Extra-pair paternity and song characteristics in the willow warbler Phylloscopus trochilus

Complexity in bird song is often argued to be advantageous in processes of sexual selection, and numerous studies show that characteristics of song are associated with increased success in territory defence or mate attraction. Less evidence exists on the relationship between bird song characteristics and patterns of extra-pair paternity. We tested whether males that suffered from extra-pair paternity differed in song characteristics from males with no extra pair paternity in the willow warbler Phylloscopus trochilus . In the Scottish population that we studied, we found that 23.5% of young were not related to the social father, and that 47% of nests contained at least one extra pair young. Older males were less likely to suffer paternity loss. While song repertoire size was not related to loss of paternity, males with short songs suffered higher paternity loss than males with long songs. Although arrival date is a good correlate of social mate choice in this population, it was not related to extra-pair paternity. These results suggest that females use song length, or a trait correlated with this song characteristic, as a cue for choosing extra-pair partners in this species, or alternatively that variance in the success of mate guarding or female coercion is related to this song variable.     

Do willow warblers Phylloscopus trochilus of northern origin start their autumn migration at an earlier age than their southern conspecifics?

Data on hatching time in the willow warbler for different regions (S Germany, S Sweden and N Sweden) are compared with the median of the corresponding migration diagrams. By doing so it is possible to calculate the age, at which the young of the year start their autumn migration. Our calculated values for the above mentioned regions are 68, 62–66 and 61–62 d, respectively. In our opinion this difference is too small to justify the statement, that northern birds migrate at an earlier age than their southern conspecifics. An alternative explanation is that the German migration diagrams are influenced by the passage of northern migrants, resulting in a later median and an apparently higher age for southern birds.     

Characterization of a divergent chromosome region in the willow warbler Phylloscopus trochilus using avian genomic resources

Genome scans have made it possible to find outlier markers thought to have been influenced by divergent selection in almost any wild population. However, the lack of genomic information in nonmodel species often makes it difficult to associate these markers with certain genes or chromosome regions. Furthermore, the extent of linkage disequilibrium (LD) in the genome will determine the density of markers required to identify the genes under selection. In this study, we investigated a chromosome region in the willow warbler Phylloscopus trochilus surrounding a single marker previously identified in a genome scan. We first located the marker in the assembled genome of another species, the zebra finch Taeniopygia guttata, and amplified surrounding sequences in Fennoscandian willow warblers. Within an investigated chromosome region of 7.3 Mb as mapped to the zebra finch genome, we observed elevated genetic differentiation between a southern and a northern population across a 2.5-Mb interval comprising numerous coding genes. Within the southern and northern populations, higher values of LD were mostly found between SNPs within the same locus, but extended across distantly situated loci when the analyses were restricted to sampling sites showing intermediate allele frequencies of southern and northern alleles. Our study shows that cross-species genome information is a useful resource to obtain candidate sequences adjacent to outlier markers in nonmodel species.     

Weak geographical structure in sperm morphology across the range of two willow warbler Phylloscopus trochilus subspecies in Scandinavia

Sperm morphology is highly diversified among species and at higher taxonomic levels. In birds, there is also increasing evidence of geographical differentiation in sperm traits within species, especially in those with strong sperm competition. Geographical divergences in sperm traits might imply the formation of a reproductive barrier in a speciation process. Here we study sperm morphology variation of willow warblers Phylloscopus trochilus in a geographical context in Scandinavia, across the range of two subspecies that are differentiated in certain genetic markers, morphology and migratory routes. We describe geographical patterns in genotypes (two previously described single‐nucleotide polymorphism (SNP) markers and four polymorphic microsatellites); in wing length, tarsus length and body mass; and in sperm traits of 330 male willow warblers sampled at 33 localities across Norway (58°N–69°N). Birds were on average larger and longer‐winged in the north (spp. acredula) than in the south (spp. trochilus), and showed a sigmoid change in the SNP allele frequencies and body morphology around 65°N. We found no evidence of genetic structuring in the microsatellites. There was no geographical variation in sperm traits across Norway, except that sperm heads were on average longer in the south. Sperm head length was also associated with the two SNP markers, with longer sperm heads for the southern alleles, and midpiece length was weakly associated with one of the SNP markers. Similar among‐male variances in total sperm length among the 33 sampling sites indicate uniform levels of sperm competition across Norway. We conclude that sperm morphology remains a rather undifferentiated trait between the two willow warbler subspecies in Scandinavia, which is consistent with a pattern of a shallow genetic divergence. This indicates that sperm morphology is not a reproductive barrier maintaining the narrow hybrid zone.     

