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Domestic chickens are less fearful, have a faster sexual development, grow bigger, and lay more eggs than their primary ancestor, the red junglefowl. Several candidate genetic variants selected during domestication have been identified, but only a few studies have directly linked them with distinct phenotypic traits. Notably, a variant of the thyroid stimulating hormone receptor (TSHR) gene has been under strong positive selection over the past millennium, but its function and mechanisms of action are still largely unresolved. We therefore assessed the abundance of the domestic TSHR variant and possible genomic selection signatures in an extensive data set comprising multiple commercial and village chicken populations as well as wild-living extant members of the genus Gallus. Furthermore, by mean of extensive backcrossing we introgressed the wild-type TSHR variant from red junglefowl into domestic White Leghorn chickens and investigated gene expression, hormone levels, cold adaptation, and behavior in chickens possessing either the wild-type or domestic TSHR variant. While the domestic TSHR was the most common variant in all studied domestic populations and in one of two red junglefowl population, it was not detected in the other Gallus species. Functionally, the individuals with the domestic TSHR variant had a lower expression of the TSHR in the hypothalamus and marginally higher in the thyroid gland than wild-type TSHR individuals. Expression of TSHB and DIO2, two regulators of sexual maturity and reproduction in birds, was higher in the pituitary gland of the domestic-variant chickens. Furthermore, the domestic variant was associated with higher activity in the open field test. Our findings confirm that the spread of the domestic TSHR variant is limited to domesticated chickens, and to a lesser extent, their wild counterpart, the red junglefowl. Furthermore, we showed that effects of genetic variability in TSHR mirror key differences in gene expression and behavi, Funding Agencies|Swedish Research Council (VR)Swedish Research Council [2015-05444, 2015-4870, 2017-06218]; European Research Council (ERC)European Research Council (ERC)European Commission [322206 GENEWELL]

Domestic animals are adapted to conditions vastly different from those of their wild ancestors, and this is particularly true for their diets. The most numerous of all domestic species, the chicken, originated from the Red Junglefowl (RJF), a native of subtropical forests in Southeast Asia. Surprisingly however, in domestic chicken breeds, a common haplotype of the beta-carotene oxygenase 2 (BCO2) gene, which is involved in carotenoid metabolism, is introgressed from a related species, the Gray Junglefowl, and has been under strong selective pressure during domestication. This suggests that a hybridization event may have conferred a fitness advantage on chickens carrying the derived allele. To investigate the possible biological function of the introgressed BCO2 allele in chicken, we introgressed the ancestral BCO2 allele into domestic White Leghorn chickens. We measured gene expression as well as carotenoid accumulation in skin and eggs of chickens carrying either the ancestral or the derived BCO2 allele. The derived haplotype was associated with down-regulation of BCO2 in skin, muscle, and adipose tissue, but not in liver or duodenum, indicating that carotenoid accumulation occurred in the tissues with reduced gene expression. Most importantly, we found that hens with the derived BCO2 genotype were capable of allocating stored carotenoids to their eggs, suggesting a functional benefit through buffering any shortage in the diet during egg production. Nevertheless, it is of interest that loss of function mutations in BCO2 gene are prevalent in other domesticates including cows, rabbits, and sheep, and, given the importance of carotenoids in development, reproduction, and immunity, it is possible that derived BCO2 alleles may provide a general mechanism in multiple domestic species to deal with higher demand for carotenoids in an environment with carotenoid shortage in the diet.

Domesticated animals share a unique set of morphological and behavioral traits, jointly referred to as the domesticated phenotype. Striking similarities amongst a range of unrelated domesticated species suggest that similar regulatory mechanisms may underlie the domesticated phenotype. These include color pattern, growth, reproduction, development and stress response. Although previous studies have focused on the brain to find mechanisms underlying domestication, the potential role of the pituitary gland as a target of domestication is highly overlooked. Here, we study gene expression in the pituitary gland of the domesticated White Leghorn chicken and its wild ancestor, the Red Junglefowl. By overlapping differentially expressed genes with a previously published list of functionally important genes in the pituitary gland, we narrowed down to 34 genes. Amongst them, expression levels of genes with inhibitory function on pigmentation (ASIP), main stimulators of metabolism and sexual maturity (TSHB and DIO2), and a potential inhibitor of broodiness (PRLR), were higher in the domesticated breed. Additionally, expression of 2 key inhibitors of the stress response (NR3C1, CRHR2) was higher in the domesticated breed. We suggest that changes in the transcription of important modulatory genes in the pituitary gland can account not only for domestication of the stress response in domestic chickens, but also for changes in pigmentation, development, and reproduction. Given the pivotal role of the pituitary gland in the regulation of multiple shared domesticated traits, we suggest that similar changes in pituitary transcriptome may contribute to the domesticated phenotype in other species as well., Funding agencies: Swedish Research Council (VR) [2015-05444]; Swedish Research Council Formas [2016-00645]; European Research Council (ERC) [322206 GENEWELL]

