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The aerial flocking of birds, or murmurations, has fascinated observers while presenting many challenges to behavioral study and simulation. We examine how the periphery of murmurations remain well bounded and cohesive. We also investigate agitation waves, which occur when a flock is disturbed, developing a plausible model for how they might emerge spontaneously. To understand these behaviors a new model is presented for orientation-based social flocking. Previous methods model inter-bird dynamics by considering the neighborhood around each bird, and introducing forces for avoidance, alignment, and cohesion as three dimensional vectors that alter acceleration. Our method introduces orientation-based social flocking that treats social influences from neighbors more realistically as a desire to turn, indirectly controlling the heading in an aerodynamic model. While our model can be applied to any flocking social bird we simulate flocks of starlings, Sturnus vulgaris, and demonstrate the possibility of orientation waves in the absence of predators. Our model exhibits spherical and ovoidal flock shapes matching observation. Comparisons of our model to Reynolds' on energy consumption and frequency analysis demonstrates more realistic motions, significantly less energy use in turning, and a plausible mechanism for emergent orientation waves., Competing Interests: Declaration of competing interest I declare that I have no known competing financial interests or personal relationships that could have influenced the work reported in this article. I declare that I have no financial or non-financial interests which may be considered as a conflict of interest with this work. This work was developed as independent research without external funding., (Copyright © 2024 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Exploring the evolutionary architecture of female sexual traits and their potential evolvability is important to understand their possible role as postmating sexual signals. Egg colouration has been proposed to be one of these postmating sexual signals, honestly advertising female quality in birds, especially in blue-green laying species. In this study, we used an animal model in a Bayesian framework to estimate the evolvability of multiple descriptors of blue-green egg colouration and egg size in a wild long-term monitored population of spotless starlings (Sturnus unicolor). Our results show low to moderate heritability (h2 = 0.31-0.44) for 3 egg colour descriptors (blue-green chroma, chroma, and lightness) and egg size. Using the coefficient of additive genetic variance (CVA) and the evolvability (IA) as proxies of evolutionary potential of all components of this trait, we found low values of CVA for all these variables, suggesting a small evolutionary potential of these phenotypic traits, contrasting to previous results reported in another blue-green egg laying species. Our results indicate a modest raw genetic material of this trait on which sexual selection can act upon and, therefore, a small probability for these traits to respond easily to selection., (© The Author(s) 2024. Published by Oxford University Press on behalf of the European Society of Evolutionary Biology.)

Rough-and-tumble play in juvenile rats and song in flocks of adult songbirds outside a breeding context (gregarious song) are two distinct forms of non-sexual social behavior. Both are believed to play roles in the development of sociomotor skills needed for later life-history events, including reproduction, providing opportunities for low-stakes practice. Additionally, both behaviors are thought to be intrinsically rewarded and are associated with a positive affective state. Given the functional similarities of these behaviors, this study used RNA-sequencing to identify commonalities in their underlying neurochemical systems within the medial preoptic area. This brain region is implicated in multiple social behaviors, including song and play, and is highly conserved across vertebrates. DESeq2 and rank-rank hypergeometric overlap analyses identified a shared neurotranscriptomic profile in adult European starlings singing high rates of gregarious song and juvenile rats playing at high rates. Transcript levels for several glutamatergic receptor genes, such as GRIN1, GRIN2A, and GRIA1, were consistently upregulated in highly gregarious (i.e., playful/high singing) animals. This study is the first to directly investigate shared neuromodulators of positive, non-sexual social behaviors across songbirds and mammals. It provides insight into a conserved brain region that may regulate similar behaviors across vertebrates., (© 2024 The Author(s). Genes, Brain and Behavior published by International Behavioural and Neural Genetics Society and John Wiley & Sons Ltd.)

Glucocorticoid (GC) hormones have traditionally been interpreted as indicators of stress, but the extent to which they provide information on physiological state remains debated. GCs are metabolic hormones that amongst other functions ensure increasing fuel (i.e. glucose) supply on the face of fluctuating energetic demands, a role often overlooked by ecological studies investigating the consequences of GC variation. Furthermore, because energy budget is limited, in natural contexts where multiple stimuli coexist, the organisms' ability to respond physiologically may be constrained when multiple triggers of metabolic responses overlap in time. Using free-living spotless starling (Sturnus unicolor) chicks, we experimentally tested whether two stimuli of different nature known to trigger a metabolic or GC response, respectively, cause a comparable increase in plasma GCs and glucose. We further tested whether response patterns differed when both stimuli occurred consecutively. We found that both experimental treatments caused increases in GCs and glucose of similar magnitude, suggesting that both variables fluctuate along with variation in energy expenditure, independently of the trigger. Exposure to the two stimuli occurring subsequently did not cause a difference in GC or glucose responses compared with exposure to a single stimulus, suggesting a limited capacity to respond to an additional stimulus during an ongoing acute response. Lastly, we found a positive and significant correlation between plasma GCs and glucose after the experimental treatments. Our results add to the increasing research on the role of energy expenditure on GC variation, by providing experimental evidence on the association between plasma GCs and energy metabolism., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Many bird species commonly aggregate in flocks for reasons ranging from predator defense to navigation. Available evidence suggests that certain types of flocks-the V and echelon formations of large birds-may provide a benefit that reduces the aerodynamic cost of flight, whereas cluster flocks typical of smaller birds may increase flight costs. However, metabolic flight costs have not been directly measured in any of these group flight contexts [Zhang and Lauder, J. Exp. Biol. 226 , jeb245617 (2023)]. Here, we measured the energetic benefits of flight in small groups of two or three birds and the requirements for realizing those benefits, using metabolic energy expenditure and flight position measurements from European Starlings flying in a wind tunnel. The starlings continuously varied their relative position during flights but adopted a V formation motif on average, with a modal spanwise and streamwise spacing of [0.81, 0.91] wingspans. As measured via CO 2 production, flight costs for follower birds were significantly reduced compared to their individual solo flight benchmarks. However, followers with more positional variability with respect to leaders did less well, even increasing their costs above solo flight. Thus, we directly demonstrate energetic costs and benefits for group flight followers in an experimental context amenable to further investigation of the underlying aerodynamics, wake interactions, and bird characteristics that produce these metabolic effects., Competing Interests: Competing interests statement:The authors declare no competing interest.