Point of no return – absence of returning birds in the otherwise philopatric willow warbler Phylloscopus trochilus

The return of individual birds to a specific area in successional years, i.e. philopatry, is a remarkable behavioural trait. Here we report on the remarkably reversed: the complete absence of returning individuals of a migratory passerine with otherwise pronounced philopatry. At a high latitude study site in Abisko (68°32?N, 18°80?E) in northern Sweden none of the banded adult willow warblers Phylloscopus trochilus returned to breed 2011–2014. This is in stark contrast to all other reports in the literature and also to our two southern study sites (at 56°56?N, 18°10?E and at 58°94?N, 17°14?E) where 18–38% of adults returned. We investigated this aberrant pattern found in Abisko by analysing three parameters known to influence philopatry; nest predation, breeding success and breeding density, and predicted that absence of philopatry should co‐occur with low breeding success, low breeding density and/or high nest predation. The results did not corroborate this, except that breeding density was lower at Abisko (49–71 pairs km^–2) than at the southern sites (106 pairs km^–2, 101 pairs km^–2). Instead, we suggest the hypothesis that the absence of philopatry is caused by an influx of southern, dispersal‐prone individuals deploying another breeding strategy and that this intra‐specific range expansion is enabled by milder climate and low population density.     

Adaptive features of the ecology and annual cycle of the willow warbler (Phylloscopus trochilus L.) at the northern boundary of the Siberian part of the range

The ecology of the willow warbler in the north of Western Siberia is considered, and the adaptations that enable the spread of this species to the Subarctic are analyzed. It is established that one of the key factors that caused the change in the range of this species is the northward distribution of shrubs and, hence, the biomass of insects (available food items of these birds).     

Heat transfer and the energetic cost of singing by canaries <Emphasis Type="Italic">Serinus canaria</Emphasis>

Sexually selected displays, such as male passerine bird song, are predicted to be costly. However, most measurements calculated the rate of oxygen consumption during singing using respirometry have shown that bird song has a low energetic cost. Since birds are reluctant to sing when enclosed inside a respirometry chamber, the energetic cost of singing could differ from that under more normal circumstances. We used heat transfer modelling, based on thermal images, to estimate the energetic cost of singing by canaries (Serinus canaria) that were not enclosed in respirometry chambers. Metabolic rate calculated from heat transfer modelling was 0.70±0.02 W (N=10 birds) during singing, which was 14±5% greater than during standing (0.62±0.02 W). The energetic cost of singing did not differ significantly from that measured previously using respirometry when we took into account that birds sang for a greater proportion of the time during the current experiments. These conclusions were not sensitive to potential errors in the heat transfer model. Heat transfer modelling would be especially useful to obtain measurements of the energetic cost of activities that animals do not perform readily inside respirometry chambers, such as singing in birds.     

A biomechanical model for size, speed and anatomical variations of the energetic costs of running mammals

Here we propose a model of energetic costs and the muscle-tendon unit function on running mammals. The main goal is to set a simple theoretical framework which gives an understanding of the biomechanical principles behind the size, speed and anatomical variations of the energetic costs of running mammals. The model is a point-like mass withstood by a two-segment leg with an extensor muscle serially attached to a tendon. We considered withstanding body weight during the stance phase as the main role of the muscle-tendon unit during fast locomotion. The ground reaction force dependence on speed and the time of stance phase as well as other biomechanical characteristics were taken from previous empirical studies of running. At the same time, the morphological variations with body mass were taken from empirically well-established allometric equations for mammals. The metabolic cost was estimated from an empirical equation relating metabolic power with muscular force and speed in shortening and stretching. Our model predicts the pattern of mass specific metabolic rate variations with both speed and body mass. It also gives an explanation of the experimentally reported linear inverse relationship between the rate of energy used for running and the time of application of force by the foot to the ground during each stride. It also suggests an explanation of the unusual energy saving adaptations of large macropodids. It provides some predictions on the relationship, between energy costs and muscle-tendon unit characteristics, testable on further experiments.     