Ketone bodies such as β-hydroxybutyrate are important indicators of metabolic condition in birds and are linked to a suite of ecologically relevant factors including migratory decision-making, hunger level and ectoparasite load. Portable point-of-care (POC) devices designed to measure ketones in humans offer a cheap and easy solution to field physiologists in comparison with previous laboratory methods; however, their accuracy for use in birds has received scant attention. Here, we assessed the accuracy of a POC ketone meter (FreeStyle Precision Neo, Abbott, IL, USA) using samples from intermittently fed Red Junglefowl Gallus gallus. Although the device overestimated ketone levels in comparison with laboratory-derived values, random error was low and laboratory vs. device values correlated well, indicating that the Precision Neo is of sufficient accuracy for use in the field and is a pragmatic choice for avian physiologists.

Flight capacity is one of the most important innovations in animal evolution; it only evolved in insects, birds, mammals and the extinct pterodactyls. Given that powered flight represents a demanding aerobic activity, an efficient cardiovascular system is essential for the continuous delivery of oxygen to the pectoral muscles during flight. It is well known that the limiting step in the circulation is stroke volume (the volume of blood pumped from the ventricle to the body during each beat), which is determined by the size of the ventricle. Thus, the fresh mass of the heart represents a simple and repeatable anatomical measure of the aerobic power of an animal. Although several authors have compared heart masses across bird species, a phylogenetic comparative analysis is still lacking. By compiling heart sizes for 915 species and applying several statistical procedures controlling for body size and/or testing for adaptive trends in the dataset (e.g. model selection approaches, phylogenetic generalized linear models), we found that (residuals of) heart size is consistently associated with four categories of flight capacity. In general, our results indicate that species exhibiting continuous hovering flight (i.e. hummingbirds) have substantially larger hearts than other groups, species that use flapping flight and gliding show intermediate values, and that species categorized as poor flyers show the smallest values. Our study reveals that on a broad scale, routine flight modes seem to have shaped the energetic requirements of birds sufficiently to be anatomically detected at the comparative level., Funding Agencies|Fondo Nacional de Desarrollo Cientifico y Tecnologico [1130750, 11160271]; Linkopings Universitet; Svenska Forskningsradet Formas; Direccion General de Investigacion Cientifica y Tecnica [CGL2012-36345]; Comision Nacional de Investigacion Cientifica y Tecnologica

Fisheries interactions are the most serious threats for sea turtle populations. Despite the existence of some rescue centres providing post-traumatic care and rehabilitation, adequate treatment is hampered by the lack of understanding of the problems incurred while turtles remain entrapped in fishing gears. Recently it was shown that bycaught loggerhead sea turtles (Caretta caretta) could experience formation of gas emboli (GE) and develop decompression sickness (DCS) after trawl and gillnet interaction. This condition could be reversed by hyperbaric O-2 treatment (HBOT). The goal of this study was to assess how GE alters respiratory function in bycaught turtles before recompression therapy and measure the improvement after this treatment. Specifically, we assessed the effect of DCS on breath duration, expiratory and inspiratory flow and tidal volume (VT), and the effectiveness of HBOT to improve these parameters. HBOT significantly increased respiratory flows by 32-45% while VT increased by 33-35% immediately after HBOT. Repeated lung function testing indicated a temporal increase in both respiratory flow and VT for all bycaught turtles, but the changes were smaller than those seen immediately following HBOT. The current study suggests that respiratory function is significantly compromised in bycaught turtles with GE and that HBOT effectively restores lung function. Lung function testing may provide a novel means to help diagnose the presence of GE, be used to assess treatment efficacy, and contribute to sea turtle conservation efforts.

