginally significant influence on nest vulnerability; no responses were apparent to auditory or olfactory stimuli. Our findings demonstrate that fish crows are visually-oriented nest predators that may preferentially prey on, or more readily encounter, above-ground nests. Moreover, the experimental design provides a new method for evaluating predator-prey interactions between nests and their predators. This study also illustrates how sensory capabilities of predators can interact with nest types to determine nest predation patterns.     

Reproduction in a population of the hooded crow Corvus cornix

Nest density was stable during the 8-year study period. The mean clutch size was 4.3, and the production of fledglings was 2.0 per pair. Incubation started, on average, when there were still two eggs to be laid and lasted for 18 days. Few eggs were lost but 23% of all broods of nestlings were lost to predators. Single nestlings in a brood were not lost to predators but often starved to death, this applied to 25% of all nestlings. A large proportion of the nestlings were probably infected by Syngamus tracheae , and some of these succumbed during the first weeks after fledging. A comparison with other studies of crow populations shows that the major causes of breeding failure are different in different populations. The causes of variation in breeding success between populations are discussed, especially considering the role of nonbreeding crows.     

Carotenoid-based coloration, condition, and immune responsiveness in the nestlings of a sexually dimorphic bird of prey

In many birds, nestlings exhibit brightly colored traits that are pigmented by carotenoids. Carotenoids are diet limited and also serve important health-related physiological functions. The proximate mechanisms behind the expression of these carotenoid-pigmented traits are still poorly known, especially in nestlings with sexual size dimorphism. In these nestlings, intrabrood competition levels and growth strategies likely differ between sexes, and this may in turn influence carotenoid allocation rules. We used dietary carotenoid supplementation to test whether wild marsh harrier (Circus aeruginosus) nestlings were carotenoid limited and whether carotenoid allocation strategies varied between sexes, which differ in their size and growth strategies. When supplemented, nestlings used the supplemental carotenoids to increase their coloration independently of their sex. We showed that the condition dependence of the carotenoid level and the response to an immune challenge (phytohemagglutinin test) differed between sexes, possibly because sexual size dimorphism influences growth strategies and/or intrabrood competition levels and access to different types of food. In this species, which often feeds on mammals, a trade-off likely exists between food quantity (energy) and quality (carotenoid content). Finally, carotenoid-based coloration expressed in marsh harrier nestlings appeared to be indicative of immune responsiveness rather than condition, therefore potentially advertising to parents nestling quality or value rather than nutritional need.     

Testosterone metabolites differentially maintain adult morphology in a sexually dimorphic neuromuscular system

The lumbar spinal cord of rats contains the sexually dimorphic, steroid‐sensitive spinal nucleus of the bulbocavernosus (SNB). Androgens are necessary for the development of the SNB neuromuscular system, and in adulthood, continue to influence the morphology and function of the motoneurons and their target musculature. However, estrogens are also involved in the development of the SNB system, and are capable of maintaining function in adulthood. In this experiment, we assessed the ability of testosterone metabolites, estrogens and nonaromatizable androgens, to maintain neuromuscular morphology in adulthood. Motoneuron and muscle morphology was assessed in adult normal males, sham‐castrated males, castrated males treated with testosterone, dihydrotestosterone, estradiol, or left untreated, and gonadally intact males treated with the 5α‐reductase inhibitor finasteride or the aromatase inhibitor fadrozole. After 6 weeks of treatment, SNB motoneurons were retrogradely labeled with cholera toxin‐HRP and reconstructed in three dimensions. Castration resulted in reductions in SNB target muscle size, soma size, and dendritic morphology. Testosterone treatment after castration maintained SNB soma size, dendritic morphology, and elevated target muscle size; dihydrotestosterone treatment also maintained SNB dendritic length, but was less effective than testosterone in maintaining both SNB soma size and target muscle weight. Treatment of intact males with finasteride or fadrozole did not alter the morphology of SNB motoneurons or their target muscles. In contrast, estradiol treatment was completely ineffective in preventing castration‐induced atrophy of the SNB neuromuscular system. Together, these results suggest that the maintenance of adult motoneuron or muscle morphology is strictly mediated by androgens. © 2009 Wiley Periodicals, Inc. Develop Neurobiol 70: 206–221, 2010.     