Whether avian migrants can adapt to their changing world depends on the relative importance of genetic and environmental variation for the timing and direction of migration. In the classic series of field experiments on avian migration, A. C. Perdeck discovered that translocated juveniles failed to reach goal areas, whereas translocated adults performed 'true-goal navigation'. His translocations of > 14 000 common starlings ( Sturnus vulgaris ) suggested that genetic mechanisms guide juveniles into a population-specific direction, i.e. 'vector navigation'. However, alternative explanations involving social learning after release in juveniles could not be excluded. By adding historical data from translocation sites, data that was unavailable in Perdeck's days, and by integrated analyses including the original data, we could not explain juvenile migrations from possible social information upon release. Despite their highly social behaviour, our findings are consistent with the idea that juvenile starlings follow inherited information and independently reach their winter quarters. Similar to more solitarily migrating songbirds, starlings would require genetic change to adjust the migration route in response to global change.

Urbanization is rapidly changing the environment and creating new challenges in the lives of animals across the globe. Anthropogenic contaminants-like heavy metals-can persist within the environment for prolonged periods of time and present a widespread problem for those living near contaminated areas. Lead (Pb) was a commonly used heavy metal that continues to threaten the health of all organisms despite being phased out, especially in urban areas where historical use was more common. In this study, a common urban-adapter, the European starling (Sturnus vulgaris), was trapped to explore whether feather Pb burden is greater in birds from urban habitats than rural habitats, as well as whether Pb burdens were correlated with behavior, physiology, and feather development. Across four sites (two rural and two urban), soil Pb concentrations were measured and 197 free-living starlings were captured to measure feather Pb concentrations. Using linear mixed models, this study found that urban starling nestlings had elevated feather Pb burdens compared to rural nestlings. In contrast, there was no correlation between Pb and urbanization in adult birds whose exposure to Pb may reflect a larger spatial range compared to nestlings. For both nestlings and adults, feather Pb was uncorrelated to corticosterone, testosterone, aggressive behavior, or feather growth rates. These findings suggest that starlings may be a useful biomonitoring tool to detect Pb in the local environment, however, the age and spatial range of birds is a critical consideration in applying this tool. Further work is needed to understand the intricate relationship between heavy metals, behavior, morphological development, and physiology in free-living organisms., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Many animals produce signals that consist of vocalizations and movements to attract mates or deter rivals. We usually consider them as components of a single multimodal signal because they are temporally coordinated. Sometimes, however, this relationship takes on a more complex spatiotemporal character, resembling choreographed music. Timing is important for audio-visual integration, but choreographic concordance requires even more skill and competence from the signaller. Concordance should therefore have a strong impact on receivers; however, little is known about its role in audio-visual perception during natural interactions. We studied the effects of movement and song type concordance in audio-visual displays of the starling, Sturnus vulgaris. Starlings produce two types of movements that naturally appear in specific phrases of songs with a similar temporal structure and amplitude. In an experiment with a taxidermic robotic model, males responded more to concordant audio-visual displays, which are also naturally preferred, than to discordant displays. In contrast, the effect of concordance was independent of the specific combination of movement and song types in a display. Our results indicate that the concordance of movements and songs was critical to the efficacy of the display and suggest that the information that birds gained from concordance could not be obtained by adding information from movements and songs., (© 2024. The Author(s).)

Male songbirds produce female-directed songs in spring that convey a state of sexual motivation. Many songbirds also sing in fall flocks in affiliative/gregarious contexts in which song is linked to an intrinsic positive affective state. The periaqueductal gray (PAG) in mammals, which is organized into functional columns, integrates information from multiple brain regions and relays this information to vocal motor areas so that an animal emits a vocal signal reflective of its affective state. Here, we test the hypothesis that distinct columns in the songbird PAG play roles in the distinct affective states communicated by sexually motivated and gregarious song. We quantified the numbers of immediate early gene ZENK-positive cells in 16 PAG subregions in male European starlings (Sturnus vulgaris) after singing gregarious or sexually motivated song. Results suggest that distinct PAG columns in songbirds context-specifically regulate song, agonistic, and courtship behaviors. A second exploratory, functional tract-tracing study also demonstrated that inputs to the PAG from specific subregions of the medial preoptic nucleus may contribute to gregarious song and behaviors indicative of social dominance. Together, findings suggest that conserved PAG columns and inputs from the preoptic nucleus may play a role in context-specific vocal and other social behaviors., (© 2023 The Authors. Annals of the New York Academy of Sciences published by Wiley Periodicals LLC on behalf of The New York Academy of Sciences.)

Despite the potential for temporally dependent relationships between trait values and fitness (e.g. as juveniles approach life-stage transitions such as fledging), how developmental stage affects canalization (a measure of robustness to environmental variation) of morphological and physiological traits is rarely considered. To test the sensitivity of morphological and physiological traits to environmental variation in two developmental stages, we manipulated brood size at hatch in European starlings (Sturnus vulgaris) and cross-fostered chicks between enlarged and reduced broods approaching fledging. We measured body size (mass, tarsus, wing length) and physiological state (aerobic capacity, oxidative status) at asymptotic mass on day 15, then cross-fostered chicks between 'high' and 'low' quality environments and assessed the same traits again on day 20, after 5 days of pre-fledging mass recession. Chicks in reduced broods were heavier at asymptotic mass and had lower reactive oxygen metabolites than enlarged broods, whereas structural size, aerobic capacity and antioxidant capacity were unaffected by experimental brood size. The observed canalization of structural and physiological traits during early development was maintained after cross-fostering, during late development. However, in contrast to early development, antioxidant capacity approaching fledging appeared sensitive to environmental conditions, as trajectories varied by cross-fostering treatment. Elevated reactive oxygen metabolites observed after early development in enlarged brood chicks were maintained after cross-fostering, suggesting that canalized development in low-quality environments could produce oxidative costs that carry over between life stages, even when conditions improve. These data reveal trait-specific relationships between environmental conditions and development, and highlight how natal environment effects may vary by developmental stage., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Seasonality in songbirds includes not only reproduction but also seasonal changes in singing behavior and its neural substrate, the song control system (SCS). Prior research mainly focused on the role of sex steroids on this seasonal SCS neuroplasticity in males. In this review, we summarize the advances made in the field of seasonal neuroplasticity by applying in vivo magnetic resonance imaging (MRI) in male and female starlings, analyzing the entire brain, monitoring birds longitudinally and determining the neuronal correlates of seasonal variations in plasma hormone levels and song behavior. The first MRI studies in songbirds used manganese enhanced MRI to visualize the SCS in a living bird and validated previously described brain volume changes related to different seasons and testosterone. MRI studies with testosterone implantation established how the consequential boost in singing was correlated to structural changes in the SCS, indicating activity-induced neuroplasticity as song proficiency increased. Next, diffusion tensor MRI explored seasonal neuroplasticity in the entire brain, focusing on networks beyond the SCS, revealing that other sensory systems and even the cerebellum, which is important for the integration of sensory perception and song behavior, experience neuroplasticity starting in the photosensitive period. Functional MRI showed that olfactory, and auditory processing was modulated by the seasons. The convergence of seasonal variations in so many sensory and sensorimotor systems resembles multisensory neuroplasticity during the critical period early in life. This sheds new light on seasonal songbirds as a model for unlocking the brain by recreating seasonally the permissive circumstances for heightened neuroplasticity., (© 2022 The Authors. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology published by Wiley Periodicals LLC.)