The effect of lower limb length on the energetic cost of locomotion: implications for fossil hominins

The consequences of the relatively short lower limbs characteristic of AL 288-1 have been widely discussed, as have the causes and consequences of the short limbs of Neanderthals. Previous studies of the effect of limb length on the energetic cost of locomotion have reported no relationship; however, limb length could have accounted for as much as 19% of the variation in cost and gone undetected (Steudel and Beattie, 1995; Steudel, 1994, 1996). Kramer (1999) and Kramer and Eck (2000) have recently used a theoretical model to predict the effect of the shorter limbs of early hominids, concluding that the shorter limbs may actually have been energetically advantageous. Here, we took an experimental approach. Twenty-one human subjects, of varying limb lengths, walked on a treadmill at 2.6, 2.8, 3.0 and 3.2m.p.h., while their expired gases were analyzed. The subjects walked for 12 minutes at each speed and their rates of oxygen consumption (VO2) over four minutes were averaged to estimate VO2. We also measured each subject's height, weight and lower limb length. Lean body mass and % fat were determined using dual-energy x-ray absorptiometry. ANCOVA with total VO2 at either speed as the dependent variable and total lean mass, % fat and lower limb length as covariates resulted in all three covariates having a significant positive effect on VO2 at p<0.01. Subjects with relatively longer lower limbs had lower locomotor costs. Thus the short lower limbs characteristic of some hominid taxa would have resulted in more costly locomotion, barring some physiological anomaly. The magnitude of this effect is substantial; Neanderthals are estimated to have had locomotor costs 30% greater than those of contemporary anatomically modern humans. By contrast the increase in lower limb length seen in H. erectus would have mitigated the increase in locomotor costs produced by the increase in body size.     

Approaches to the analysis of energetic costs of intraspecific competition for space by rainbow trout (Salmo gairdneri)

Two approaches of estimating energy expended by rainbow trout ( Salmo gairdneri ) on density-dependent activities were tested. Metabolic rates were obtained from growth experiments in laboratory systems. Increases in metabolic rates at higher population densities were attributed to three factors: (1) starvation, (2) increased levels of exercise by subdominant fish were forced to maintain stations in fast water due to saturation of territories of more dominant trout, (3) higher planes of excitation. It was observed that variance of energetic factors of routine metabolism, growth rate, consumption rate, and growth efficiency tended to increase with density. These responses seem to be a good indication of intraspecific competition. Little correlation was observed between density and average daily ration, or density and aggressive behaviour.
Dominant trout, as a group grew faster, grew more efficiently, and contained greater lipid content than the rest of the population at each density. However, strictly defined, energetic advantages were not conferred on a 'peck-right' basis. At higher densities, dominance conferred less benefits than was observed at lower population levels. Space limitation per se was not an influential factor affecting growth rates. Methods used to estimate density-dependent energy costs are evaluated. Both methods give nearly identical estimates.     

Predation risk is unlikely to account for the failure of subordinate speckled warblers Chthonicola sagittata to help at the nest

The predator avoidance hypothesis suggests that the failure of subordinate birds to provision nestlings in communally breeding species is a consequence of increased predation risk. Parents exclude subordinates from the nest area and thus reduce the frequency of predator-attracting visits when the nest is most vulnerable, leading to increased reproductive success. I evaluated this hypothesis for the speckled warbler Chthonicola sagittata , a group-living member of the Pardalotidae in which subordinate males never feed nestlings or fledglings even though they are unrelated to the primary pair, compete for copulations and sometimes sire young in the brood. Parents did not modify provisioning behaviour relative to the risk of nest predation; provisioning rates to 10 d-old nestlings were similar on high and low risk territories. Furthermore, there was no evidence that parents modified the timing of deliveries or adjusted the relative size of deliveries in relation to predation risk. The condition (residual mass) of nestlings differed between high and low risk territories because nestlings on high risk territories had smaller tarsi but similar body mass to those at low risk. Tarsus length was the result of parental phenotype, not modified provisioning behaviour. Given that parents were unresponsive to predation risk, it seems unlikely that predation can account for the failure of subordinates to provision at the nest.     

Evidence for a cost of immunity when the crustacean Daphnia magna is exposed to the bacterial pathogen Pasteuria ramosa

The deployment of the immune system has the obvious potential to ameliorate infection outcomes, but immune responses can also harm hosts by either damaging host tissues or monopolizing resources, leading to enhanced mortality. To gain insight into such a 'cost of immunity' when the crustacean Daphnia magna is challenged with the bacterium Pasteuria ramosa, we measured survivorship among hosts that resisted infection following exposure to various strains and doses of the parasite. In the first of two experiments, these exposures were: single exposures with relatively non-aggressive strains, double exposures with non-aggressive strains, and exposure to aggressive strains. Mortality increased across this gradient of exposure. In a second experiment, we varied the dose of the most aggressive P. ramosa strain and found that resisting infection when a large dose was applied resulted in greater mortality than when a medium or low dose was applied. Assuming that resistance is accomplished with an immune response, and that more aggressive parasites and/or larger doses of parasites are more immunostimulatory, these data are compatible with a cost of immunity. Indeed, in terms of survival, resisting parasites can be more harmful than infection.     