Genetic control of sexually dimorphic axon morphology in Drosophila sensory neurons

The mechanism by which orderly axonal projections are formed during development remains an important and largely unsolved problem in neurobiology. It may be possible to examine the control of axon growth in Drosophila and take advantage of genetic tools to better understand the phenomenon. We show here that some gustatory axons in Drosophila are sexually dimorphic and that genes involved in sex determination control the anatomy of these axons. Both males and females possess gustatory receptors on their legs but males possess more of these receptors than females. More significantly, the axons of the male receptors usually cross the midline and they never do so in females, indicating a central zone of bilateral input in the male but not in the female nervous system. In chromosomal females, expressing a tra or Sxl mutation, the gustatory system is transformed toward the male phenotype. Mutant XX adults resemble normal males externally, because they have more gustatory receptors, and internally, because their axons cross the midline. Gynandromorphs show that the sex of the sensory neuron, and apparently not the central nervous system, controls the growth of the axons. We conclude that the anatomical site of control for this dimorphism is the gustatory neurons.     

Electric organ morphology of Sternopygus macrurus, a wave-type, weakly electric fish with a sexually dimorphic EOD.

In several species of electric fish with a sex difference in their pulse-type electric organ discharge (EOD), the action potential-generating cells of the electric organ (electrocytes) of males are larger and more invaginated compared to females. Androgen treatment of females and juveniles produces a longer-duration EOD pulse that mimics the mature male EOD, with a concurrent increase in electrocyte size and/or membrane infolding. In Sternopygus macrurus, which generates a wave-type EOD, androgen also increases EOD pulse duration. To investigate possible morphological correlates of hormone-dependent changes in EOD in Sternopygus, we examined electric organs from both fish collected in the field, and untreated and androgen-treated specimens in the laboratory. The electrocytes are cigar shaped, with prominent papillae on the posterior, innervated end. Electrocytes of field-caught specimens were significantly larger in all parameters than were electrocytes of specimens maintained in the laboratory. EOD pulse duration and frequency were highly correlated, and were significantly different between the sexes in sexually mature fish. Nevertheless, no sex difference in electrocyte morphology was observed, nor did any parameters of electrocyte morphology correlate with EOD pulse duration or frequency. Further, whereas androgen treatment significantly lowered EOD frequency and broadened EOD pulse duration, there was no difference in electrocyte morphology between hormone-treated and control groups. Thus, in contrast to results from studies on both mormyrid and gymnotiform pulse fish, electrocyte morphology is not correlated with EOD waveform characteristics in the gymnotiform wave-type fish Sternopygus. The data, therefore, suggest that sex differences in EOD are dependent on changes in active electrical properties of electrocyte membranes.     

Estrogen alters excitability but not morphology of a sexually dimorphic neuromuscular system in adult rats

In rats, motoneurons of the spinal nucleus of the bulbocavernosus (SNB) innervate the bulbocavernosus (BC) muscle, which surrounds the base of the penis. The SNB/BC is a sexually dimorphic, steroid-sensitive neuromuscular system, which is critically important in male reproductive behavior. Androgens are necessary for the development, morphology, and function of the SNB/BC system. However, estradiol (E) is also necessary for the development of the SNB/BC system, and E is capable of maintaining BC EMG activity in adulthood. In this study, we used electrophysiological and anatomical methods to examine estrogenic effects on BC EMG activity. We used a modified H-reflex testing method to investigate polysynaptic reflex characteristics in intact males, castrates, and castrates treated short term with estradiol benzoate (EB). Measures of EMG activity, response latency, and spike count were altered in castrates, but maintained in EB-treated castrates to the levels of intact males. Furthermore, estrogenic effects were found in EMG activity that could be isolated to the periphery of the SNB/BC system. BC NMJ size and muscle fiber area have been demonstrated to be hormone sensitive, and we examined these for possible correlates of E's effects on BC EMG activity. BC muscles of intact males, castrates, and short-term EB-treated castrates were fixed and stained with zinc iodide and osmium tetroxide. NMJ size and muscle fiber area did not differ between groups. Together, these data suggest that E treatment results in changes in the neuromuscular periphery that maintain BC EMG activity, but this effect cannot be accounted for by changes in NMJ size or muscle fiber area.     