Migratory birds undergo seasonal changes to muscle biochemistry. Nonetheless, it is unclear to what extent these changes are attributable to the exercise of flight itself versus endogenous changes. Using starlings (Sturnus vulgaris) flying in a wind tunnel, we tested the effects of exercise training, a single bout of flight and dietary lipid composition on pectoralis muscle oxidative enzymes and lipid transporters. Starlings were either unexercised or trained over 2 weeks to fly in a wind tunnel and sampled either immediately following a long flight at the end of this training or after 2 days recovery from this flight. Additionally, they were divided into dietary groups that differed in dietary fatty acid composition (high polyunsaturates versus high monounsaturates) and amount of dietary antioxidant. Trained starlings had elevated (19%) carnitine palmitoyl transferase and elevated (11%) hydroxyacyl-CoA dehydrogenase in pectoralis muscle compared with unexercised controls, but training alone had little effect on lipid transporters. Immediately following a long wind-tunnel flight, starling pectoralis had upregulated lipid transporter mRNA (heart-type fatty acid binding protein, H-FABP, 4.7-fold; fatty acid translocase, 1.9-fold; plasma membrane fatty acid binding protein, 1.6-fold), and upregulated H-FABP protein (68%). Dietary fatty acid composition and the amount of dietary antioxidants had no effect on muscle catabolic enzymes or lipid transporter expression. Our results demonstrate that birds undergo rapid upregulation of catabolic capacity that largely becomes available during flight itself, with minor effects due to training. These effects likely combine with endogenous seasonal changes to create the migratory phenotype observed in the wild., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

Cooperatively breeding vertebrates are common in unpredictable environments where the costs and benefits of providing offspring care fluctuate temporally. To balance these fitness outcomes, individuals of cooperatively breeding species often exhibit behavioural plasticity according to environmental conditions. Although individual variation in cooperative behaviours is well-studied, less is known about variation in plasticity of social behaviour. Here, we examine the fitness benefits, plasticity and repeatability of nest guarding behaviour in cooperatively breeding superb starlings ( Lamprotornis superbus ). After demonstrating that the cumulative nest guarding performed at a nest by all breeders and helpers combined is a significant predictor of reproductive success, we model breeder and helper behavioural reaction norms to test the hypothesis that individuals invest more in guarding in favourable seasons with high rainfall. Variation in nest guarding behaviour across seasons differed for individuals of different reproductive status: breeders showed plastic nest guarding behaviour in response to rainfall, whereas helpers did not. Similarly, we found that individual breeders show repeatability and consistency in their nest guarding behaviour while individual helpers did not. Thus, individuals with the potential to gain direct fitness benefits exhibit greater plasticity and individual-level repeatability in cooperative behaviour.

The study examined the changes in intraperitoneal body temperature of laboratory mice, Jungar hamsters, European greenfinch Chloris chloris, and starlings. In a few minutes range, these changes significantly correlated not only between the animals of the same species, but also between the different classes such as birds and mammals, which were isolated from each other and maintained under different illumination regimen. This phenomenon indicates some external influence(s) on the central mechanisms of the thermal control system not related to illumination regiment. In 80% cases, the phases of most pronounced rhythms of body temperature oscillating with the periods of 8-9 and 12-13 min coincided with those of geomagnetic field within the accuracy of ±1 min. However, the amplitude of body temperature oscillations did not depend on the amplitude of geomagnetic field (GMF) oscillations. Synchronicity of the changes in body temperature and GMF was observed at the amplitude of GMF oscillation of 0.4 nT, which is extremely low value. In contrast, there was no reaction of body temperature to greater (6-10 nT) but irregular and abrupt perturbations of GMF., (© 2021. Springer Science+Business Media, LLC, part of Springer Nature.)

Traditionally, research unraveling seasonal neuroplasticity in songbirds has focused on the male song control system and testosterone. We longitudinally monitored the song behavior and neuroplasticity in male and female starlings during multiple photoperiods using Diffusion Tensor and Fixel-Based techniques. These exploratory data-driven whole-brain methods resulted in a population-based tractogram confirming microstructural sexual dimorphisms in the song control system. Furthermore, male brains showed hemispheric asymmetries in the pallium, whereas females had higher interhemispheric connectivity, which could not be attributed to brain size differences. Only females with large brains sing but differ from males in their song behavior by showing involvement of the hippocampus. Both sexes experienced multisensory neuroplasticity in the song control, auditory and visual system, and cerebellum, mainly during the photosensitive period. This period with low gonadal hormone levels might represent a 'sensitive window' during which different sensory and motor systems in the cerebrum and cerebellum can be seasonally re-shaped in both sexes., Competing Interests: JO, EC, JH, EJ, VD, MV, AV No competing interests declared, (© 2021, Orije et al.)

The capacity for sensory systems to encode relevant information that is invariant to many stimulus changes is central to normal, real-world, cognitive function. This invariance is thought to be reflected in the complex spatiotemporal activity patterns of neural populations, but our understanding of population-level representational invariance remains coarse. Applied topology is a promising tool to discover invariant structure in large datasets. Here, we use topological techniques to characterize and compare the spatiotemporal pattern of coactive spiking within populations of simultaneously recorded neurons in the secondary auditory region caudal medial neostriatum of European starlings ( Sturnus vulgaris ). We show that the pattern of population spike train coactivity carries stimulus-specific structure that is not reducible to that of individual neurons. We then introduce a topology-based similarity measure for population coactivity that is sensitive to invariant stimulus structure and show that this measure captures invariant neural representations tied to the learned relationships between natural vocalizations. This demonstrates one mechanism whereby emergent stimulus properties can be encoded in population activity, and shows the potential of applied topology for understanding invariant representations in neural populations. SIGNIFICANCE STATEMENT Information in neural populations is carried by the temporal patterns of spikes. We applied novel mathematical tools from the field of algebraic topology to quantify the structure of these temporal patterns. We found that, in a secondary auditory region of a songbird, these patterns reflected invariant information about a learned stimulus relationship. These results demonstrate that topology provides a novel approach for characterizing neural responses that is sensitive to invariant relationships that are critical for the perception of natural stimuli., (Copyright © 2021 the authors.)