Flight capacity is one of the most important innovations in animal evolution; it only evolved in insects, birds, mammals and the extinct pterodactyls. Given that powered flight represents a demanding aerobic activity, an efficient cardiovascular system is essential for the continuous delivery of oxygen to the pectoral muscles during flight. It is well known that the limiting step in the circulation is stroke volume (the volume of blood pumped from the ventricle to the body during each beat), which is determined by the size of the ventricle. Thus, the fresh mass of the heart represents a simple and repeatable anatomical measure of the aerobic power of an animal. Although several authors have compared heart masses across bird species, a phylogenetic comparative analysis is still lacking. By compiling heart sizes for 915 species and applying several statistical procedures controlling for body size and/or testing for adaptive trends in the dataset (e.g. model selection approaches, phylogenetic generalized linear models), we found that (residuals of ) heart size is consistently associated with four categories of flight capacity. In general, our results indicate that species exhibiting continuous hovering flight (i.e. hummingbirds) have substantially larger hearts than other groups, species that use flapping flight and gliding show intermediate values, and that species categorized as poor flyers show the smallest values. Our study reveals that on a broad scale, routine flight modes seem to have shaped the energetic requirements of birds sufficiently to be anatomically detected at the comparative level.

Restricted feeding of broiler breeders is required for improved long-term health and welfare. Because feeding frustration and hunger are major welfare concerns during rearing, many suggestions have been made to decrease the negative feelings of hunger while maintaining suitable growth rates and reproductive health. Non-daily ("skip-a-day") feeding schedules are commonly used around the world to increase portion sizes at meal times while restricting intake but these practices are prohibited in many countries due to welfare concerns on fasting days. We compared birds raised on a non-daily feeding schedule (2 non-consecutive fasting days per week, 5:2), previously suggested as a welfare-friendlier non-daily alternative, to birds raised on daily feed restriction. We found signs of increased physiological stress levels in 5:2 birds, including elevated heterophil to lymphocyte ratios (1.00 for 5:2 vs. 0.75 for daily fed at 12 weeks of age), increased adiposity (0.21% lean body weight [LBW] for 5:2 vs. 0.13% LBW for daily fed), and reduced muscle growth (pectoral muscle 5.94% LBW for 5: 2 vs. 6.52% LBW for daily fed). At the same time, 5:2 birds showed signs of lower anxiety before feeding times (activity was reduced from 10.30 in daily fed to 4.85) which may be a result of the lower feed competition associated with larger portion sizes. Although we found no difference in latency to first head movement in tonic immobility between the treatments (136.5 s on average for both groups), 5:2 birds generally showed more interest in a novel object in the home pen which indicated increased risk taking and reduced fear while fasting. The 5:2 birds in this study showed no signs of learning the feeding schedule, and this unpredictability may also increase stress. Taken together, the effects of 5:2 vs. daily feed restriction on the welfare of broiler breeder pullets remain inconclusive and differ between feeding and fasting days. In addition to reducing stress by minimizing the number, Funding Agencies|Swedish research council Formas [2013-293]; Swedish Centre of Excellence in Animal Welfare Science

The ability to regulate food intake is critical to survival. The hypothalamus is central to this regulation, integrating peripheral signals of energy availability. Although our understanding of hunger in rodents is advanced, an equivalent understanding in birds is lacking. In particular, the relationship between peripheral energy indices and hypothalamic hunger peptides, agouti-related protein (AgRP), proopiomelanocortin (POMC) and neuropeptide Y (NPY) is poorly understood. Here, we compare AgRP, POMC and NPY RNA levels in the hypothalamus of Red Junglefowl chicks raised under ad libitum, chronic restriction and intermittent feeding regimens. Hypothalamic gene expression differed between chronically and intermittently restricted birds, confirming that different restriction regimens elicit different patterns of hunger. By assessing the relationship between hypothalamic gene expression and carcass traits, we show for the first time in birds that AgRP and POMC are responsive to fat-related measures and therefore represent long-term energy status. Chronically restricted birds, having lower indices of fat, show elevated hunger according to AgRP and POMC. NPY was elevated in intermittently fasted birds during fasting, suggesting a role as a short-term index of hunger. The different physiological and neuroendocrine responses to quantitative versus temporal feed restriction provide novel insights into the divergent roles of avian hunger neuropeptides., Funding Agencies|Swedish research council Formas [2013-293]; Swedish Centre of Excellence in Animal Welfare Science; Carl Tryggers Stiftelse for Vetenskaplig Forskning