Complementarity of stability patches at the interfaces of protein complexes: Implication for the structural organization of energetic hot spots

A rational design of protein complexes with defined functionalities and of drugs aimed at disrupting protein–protein interactions requires fundamental understanding of the mechanisms underlying the formation of specific protein complexes. Efforts to develop efficient small‐molecule or protein‐based binders often exploit energetic hot spots on protein surfaces, namely, the interfacial residues that provide most of the binding free energy in the complex. The molecular basis underlying the unusually high energy contribution of the hot spots remains obscure, and its elucidation would facilitate the design of interface‐targeted drugs. To study the nature of the energetic hot spots, we analyzed the backbone dynamic properties of contact surfaces in several protein complexes. We demonstrate that, in most complexes, the backbone dynamic landscapes of interacting surfaces form complementary “stability patches,” in which static areas from the opposing surfaces superimpose, and that these areas are predominantly located near the geometric center of the interface. We propose that a diminished enthalpy–entropy compensation effect augments the degree to which residues positioned within the complementary stability patches contribute to complex affinity, thereby giving rise to the energetic hot spots. These findings offer new insights into the nature of energetic hot spots and the role that backbone dynamics play in facilitating intermolecular recognition. Mapping the interfacial stability patches may provide guidance for protein engineering approaches aimed at improving the stability of protein complexes and could facilitate the design of ligands that target complex interfaces.     

Mechanisms responsible for increase in circulating inhibin levels at the time of ovulation in mares

In female mammals, inhibin is secreted by the granulosa cells and selectively inhibits secretion of FSH. Although circulating immunoreactive (ir)-inhibin levels decrease after ovulation as a result of the disappearance of its main source, they abruptly increase at the time of ovulation in mares. To investigate the mechanisms responsible for this increase, 50 ml of equine follicular fluid (eFF) was administered into the abdominal cavity of mares during the luteal phase (eFF, n = 4). One hour after treatment, plasma levels of ir-inhibin and inhibin pro-alphaC (but not estradiol-17beta) were significantly higher in eFF-treated mares than in control mares (n = 4). The hormone profiles in eFF-treated mares were similar to those in mares with the spontaneous or hCG induced ovulations. The present study demonstrates that the release of follicular fluid into the abdominal cavity when the follicle ruptures is responsible for the ovulatory inhibin surge in the mare. These findings also suggest that circulating inhibin pro-alphaC may be useful for determining the time of ovulation in the mare.     

Complexation of iron hexacyanides by cytochrome c. Evidence for electron exchange at the exposed heme edge.

Electrostatic binding of at least two anionic iron hexacyanides to cationic horse heart cytochrome c was demonstrated by equilibrium dialysis measurements. No binding was detected following trifluoroacetylation of all of the 19 lysine residues. Replacement of the natural heme iron ligand methionine 80 by the alternative intrinsic ligand lysine 79 but not the extrinsic ligand imidazole resulted in the loss of one hexacyanide binding site. It is proposed that this site is located at the exposed heme edge and is functional in electron exchange.     

Development of unit cost for the health services offered at King FAHD Central hospital Jazan,Saudi Arabia

BackgroundEfficiency remains one of the most important drivers of decision making in health care system. Fund allocators need to receive structured information about the cost healthcare services from hospitals for better decisions related to resource allocation and budgeting. The objective of the study was to estimate the unit cost for health services offered to inpatients in King Fahd Central hospital (KFCH) Jazan during the financial year 2018.MethodsWe applied a retrospective approach using a top-down costing method to estimate the cost of health care services. Clinical and Administrative departments divided into cost centres, and the unit cost was calculated by dividing the total cost of final care cost centres into the total number of patients discharged in one year. The average cost of inpatient services was calculated based on the average cost of each ward and the number of patients treated.ResultsThe average cost per patient stayed in KFCH was SAR 19,034, with the highest cost of SAR 108,561 for patients in the Orthopedic ward. The average cost of the patient in the Surgery ward, Plastic surgery, Neurosurgery, Medical ward, Pediatric ward and Gynecology ward was SAR 33,033, SAR 29,425, SAR 23,444, SAR 20,450, SAR 9579 and SAR 8636 respectively.ConclusionThis study provides necessary information about the cost of health care services in a tertiary care setting. This information can be used as a primary tool and reference for further studies in other regions of the country. Hence, this data can help to provide a better understanding of tertiary hospital costing in the region to achieve the privatization objective.     