Training and diet are hypothesized to directly stimulate key molecular pathways that mediate animal performance, and flight training, dietary fats, and dietary antioxidants are likely important in modulating molecular metabolism in migratory birds. This study experimentally investigated how long-distance flight training, as well as diet composition, affected the expression of key metabolic genes in the pectoralis muscle and the liver of European starlings ( Sturnus vulgaris , n = 95). Starlings were fed diets composed of either a high or low polyunsaturated fatty acid (PUFA; 18:2n-6) and supplemented with or without a water-soluble antioxidant, and one-half of these birds were flight trained in a wind-tunnel while the rest were untrained. We measured the expression of 7 (liver) or 10 (pectoralis) key metabolic genes in flight-trained and untrained birds. Fifty percent of genes involved in mitochondrial metabolism and fat utilization were upregulated by flight training in the pectoralis ( P < 0.05), whereas flight training increased the expression of only one gene responsible for fatty acid hydrolysis [lipoprotein lipase (LPL)] in the liver ( P = 0.04). Dietary PUFA influenced the gene expression of LPL and fat transporter fatty acid translocase (CD36) in the pectoralis and one metabolic transcription factor [peroxisome proliferator-activated receptor (PPAR)-α (PPARα)] in the liver, whereas dietary antioxidants had no effect on the metabolic genes measured in this study. Flight training initiated a simpler causal network between PPARγ coactivators, PPARs, and metabolic genes involved in mitochondrial metabolism and fat storage in the pectoralis. Molecular metabolism is modulated by flight training and dietary fat quality in a migratory songbird, indicating that these environmental factors will affect the migratory performance of birds in the wild.

Group cohesion relies on the ability of its members to process social signals. Songbirds provide a unique model to investigate links between group functioning and brain processing of social acoustic signals. In the present study, we performed both behavioral observations of social relationships within a group of starlings and individual electrophysiological recordings of HVC neuronal activity during the broadcast of either familiar or unfamiliar individual songs. This allowed us to evaluate and compare preferred partnerships and individual electrophysiological profiles. The electrophysiological results revealed asymmetric neuronal activity in the HVC and higher responsiveness to familiar than to unfamiliar songs. However, most importantly, we found a correlation between strength of cerebral asymmetry and social integration in the group: the more preferred partners a bird had, the more its HVC neuronal activity was lateralized. Laterality is likely to give advantages in terms of survival. Our results suggest that these include social skill advantages. Better knowledge of links between social integration and lateralization of social signal processing should help understand why and how lateralization has evolved.

The integration of normative and descriptive analyses of decision processes in humans struggles with the fact that measuring preferences by different procedures yields different rankings and that humans appear irrationally impulsive (namely, show maladaptive preference for immediacy). Failure of procedure invariance has led to the widespread hypothesis that preferences are constructed "on the spot" by cognitive evaluations performed at choice time, implying that choices should take extra time in order to perform the necessary comparisons. We examine this issue in experiments with starlings (Sturnus vulgaris) and show that integrating normative and descriptive arguments is possible and may help reinterpreting human decision results. Our main findings are that (1) ranking alternatives through direct rating (response time) accurately predicts preference in choice, overcoming failures of procedure invariance; (2) preference is not constructed at choice time nor does it involve extra time (we show that the opposite is true); and (3) starlings' choices are not irrationally impulsive but are instead directly interpretable in terms of profitability ranking. Like all nonhuman research, our protocols examine decisions by experience rather than by description, and hence support the conjecture that irrationalities that prevail in research with humans may not be observed in decisions by experience protocols., Competing Interests: The authors have declared that no competing interests exist.

Introduced bird species can become invasive in agroecosystems and their management is inhibited if their origin and movements are not well understood. Stable isotope measurements of feathers can be used to infer molt origins and interstate movements in North America. We analyzed stable-hydrogen (δ2H), carbon (δ13C) and nitrogen (δ 15N) isotope ratios in feathers to better understand the molt origin of European starlings (Sturnus vulgaris) collected at dairies and feedlots throughout the United States. Primary feathers were used from 596 adult and 90 juvenile starlings collected during winter at dairies and feedlots that experience starling damages in Arizona, California, Colorado, Idaho, Illinois, Iowa, Kansas, Minnesota, Missouri, Nebraska, Nevada, New Hampshire, New Mexico, New York, North Carolina, Oregon, Texas, Vermont, Washington and Wisconsin. The best-fit model indicated that the combination of feather δ2H, δ13C and δ15N values best predicted the state where samples were collected and thus supported use of this approach for tracing molt origins in European starlings. Interestingly, molt origins of starlings collected at dairies and feedlots generally west of -90° longitude (i.e. 11 of 15 states west of the Mississippi River, including Wisconsin) were assigned to the collection state and/or the state adjacent to the collection state. In contrast, molt origins of starlings collected generally east of -90° longitude (four of five eastern states) were not assigned to the collection state and/or the state adjacent to the collection state. Among all starlings (N = 686), 23% were assigned to the collection state and 19% were assigned to the state adjacent to the collection state. Among all males (N = 489) and all females (N = 197), 23% and 26% were assigned to the collection state and 19% and 13% were assigned to the state adjacent to the collection state, respectively. We observed a greater proportion (88%) of juvenile starlings assigned to states other than their collection state (i.e. potentially a result of natal dispersal) than that proportion (76%) in adult starlings. This study included an unprecedented sample of feather isotopes from European starlings throughout the United States. As a novel contribution to the ecology and management of invasive and migratory passerines, we demonstrate how such feather isoscapes can be used to predict molt origin and, potentially, interstate movements of European starlings for subsequent ecological and management investigations., Competing Interests: The authors have declared that no competing interests exist.

Many songbirds sing in non-reproductive contexts while in flocks. Singing in such gregarious contexts is critical for maintaining and learning songs; however, song is not directed towards other individuals and has no obvious, immediate social consequences. Studies using conditioned place preference (CPP) tests of reward indicate that song production in gregarious contexts correlates positively with a bird's intrinsic reward state and with opioid markers in the medial preoptic nucleus (mPOA). However, the causal involvement of opioids in gregarious song is unknown. Here we report that the selective mu opioid receptor (MOR) agonist fentanyl dose-dependently facilitates gregarious song and reduces stress/anxiety-related behavior in male and female European starlings. Furthermore, infusion of siRNA targeting MORs specifically in mPOA both suppresses gregarious song and disrupts the positive association between affective state and singing behavior, as revealed using CPP tests of song-associated reward. Results strongly implicate opioids in gregarious song and suggest that endogenous opioids in the mPOA may facilitate song by influencing an individual's intrinsic reward state.