Metal interactions within and between tissues of nestling rooks Corvus frugilegus

We analyzed interactions of concentrations of 11 essential and nonessential elements, including toxic metals within and between internal organs (liver, kidney and lung), muscles and bones of nestling rooks Corvus frugilegus with acute cadmium contamination and elevated level of lead. The number of statistically significant (P ≤ 0.05) metal-metal relationships (positive/negative) within particular tissues was the highest in the kidney (7/6), following in the bone (9/2), liver (6/4), lung (5/2) and muscle (5/2). We found eight significant interactions of lead with other metals, and only two of cadmium (only with lead and cobalt, which probably mirrored a greater ability of lead (than in the case of cadmium) to functional and kinetic interaction with other metals, and/or inhibiting effect of lead or cadmium in co-accumulation. Furthermore, a positive relationship between concentration of cadmium and lead in the kidney could hint at the key importance of this organ in detoxification of both toxic metals. Analyses of relationships of individual metals between examined tissues show only positive results in the case of copper (n = 8), following potassium (n = 3), zinc and iron (in both cases n = 2) and a single ones for calcium and magnesium. We concluded that the lack of significant relationships of individual toxic metals (cadmium or lead) between analyzed tissues could result from high levels of these metals, which destroyed detoxifying capacity of kidney, and ultimately enabled a rapid bioaccumulation of these inorganic contaminations in all tissues of examined nestlings. An explanation of concentration of toxic metals in tissues of animals, especially in the case of their high level, require an identification of the actual level of essential elements associated with physiological status of organism.     

Prenatal sex ratios and expression of sexually dimorphic traits in three snake species

Variation in intrauterine exposure to hormones associated with variation in the sex of litter mates has well-established and far-reaching effects on sexual development in some mammals. Research on this phenomenon in reptiles is scant, but suggests that lizards may follow the mammalian model whereas snakes may be affected differently. We examined sex-specific expression of four sexually dimorphic traits (tail length, head length, ventral scale count, swimming speed) in three species of snakes (Nerodia sipedon, Thamnophis sirtalis, T. sauritus) relative to litter sex ratios. We found little evidence that traits in either sex were masculinized or feminized in response to variation in litter sex ratio. The one significant result appeared best explained as a statistical artifact attributable to a single litter. Our results indicate that snakes are different from the one lizard studied to date. Unlike previous suggestions that prenatal hormonal mechanisms operate differently in snakes and lizards, however, the difference appears to be that development of sexually dimorphic traits in lizards is affected by litter sex ratios whereas in snakes it is not.     

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.     

No evidence of current sexual selection on sexually dimorphic traits in a bird with high variance in mating success

Sexual dimorphism, particularly in ornamental traits, is likely to have arisen by sexual selection. Most empirical and theoretical studies of sexual dimorphism assume that ongoing sexual selection also maintains the dimorphism. Over four seasons, I measured the sexual selection acting on three sexually dimorphic attributes (epaulet size, body size, and the blackness of the body plumage) of male red-winged blackbirds and found no consistent directional or stabilizing selection on any of them. Correlational selection was also negligible. I used path analysis to explore potential relationships in more detail but found no direct or indirect effects of male traits on either within- or extrapair success. Males who were resident on the marsh for more years had higher within-pair success, primarily because they spent more of the season on their territory. Experimental manipulations of epaulet size and color and the extent of nonblack feathers in the black body plumage had no detectable effect on the number of within-pair mates, paternity, or the number of extrapair offspring sired in nearby territories. These results combine with data from other studies of red-winged blackbirds to suggest that, despite high variation in male mating success and hence a strong opportunity for sexual selection, several morphological attributes that differ between the sexes and vary among males are not under current sexual selection. The possible explanations for why add complexity to our understanding of how sexual selection operates.     