Conditions of the cervix for the increase of plasma levels of 13,14-dihydro-15-keto-prostaglandin F2 alpha after amniotomy at term

Amniotomy was performed in 12 multiparas at term but not in labor. In 6 of these patients (group I), the fetal head and cervix condition were favorable for amniotomy, and in the other 6 (group II), they were not favorable. In all group I patients, a sudden and progressive descent of the fetal head, and onset and progress of labor were noted within 5 hours. Plasma 13,14-dihydro-15-keto-prostaglandin F2 alpha (PGFM) levels increased significantly (P less than 0.05) in 4 of these cases with time. In group II patients, descent of the head was less than that in group I patients (P less than 0.05), and neither strong labor nor rise of PGFM levels was noted within 5 hours. These data support our view that amniotomy at an appropriate time results in the onset and progress of labor, and the rise of plasma PGFM in virtue of the sudden and exponential increase of the head to cervix force, but amniotomy at an inappropriate time does not, because this force is unchanged.     

A change of the hepatocyte population is responsible for the progressive increase of vitellogenin synthetic capacity at and after metamorphosis of Xenopus laevis

Synthesis of the egg yolk precursor protein, vitellogenin, can be induced in adult, but not in larval, amphibian hepatocytes by estrogen treatment. The transition process for this inducibility of hepatocytes during development of Xenopus laevis was examined, using primary cultures of hepatocytes. This was found to occur at about the metamorphic climax of stage 62, although the level of vitellogenin production was very limited at this stage. This low level seemed due neither to insufficient estradiol-17 beta nor to high estrogen-degrading activity. The level of synthesis gradually increased following metamorphosis. Immunohistochemical analysis showed that fewer than 5% of the hepatocytes at stage 62 could be stained with antivitellogenin antibody and that the stained cell fraction subsequently increased gradually for several months after metamorphosis. These findings indicate that adult-type cells capable of synthesizing vitellogenin appear at metamorphosis and then expand their population in the liver during postmetamorphic maturation.     

不同温湿度条件下飞虱虫疠霉对桃蚜生殖力和内禀增长力的影响

对7日龄桃蚜 Myzus persicae用高剂量飞虱虫疠霉Pandora delphacis接种后,饲养于不同温(10～30℃)、湿(74%～100%RH)度组合条件下观察其生殖力及内禀增长力(r_m) 的变化。结果表明,被接种个体生殖力比对照显著降低,且下降幅度受温度、湿度及温湿互作的影响。在各处理组合中,尤其以20～30℃和95%～100%RH的条件下下降更为明显。各处理r_m在不同温度间差异显著,均随温度呈抛物线形变化,25℃下达最高,但不同湿度间差异不显著。与对照相比,被接种蚜的r_m>除10℃与各湿度组合间无显著差异外,其余温湿组合下均显著低于对照。上述分析表明,飞虱虫疠霉的侵染能显著影响桃蚜生殖力及r_m,尤其是在桃蚜繁殖的最适温度范围内效果最为明显。     

The capacity for increase at a low temperature of some Australian populations of the granary weevil,Sitophilus granarius (L.)

The capacity for increase (r_c) of one laboratory and seven field populations of young adult S. granarius from different sites in Australia was determined over thirty-two weeks at 15°C in wheat of 14% moisture content. The mean value of r_c was 0.0704 ±0.0016 and the populations differed significantly with respect to this parameter. Variation in the net reproductive rate (R₀), which averaged 25.4± 1.29, had a greater effect on the value of I_c than did variation in the cohort generation time (T_c), which averaged 45.7±0.37 weeks. The populations did not differ significantly in terms of adult survivorship and 94% of females were still alive at thirty-two weeks. The maximum rate of oviposition, about ten eggs per female per week, was achieved in the eighteen-twenty week age-interval. About 41% of the immature stages survived to adulthood. Estimates of r_c over a twenty-four week period were only slightly lower than those over thirty-two weeks. The capacity for increase at 15°C of a given population was shown to be correlated with its fertility at 29°C, an optimal temperature, and with its body weight rather than with its cold tolerance, as indicated by its chill-coma temperature, or its previous temperature-history. The temperature experienced by the immature stages had a marked influence on r_c in that weevils reared at 15°C and 27°C had respective values of 0.0654 and 0.0786 when subsequently held at 15°C. The differences in the survivorship and fertility of S. granarius and S. oryzae (L.) when both species were reared at 15°C are considered.