Group living is predicted to arise only when the fitness benefits outweigh the costs of sociality. Group-living species-including cooperatively breeding and family-living birds and mammals-occur most frequently in environments where climatic conditions fluctuate unpredictably from year to year. The fitness consequences of group living are thus expected to vary with changing environmental conditions, though few studies have examined this possibility. We examined whether living in large social groups improves adult survivorship in cooperatively breeding superb starlings ( Lamprotornis superbus ). We also tested the hypothesis that larger groups buffer against harsh conditions by increasing survivorship most under periods of low rainfall. We found that group size was positively correlated with adult survival but in a sex-specific manner: female survival increased with group size across all environmental conditions, whereas male survival increased with group size only in wet years. Together with previous work in this system, our results suggest that larger groups confer survival benefits by reducing predation, rather than by improving access to food or buffering against physiological stress. Although group living does not appear to buffer against harsh conditions in adult starlings living in a fluctuating environment, living in larger groups does confer a survival advantage.

Evolutionary theory expects social, communicative species to eavesdrop most on other species' alarm calls [e.g., 1, 2] but also that solitary-living species benefit most from eavesdropping [3, 4]. Examples of solitary species responding to the alarm calls of other species, however, are limited and unconvincing [3-5]. The Swahili name for the red-billed oxpecker (Buphagus erythrorynchus) is Askari wa kifaru, the rhinos' guard [6]. Black rhino (Diceros bicornis) are a solitary-living, non-vocal species and are critically endangered through hunting. We searched Hluhluwe-iMfolozi Park, South Africa, for rhinoceros for 27 months with and without the aid of radio telemetry and conducted 86 experimental, unconcealed approaches to 11 rhino, without or with varying numbers of resident oxpecker. Oxpeckers enabled rhinos to evade detection by us in 40% to 50% of encounters. Alarm-calling by oxpeckers significantly improved the rate and distance that rhinos detected our approach from 23% to 100% and 27 ± 6 m to 61 ± 4 m, respectively. Every additional oxpecker improved detection distance by 9 m. Rhinos alerted by oxpeckers' alarm calls never re-oriented in our direction but moved to face downwind. Thus, oxpeckers' calls communicate only threat proximity, not direction, and rhinos assume the hunter is stalking from downwind. We confirm that oxpeckers guard rhinos and the importance of depredation, not sociality, in the evolution of eavesdropping [4, 7]. Conservationists should consider reintroducing oxpeckers to rhino populations, reinstating their anti-human sentinel [8]. VIDEO ABSTRACT., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Sleep is considered to be of crucial importance for performance and health, yet much of what we know about sleep is based on studies in a few mammalian model species under strictly controlled laboratory conditions. Data on sleep in different species under more natural conditions may yield new insights in the regulation and functions of sleep. We therefore performed a study with miniature electroencephalogram (EEG) data loggers in starlings under semi-natural conditions, group housed in a large outdoor enclosure with natural temperature and light. The birds showed a striking 5-h difference in the daily amount of non-rapid-eye-movement (NREM) sleep between winter and summer. This variation in the amount of NREM sleep was best explained by night length. Most sleep occurred during the night, but when summer nights became short, the animals displayed mid-day naps. The decay of NREM sleep spectral power in the slow-wave range (1.1-4.3 Hz) was steeper in the short nights than in the longer nights, which suggests that birds in summer have higher sleep pressure. Additionally, sleep was affected by moon phase, with 2 h of NREM sleep less during full moon. The starlings displayed very little rapid-eye-movement (REM) sleep, adding up to 1.3% of total sleep time. In conclusion, this study demonstrates a pronounced phenotypical flexibility in sleep in starlings under semi-natural conditions and shows that environmental factors have a major impact on the organization of sleep and wakefulness., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

The dynamics of body temperature was used to determine the food intake synchronization in starlings located at a distance of 105 km from each other. Thermal accumulators were implanted intraperitoneally to the birds; the birds had free access to food or were kept under the conditions of food deprivation. It was found that food intake moments significantly more often coincided with an accuracy of up to a minute even in the birds separated by a considerable distance. The difference in time of sunrise of a few minutes, as well as the conditions of constant lighting did not affect the synchronism of food intake. On the basis of the data obtained, it can be assumed that there is a certain external factor, probably of an electromagnetic nature, which synchronizes the functional activity of the body, including food intake.

In previous research, pigeons and hill mynas that performed differently on an object permanence task were presumed to attend to objects in different ways (Plowright, Reid, & Kilian, 1998). In the current study, we conducted 4 experiments to investigate if the attention of hill mynas and pigeons is object-based and if there are species differences in their visual-attentional processes. In Experiment 1, pigeons were tested in a spatial cueing task requiring them to respond sequentially to a cue and a target that appeared at 1 of the 4 ends of two rectangles. Both when the response to a fixation stimulus was required before target presentation (Experiment 1A) and when such a response was not required (Experiment 1B), there were no significant differences in reaction times to the targets appearing at cued and noncued rectangles; these results provided no evidence of object-based attention in pigeons. In Experiment 2, for 2 of the 3 hill mynas tested in a procedure similar to that for the pigeons in Experiment 1B, reaction times were shorter to the target appearing on the cued rectangle than to the target appearing on noncued rectangle, suggesting the operation of object-based attention, as in humans. In Experiment 3, we tested naive pigeons by means of the procedure used for hill mynas in Experiment 2. However, again pigeons showed no evidence of object-based attention, suggesting a species difference in attentional processes. The generality of the current results and evolution of the possible species differences were discussed. (PsycINFO Database Record (c) 2020 APA, all rights reserved).

Animals often avoid novel objects, a behavior known as neophobia. We examined behavioral responses of captive European starlings to novel objects placed at their food dishes. Exposure occurred concurrently to food reintroduction following overnight fasting. Behavior was analyzed for 10 min via video recording. We expected an increase in avoidance behavior compared with trials in which food was reintroduced without a novel object. Seven of 10 novel objects increased latency to approach the dish. In contrast to our expectations, neither prior restraint nor exogenous corticosterone changed the neophobic response to novel objects. While exposure to a novel object increased approach latency, there was no additional effect of restraining animals in a cloth bag for 15 min before food reintroduction. Furthermore, the subcutaneous injection of corticosterone did not affect approach latency by itself, nor did it affect the response to a novel object. Finally, we expected repeated exposure to the same object to extinguishing the neophobic response, and that restraint stress would prevent habituation. Our results show that European starlings habituate rapidly to exposure to a novel object, as approach latency returned to baseline within three repeated exposures to the same object. When the repeated presentation of the object was combined with prior restraint, however, the latency to approach never returned to baseline. These results reveal that neophobia in starlings is object-specific and that, while neither acute stress nor corticosterone directly affects the behavioral response to a novel object, acute stress appears to have a permissive effect on neophobia by inhibiting habituation., (© 2019 Wiley Periodicals, Inc.)