Uniform growth is a desirable trait in all large-scale animal production systems because it simplifies animal management and increases profitability. In parental broiler flocks, so-called broiler breeders, low growth uniformity is largely attributed to the feed competition that arises from quantitatively restricted feeding. As feed restriction is crucial to maintaining healthy and fertile breeders, several practices for reducing feed competition and the associated growth heterogeneity have been suggested and range from nutrient dilution by increasing fiber content in feed to intermittent fasting with increased portion size (“skip a day”), but no practice appears to be entirely effective. The fact that a large part of the heterogeneity remains even when feed competition is minimized suggests that some growth variation is caused by other factors. We investigated whether this variation arises during embryonic development (as measured by size at hatch) or during posthatch development by following the growth and body composition of birds of varying hatch sizes. Our results support the posthatch alternative, with animals that later grow to be small or large (here defined as >1 SD lighter or heavier than mean BW of the flock) being significantly different in size as early as 1 d after gaining access to feed (P < 0.05). We then investigated 2 possible causes for different postnatal growth: that high growth performance is linked 1) to interindividual variations in metabolism (as measured by cloacal temperature and verified by respirometry) or 2) to higher levels of social motivation (as measured in a social reinstatement T-maze), which should reduce the stress of being reared in large-scale commercial flocks. Neither of these follow-up hypotheses could account for the observed heterogeneity in growth. We suggest that the basis of growth heterogeneity in broiler breeder pullets may already be determined at the time of hatch in the for

Mass transport can be generally defined as movement of material matter. The circulatory system then is a biological example given its role in the movement in transporting gases, nutrients, wastes, and chemical signals. Comparative physiology has a long history of providing new insights and advancing our understanding of circulatory mass transport across a wide array of circulatory systems. Here we focus on circulatory function of nonmodel species. Invertebrates possess diverse convection systems; that at the most complex generate pressures and perform at a level comparable to vertebrates. Many invertebrates actively modulate cardiovascular function using neuronal, neurohormonal, and skeletal muscle activity. In vertebrates, our understanding of cardiac morphology, cardiomyocyte function, and contractile protein regulation by Ca2+ highlights a high degree of conservation, but differences between species exist and are coupled to variable environments and body temperatures. Key regulators of vertebrate cardiac function and systemic blood pressure include the autonomic nervous system, hormones, and ventricular filling. Further chemical factors regulating cardiovascular function include adenosine, natriuretic peptides, arginine vasotocin, endothelin 1, bradykinin, histamine, nitric oxide, and hydrogen sulfide, to name but a few. Diverse vascular morphologies and the regulation of blood flow in the coronary and cerebral circulations are also apparent in nonmammalian species. Dynamic adjustments of cardiovascular function are associated with exercise on land, flying at high altitude, prolonged dives by marine mammals, and unique morphology, such as the giraffe. Future studies should address limits of gas exchange and convective transport, the evolution of high arterial pressure across diverse taxa, and the importance of the cardiovascular system adaptations to extreme environments. © 2017 American Physiological Society. Compr Physiol 7:17-66, 2017., Funding agencies: NSF CAREER award [IBN IOS-0845741]; Linkoping University

Organic chicken production is currently experiencing a boom in Sweden, but has only recently gained access to a commercial slow-growing broiler strain, the Rowan Ranger. While these chickens are reported to grow at rates better suited to organic production than previously available strains, anecdotal evidence suggested increased fearfulness in this strain and concerns were raised that this may deter the birds from accessing the range. We subjected Rowan Rangers and fast-growing Ross 308 raised on the same organic farm to a set of behavioural tests while also tracking activity and outdoor use. The results suggest that although Rowan Rangers indeed show longer durations of tonic immobility, an index of predator-related fear, both strains made an outdoor visit on average once every 1.5 h and typically spent around 5% of their days in the range. Overall however, our results support the transition to Rowan Rangers in organic chicken production in Sweden., Funding Agencies|Swedish research council Formas [2013-293]; Linkopings Universitet; Formas