Dark or short nights: differential latitudinal constraints in nestling provisioning patterns of a nocturnally hunting bird species

In diurnal bird species, individuals breeding at high latitudes have larger broods than at lower latitudes, which has been linked to differences in the daily time available for foraging. However, it remains unclear how latitude is linked with parental investment in nocturnal species. Here, we investigate nestling provisioning rates of male Tengmalm's owls in two populations at different latitudes (Czech Republic 50 °N; Finland 63 °N) with the help of cameras integrated into nest boxes. Clutch sizes were smaller in the Czech population (CZ: 5.1 ± 0.1; FIN: 6.6 ± 0.1), but given the higher nestling mortality in the Finnish population, the number of fledglings did not differ between the two populations (CZ: 3.5 ± 0.3; FIN: 3.9 ± 0.2). Nestling provisioning patterns varied within days, over the reproductive season and between the two sites. Males delivered most food at dusk and dawn, having peak delivery rates at sun angles of -11° to -15° at both sites, and males increased the prey delivery rates with higher nestling requirements. Given the longer nights during summer in the Czech Republic compared to Finland, Czech males only showed a small shift in their delivery peak during the night from -17° in April to -14° in July. In contrast, Finnish males shifted their peak of prey delivery from -11° in April to -1° in July. Consequently, Czech males had a longer hunting time per night around midsummer when feeding young (360 min) than Finnish males (270 min). This suggests that nocturnal owl species in northern populations are constrained by the short nights during the breeding season, which can limit the number of young they can raise. Moreover, owls in northern populations are additionally constrained through the unpredictable changes in food availability between years, and both these factors are likely to influence the reproductive investment between populations.     

Genetic identification of neurons controlling a sexually dimorphic behaviour

In the fruit fly Drosophila melanogaster, locomotor activity is sexually dimorphic: female flies constantly modulate their activity pattern whereas males show a steadier, stereotyped walking pace [1]. Here, we mapped the area of the brain controlling this behavioural dimorphism. Adult male Drosophila expressing a dominant feminising transgene in a small cluster of neurons in the pars intercerebralis exhibited a female-like pattern of locomotor activity. Genetic ablation of these neurons prevented the feminisation of the locomotor activity of transgenic males. The results suggest that this cluster of neurons modulates sex-specific activity, but is not involved in initiating fly locomotion. Nor does it control male courtship behaviour, because feminisation of courtship was not correlated with the feminisation of locomotor activity.     

Differential effects of dihydrotestosterone and estrogen on the development of motoneuron morphology in a sexually dimorphic rat spinal nucleus

The rat lumbar spinal cord contains a sexually dimorphic motor nucleus, the spinal nucleus of the bulbocavernosus (SNB), whose motoneurons innnervate perineal muscles involved in copulatory reflexes. Dendritic development of SNB motoneurons is biphasic and androgen dependent. During the first 4 postnatal weeks, SNB dendrites grow exuberantly, and subsequently retract to mature lengths by 7 weeks of age. After early postnatal castration, SNB dendrites fail to grow, and testosterone replacement restores this growth. In other systems, testosterone and its metabolites, dihydrotestosterone and estrogen, are important for somatic and neural sexual differentiation. The purpose of the present study was to examine the effects of castration and dihydrotestosterone or estrogen replacement on the growth of SNB motoneuron somata and dendritic arbors. Male rat pups were castrated on postnatal (P) day 7 and treated daily with either dihydrotestosterone propionate (DHTP; 2 mg) or estradiol benzoate (EB; 100 μg) until P28 or P49. By using cholera toxin horseradish peroxidase (BHRP) histochemistry, the soma size, dendritic length, dendritic extent, and arbor area of BHRP-labeled SNB motoneurons were measured and analyzed. Both DHTP and EB treatment supported the initial exuberant growth of SNB dendrites through P28, but EB treatment was ineffective in maintaining mature, adult lengths at P49. The possible sites of hormone action and functional implications of these hormonal treatments are discussed. 1994 John Wiley & Sons, Inc.