Adult neuroplasticity in the song control system of seasonal songbirds is largely driven by photoperiod-induced increases in testosterone. Prior studies of the relationships between testosterone, song performance and neuroplasticity used invasive techniques, which prevent analyzing the dynamic changes over time and often focus on pre-defined regions-of-interest instead of examining the entire brain. Here, we combined (i) in vivo diffusion tensor imaging (DTI) to assess structural neuroplasticity with (ii) repeated monitoring of song and (iii) measures of plasma testosterone concentrations in thirteen female photosensitive starlings (Sturnus vulgaris) who received a testosterone implant for 3 weeks. We observed fast (days) and slower (weeks) effects of testosterone on song behavior and structural neuroplasticity and determined how these effects correlate on a within-subject level, which suggested separate contributions of the song motor and anterior forebrain pathways in the development of song performance. Specifically, the increase in testosterone correlated with a rapid increase of song rate and RA volume, and with changes in Area X microstructure. After implant removal, these variables rapidly reverted to baseline levels. In contrast, the more gradual improvement of song quality was positively correlated with the fractional anisotropy values (DTI metric sensitive to white matter changes) of the HVC-RA tract and of the lamina mesopallialis, which contains fibers connecting the song control nuclei. Thus, we confirmed many of the previously reported testosterone-induced effects, like the increase in song control nuclei volume, but identified for the first time a more global picture of the spatio-temporal changes in brain plasticity., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Vocal communication plays an important role in the regulation of social interactions and the coordination of activities in many animal species. Synchrony is an essential part of the establishment and maintenance of pair bonds, but few reports have investigated decision-making at the pair level. We investigated temporal characteristics of call exchanges in pale-winged starlings (Onychognathus nabouroup) that could predict whether one, two, or neither members of a pair would take off. Our analysis of these interactions revealed that the overall rhythm of a call exchange, as well as the acceleration towards the end of an interaction, were significantly associated with the type of behavioural outcome. Faster rhythms were associated with higher probabilities that both birds would fly away. Our results confirm the findings of previous studies showing that higher rates of alarm calls indicate imminent departure and highlight the relationship between temporal features of vocal interactions and their outcome.

Estimating the potential distribution of invasive species has been primarily achieved by employing species distribution models (SDM). Recently introduced joint species distribution models (JSDM) that include species interactions are expected to improve model output. Here we compare the predictive ability of SDM and JSDM by modelling the distribution of one of the most prolific avian invaders in the world, the common myna (Acridotheres tristis), in a recent introduction in Israel. Our results indicate that including information on the local species composition did not improve model accuracy, possibly because of the unique characteristics of this species that include broad environmental tolerance and behavior flexibility. However, the JSDM provided insights into co-occurrence patterns of common mynas and their local heterospecifics, suggesting that at this time point, there is no evidence of species exclusion by common mynas. Our findings suggest that the invasion potential of common mynas depends greatly on urbanization and less so on the local species composition and reflect the major role of anthropogenic impact in increasing the distribution of avian invaders.

Wild-caught European starlings (Sturnus vulgaris) were exposed to a learning task to determine whether heart rate (HR) and behavior responses to the learning activated the sympathetic nervous system. Birds learned to discriminate between images of opposite convexity (concave and convex) based on shading cues in a closed economy (food only available through task completion). Once learned, the task was changed in three ways: (a) manipulating the angle and shape of the image; (b) altering the availability of the task; and (c) reversing the positive stimulus. HR, used as an index of catecholamine release, was measured during each change to determine whether having to alter previously established behaviors to learn new behaviors elicited a sympathetic response. Starlings decreased their HR during the initial discrimination training and did not alter their HR when presented with modified images or when the positive stimulus was reversed. However, HR increased when the task became unavailable and decreased upon its return, suggesting that preventing task performance was perceived as stressful. Birds also modified their behavior when tasks were changed. The number of trials per minute decreased during the reversal treatment, as did the success rate, suggesting that starlings may try to conserve energy when access to food diminishes. This is also supported by the decrease in perch hops per minute when the task was unavailable and the subsequent increase upon its return. Overall, these results suggest that learning per se does not activate the sympathetic nervous system and, therefore, is not a stressor for wild birds., (© 2019 Wiley Periodicals, Inc.)

Telomere length can be considered as an indicator of an organism's somatic state, long telomeres reflecting higher energy investment in self-maintenance. Early-life is a period of intense investment in somatic growth and in physiological maturation but how this is reflected in telomere length remains unclear. Using European starling chicks we tested: (i) how telomere length measured at asymptotic mass is related to proxies of somatic growth and physiological maturity in 17-day-old nestlings; (ii) how telomere length measured at 17 days then predicts the changes in somatic and physiological maturity occurring in fledglings (between 17 and 21 days); (iii) how growth and telomere length co-vary when chicks are under experimentally good (fed) growth conditions. Depending on environmental conditions, our data suggest links between somatic growth, physiological maturation and body maintenance parameters (positive with oxidative stress and negative with telomere length) in nestlings. Telomere length measured at day 17 predicted a subsequent change in physiological maturation variables observed in fledglings, but only in second-brood chicks: chicks with shorter telomeres had a higher pre-fledging rate of increase in haematocrit and haemoglobin content and a greater decrease in reticulocyte count. Finally, food supplementation of chicks did not change telomere length compared with that in control siblings. Our results suggest that physiological maturation prior to fledging may occur at the expense of telomere length but only when environmental conditions are sub-optimal., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

With the approval of exenatide in 2005, physicians had a new class of hypoglycemic agents available for the treatment of type 2 diabetes-the glucagon-like peptide-1 receptor agonists (or GLP-1 receptor agonists). As of this writing, there are seven drugs in this class available in the United States. In addition to demonstrating either cardiovascular risk neutrality or overt benefit, as now mandated by the United States Food and Drug Administration (FDA), many of these drugs have other, unexpected actions. It is our goal to outline these actions, some beneficial, some not. We have reviewed English-language articles in this area, not for an exhaustive study, but rather a broad search to define current understanding and perhaps generate further investigation.