The ventricles of non-crocodilian reptiles are incompletely dividedand provide an opportunity for mixing of oxygen-poor blood andoxygen-rich blood (intracardiac shunting). However, both cardiacmorphology and in vivo shunting patterns exhibit considerableinterspecific variation within reptiles. In the present study, wedevelop an in situ double-perfused heart approach to characterisethe propensity and capacity for shunting in five reptile species: theturtle Trachemys scripta, the rock python Python sebae, the yellowanaconda Eunectes notaeus, the varanid lizard Varanusexanthematicus and the bearded dragon Pogona vitticeps. Tosimulate changes in vascular bed resistance, pulmonary andsystemic afterloads were independently manipulated and changesin blood flow distribution amongst the central outflow tracts weremonitored. As previously demonstrated in Burmese pythons, rockpythons and varanid lizards exhibited pronounced intraventricularflow separation. As pulmonary or systemic afterload was raised, flowin the respective circulation decreased. However, flow in the othercirculation, where afterload was constant, remained stable. Thiscorrelates with the convergent evolution of intraventricular pressureseparation and the large intraventricular muscular ridge, whichcompartmentalises the ventricle, in these species. Conversely, inthe three other species, the pulmonary and systemic flows werestrongly mutually dependent, such that the decrease in pulmonaryflow in response to elevated pulmonary afterload resulted inredistribution of perfusate to the systemic circuit (and vice versa).Thus, in these species, the muscular ridge appeared labile and bloodcould readily transverse the intraventricular cava. We conclude thatrelatively minor structural differences between non-crocodilianreptiles result in the fundamental changes in cardiac function.Further, our study emphasises that functionally similar intracardiacflow separation evolved independently in lizards (varanids) andsnakes (pytho, Funding agencies: Danish Research Council (Det Frie Forskningsrad \ Natur og Univers); Swedish Research Council (Vetenskapsradet)

Since most animal species have been recognized as sentient beings, emotional state may be a good indicator ofwelfare in animals. The goal of this study was to manipulate the environment of nine beagle research dogs tohighlight physiological responses indicative of different emotional experiences. Stimuli were selected to be amore or a less positive food (meatball or food pellet) or social reward (familiar person or less familiar person).That all the stimuli were positive and of different reward value was confirmed in a runway motivation test.Dogs were tested individually while standing facing a display theatre where the different stimuli could beshown by lifting a shutter. The dogs approached and remained voluntarily in the test system. They were testedin four sessions (of 20 s each) for each of the four stimuli. A test session consisted of four presentation phases(1st exposure to stimulus, post exposure, 2nd exposure, and access to reward). Heart rate (HR) and heart ratevariability (HRV) responses were recorded during testing in the experimental room and also when lying restingin a quiet familiar room. A newmethod of ‘stitching’ short periods of HRV data together was used in the analysis.When testing different stimuli, no significant differenceswere observed in HR and LF:HF ratio (relative power inlow frequency (LF) and the high-frequency (HF) range), implying that the sympathetic tone was activated similarlyfor all the stimuli and may suggest that dogs were in a state of positive arousal. A decrease of HF was associatedwith the meatball stimulus compared to the food pellet and the reward phase (interacting with the personor eating the food) was associated with a decrease in HF and RMSSD (root mean square of successive differencesof inter-beat intervals) compared to the preceding phase (looking at the person or food). This suggests that parasympatheticdeactivation is associated with a more positive emotional state in the dog. A similar reduction in HFandRMSSDwas fo, Funding agencies: Swedish University of Agricultural Sciences [50200]; Swedish Research Council Formas [221-2010-35]

Electroactive biomaterials are widely explored as bioelectrodes and as scaffolds for neural and cardiac regeneration. Most electrodes and conductive scaffolds for tissue regeneration are based on synthetic materials that have limited biocompatibility and often display large discrepancies in mechanical properties with the surrounding tissue causing problems during tissue integration and regeneration. This work shows the development of a biomimetic nanocomposite material prepared from self-assembled collagen fibrils and silver nanowires (AgNW). Despite consisting of mostly type I collagen fibrils, the homogeneously embedded AgNWs provide these materials with a charge storage capacity of about 2.3 mC cm(-2) and a charge injection capacity of 0.3 mC cm(-2), which is on par with bioelectrodes used in the clinic. The mechanical properties of the materials are similar to soft tissues with a dynamic elastic modulus within the lower kPa range. The nanocomposites also support proliferation of embryonic cardiomyocytes while inhibiting the growth of both Gram-negative Escherichia coli and Gram-positive Staphylococcus epidermidis. The developed collagen/AgNW composites thus represent a highly attractive bioelectrode and scaffold material for a wide range of biomedical applications., Funding Agencies|Linkoping University; Swedish Foundation for Strategic Research (SSF).The previous status of this article was Manuscript and the working title was Collagen-Silver Nanowire Composites as Electrically Activeand Antibacterial Scaffolds for Embryonic Cardiac Cell Proliferation

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