Pigments are largely responsible for the appearance of organisms. Most biological pigments derive from the metabolism of shikimic acid (melanins), mevalonic acid (carotenoids) or levulinic acid (porphyrins), which thus generate the observed diversity of external phenotypes. Starlings are generally dark birds despite iridescence in feathers, but 10% of species have evolved plumage pigmentation comprising bright colors that are known to be produced only by carotenoids. However, using micro-Raman spectroscopy, we have discovered that the bright yellow plumage coloration of one of these species, the Afrotropical golden-breasted starling Cosmopsarus regius , is not produced by carotenoids, but by vitamin A (all- trans -retinol). This is the first organism reported to deposit significant amounts of vitamin A in its integument and use it as a body pigment. Phylogenetic reconstructions reveal that the retinol-based pigmentation of the golden-breasted starling has independently appeared in the starling family from dark ancestors. Our study thus unveils a unique evolution of a new class of external pigments consisting of retinoids., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

A three-species population system under a facultative mutualistic relationship of one of the species is studied. The considered interactions are as follows: facultative between the first species and the second species, obligatory mutualism between the second species and the first one, and the third species is a predator of the first species. For this purpose, we extend the model proposed by Morozov et al., originally used to describe obligatory mutualism, to consider obligatory and facultative mutualism and prove that under adequately selected parameters this system produces a spatial patchy spread of populations or continuous wave fronts. Since the analytical treatment of a three-species model is often prohibitive, we first analyze the interaction between two mutualist species without diffusion and without the presence of the predator. Some parameters are fixed in the bistable regime of the mutualistic species to further consider the influence of the third species. The remaining parameters are then selected to produce patchy patterns under different mortality rates. Finally, the equations of the final three-species system are numerically solved to test the influence of different initial conditions in the formation of patchy populations. It is confirmed that the velocity and the profile of a traveling front are independent on the initial conditions. Our approach opens the way to study more general biological situations., (Copyright © 2019 Elsevier B.V. All rights reserved.)

In seasonally breeding animals, changes in photoperiod and sex-steroid hormones may modify sexual behavior in part by altering the activity of neuromodulators, including opioids and dopamine. In rats and birds, activation of mu-opioid receptors (MOR) and dopamine D1 receptors in the medial preoptic area (mPOA) often have opposing effects on sexual behavior, yet mechanisms by which the mPOA integrates these opposing effects to modulate behavior remain unknown. Here, we used male European starlings (Sturnus vulgaris) to provide insight into the hypothesis that MOR and D1 receptors modify sexual behavior seasonally by altering activity in the same neurons in the mPOA. To do this, using fluorescent immunohistochemistry, we examined the extent to which MOR and D1 receptors co-localize in mPOA neurons and the degree to which photoperiod and the sex-steroid hormone testosterone alter co-localization. We found that MOR and D1 receptors co-localize throughout the mPOA and the bed nucleus of the stria terminalis, a region also implicated in the control of sexual behavior. Numbers of single and co-labeled MOR and D1 receptor labeled cells were higher in the rostral mPOA in photosensitive males (a condition observed just prior to the breeding season) compared to photosensitive males treated with testosterone (breeding season condition). In the caudal mPOA co-localization of MOR and D1 receptors was highest in photosensitive males compared to photorefractory males (a post-breeding season condition). Seasonal shifts in the degree to which neurons in the mPOA integrate signaling from opioids and dopamine may underlie seasonal changes in the production of sexual behavior., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Stress during early development can induce substantial long-term effects in organisms. In the case of birds, despite growth compensations, nestlings reared under harsh conditions typically show reduced survival chances in adulthood. It has been proposed that environmental early-life stressors could affect longevity via effects on telomere length, possibly mediated through oxidative stress. However, the link between these processes is not clear. In this study, we experimentally manipulated brood size in spotless starlings (Sturnus unicolor) to test the causal relationship between early stress, oxidative and corticosterone-mediated stress and telomere shortening. Our results show that experimentally enlarged brood sizes led to a reduction in morphometric development on nestlings, the effect being stronger for females than males. Additionally, basal corticosterone levels increased with increasing brood size in female nestlings. Neither plasma antioxidant status nor malondialdehyde levels (a marker of lipid peroxidation) were affected by experimental brood size, although the levels of a key intracellular antioxidant (glutathione) decreased with increasing brood size. We found that the treatment showed a quadratic effect on nestling telomere lengths: these were shortened either by increases or by decreases in the original brood size. Our study provides experimental evidence for a link between developmental stress and telomere length, but does not support a direct causal link of this reduction with corticosterone or oxidative stress. We suggest that future studies should focus on how telomere length responds to additional markers of allostatic load., (© 2018 European Society For Evolutionary Biology. Journal of Evolutionary Biology © 2018 European Society For Evolutionary Biology.)

Judgement bias tasks are designed to provide markers of affective states. A recent study of European starlings (Sturnus vulgaris) demonstrated modest familial effects on judgement bias performance, and found that adverse early experience and developmental telomere attrition (an integrative marker of biological age) both affected judgement bias. Other research has shown that corticosterone levels affect judgement bias. Here, we investigated judgement bias using a modified Go/No Go task in a new cohort of starlings (n = 31) hand-reared under different early-life conditions. We also measured baseline corticosterone and the corticosterone response to acute stress in the same individuals. We found evidence for familial effects on judgement bias, of a similar magnitude to the previous study. We found no evidence that developmental treatments or developmental telomere attrition were related to judgement bias per se. We did, however, find that birds that experienced the most benign developmental conditions, and birds with the greatest developmental telomere attrition, were significantly faster to probe the learned unrewarded stimulus. We also found that the birds whose corticosterone levels were faster to return towards baseline after an acute stressor were slower to probe the learned unrewarded stimulus. Our results illustrate the potential complexities of relationships between early-life experience, stress and affectively mediated decision making. For judgement bias tasks, they demonstrate the importance of clearly distinguishing factors that affect patterns of responding to the learned stimuli (i.e. response inhibition in the case of the Go/No Go design) from factors that influence judgements under ambiguity.

Stimuli such as visual representations of raptors, snakes, or humans are generally assumed to be universally fear-inducing in birds and considered as a product of evolutionary perceptual bias. Both naïve and experienced birds should thus react to such stimuli with fear reactions. However, studies on different species have shown the importance of experience in the development of these fear reactions. We hypothesized that the responses of adult European starlings to fear-inducing visual stimuli may differ according to experience. We compared the reactions of Hand-raised adults with no experience of predators to those of Wild-caught adults, with potentially extensive experience with predators. Three visual stimuli (i.e. human, raptor, snake) were broadcast to 17 birds as 2D pictures (displayed via a LCD screen) with different modalities of presentation: degree of proximity and with or without movement. The results reveal that the birds were particularly sensitive to proximity and movement, with more attention towards moving stimuli and more withdrawal for close stimuli. The human stimulus elicited attention in both the distant and moving modalities but, like the other stimuli, mostly withdrawal when it was close. Developmental experience appeared to influence the emotional level, as the Hand-raised birds reacted strongly to all stimuli and all modalities, contrarily to the WC birds which performed withdrawals almost only for close stimuli and attention to moving stimuli. Stimuli proximity and movement seemed, therefore, relevant features that elicited negative reactions in Wild-caught birds. The Hand-raised birds were equally attentive to both distant and moving stimuli. Thus the young birds showed no real discrimination. Early and later experiences may, therefore, influence birds' reactions. Starlings may require experience with real threats to develop adaptive responses, i.e. limiting unnecessary loss of energy by fleeing in front of non-dangerous stimuli.

The cost of reproduction results in a life-history trade-off where investment in current reproduction via costly parental care decreases subsequent fitness. Although this trade-off is thought to occur ubiquitously across animals, there is equivocal evidence that parental care behaviours are costly. A major challenge of studying the cost of parental care has been a lack of consensus over which physiological mechanisms underlie this trade-off. Here, we compare four traits believed to mediate the cost of parental care by examining whether glucocorticoids, oxidative stress, immune function or body condition represent a cost of performing offspring care and shape subsequent fitness. We use a 4 year dataset collected in free-living cooperatively breeding superb starlings ( Lamprotornis superbus ), a species in which parental and alloparental care effort varies widely among individuals and across years. Our results showed that within-individual change in physiology was unrelated to investment in offspring care, and physiological state during chick rearing did not predict the likelihood that an individual would breed in subsequent seasons. Instead, individuals that had elevated baseline corticosterone during incubation performed more nest guarding, suggesting that this hormone may play a preparatory role for investing in offspring care. Together, our results indicate that superb starlings modify their investment in offspring care according to their physiological state during incubation, despite there being no evidence of a short-term physiological cost of parental or alloparental care. Thus, breeding cooperatively appears to provide individuals with the flexibility to adjust their investment in offspring care and overcome any potential costs of reproduction., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The cyclic ratio (CR) schedule is a behavioural assay developed to study feeding in rats, in which the number of operant responses required to obtain food reward (the ratio requirement) increases and then decreases in a repeating cycle. In a recent study, we used the CR schedule with European starlings (Sturnus vulgaris) to investigate the effects of an early-life manipulation on adult feeding behaviour. As this was the first time the CR schedule had been used with any bird species, a more in-depth evaluation is warranted. Here, we performed a fuller CR experiment with the same birds as the prior study, a year later. First, we examine the individual consistency of feeding behaviour between experimental sessions and also between CR schedules comprising different ratio requirement progressions. We found that between-session consistency was poor to moderate, and that a geometric ratio progression provided greater between-session consistency than an arithmetic ratio progression. Second, we tried to replicate some of the canonical findings from rats working on CR schedules. In contrast to findings from rats, we found that defence of feeding rates did not increase when starlings were acutely food deprived. However, as in rats, we found that the post-reinforcement pause increased linearly with the upcoming ratio requirement, suggesting that starlings were able to learn the cyclic nature of the schedule. Third, we compared the results from the present study concerning the impacts of our early-life treatment with those from our earlier study. We found that the majority of our previous findings were replicated in the same individuals one year on, reinforcing our previous conclusion that the early-life manipulation had canalised our birds into two groups with different patterns of feeding rate defence., Competing Interests: The authors have declared that no competing interests exist.

Reconsolidation theory describes memory formation as an ongoing process that cycles between labile and stable states. Though sleep is critical for the initial consolidation of a memory, there has been little evidence that sleep facilitates reconsolidation. We now demonstrate in two experiments that a sleep-consolidated memory can be destabilized if the memory is reactivated by retrieval. The destabilized memory, which can be impaired if an interference task is encountered after, but not before, the memory is reactivated, is then reconsolidated after sleep. In two additional experiments, we provide evidence suggesting that the learning of the interference task promotes the subsequent sleep-dependent enhancement of the original memory. These results provide novel insight into the complex mechanisms of memory processing, as well as critical evidence supporting the view that long-term memory formation involves a dynamic process of sleep-dependent consolidation, use-dependent destabilization, and sleep-dependent reconsolidation.

Environmental cues, such as photoperiod, regulate the timing of major life-history events like breeding through direct neuroendocrine control. Less known is how supplementary environmental cues (e.g., nest sites, food availability) interact to influence key hormones and behaviors involved in reproduction, specifically in migratory species with gonadal recrudescence largely occurring at breeding sites. We investigated the behavioral and physiological responses of male European starlings to the sequential addition of nest boxes and nesting material, green herbs, and female conspecifics and how these responses depend on the availability of certain antioxidants (anthocyanins) in the diet. As expected, cloacal protuberance volume and plasma testosterone of males generally increased with photoperiod. More notably, testosterone levels peaked in males fed the high antioxidant diet when both nest box and herbal cues were present, while males fed the low antioxidant diet showed no or only a muted testosterone response to the sequential addition of these environmental cues; thus our results are in agreement with the oxidation handicap hypothesis. Males fed the high antioxidant diet maintained a constant frequency of breeding behaviors over time, whereas those fed the low antioxidant diet decreased breeding behaviors as environmental cues were sequentially added. Overall, sequential addition of the environmental cues modulated physiological and behavioral measures of reproductive condition, and dietary antioxidants were shown to be a key factor in affecting the degree of response to each of these cues. Our results highlight the importance of supplementary environmental cues and key resources such as dietary antioxidants in enhancing breeding condition of males, which conceivably aid in attraction of high quality females and reproductive success., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Photoperiodic control of reproduction in birds is based on two processes, a positive effect leading to gonadal maturation and an inhibitory effect subsequently inducing regression. Nonphotoperiodic cues can modulate photoperiodic control, particularly the inhibitory process. In previous studies of common starlings (Sturnus vulgaris), (1) restriction of food availability to 8 h after dawn had little effect on testicular maturation but dramatically delayed subsequent regression and (2) lower ambient temperature also had little effect during maturation but delayed regression. Could the effects of food restriction and temperature share a common underlying mechanism? Four groups of starlings were kept on a simulated natural cycle in photoperiod in a 2 × 2 factorial experimental design. Two groups were held under an ambient temperature of 16°C, and the other two were held under 6°C. One of each of these groups had food provided ad lib., and in the other two groups access to food was denied 7 h after dawn. In both the ad lib. food groups and the food-restricted groups, lower temperature had little effect on testicular maturation but delayed subsequent regression and molt. In both the 16°C groups and the 6°C groups, food restriction had no effect on testicular maturation but delayed regression and molt. The daily cycle in body temperature was recorded in all groups when the photoperiod had reached 12L∶12D, the photoperiod at which regression is initiated. In both 6°C groups, nighttime body temperature was lower than in the 16°C groups, a characteristic of shorter photoperiods. In the two ad lib. food groups high daytime temperature was maintained until dusk, whereas in the two food-restricted groups body temperature began to decrease after food withdrawal. Thus, both lower temperature and food restriction delayed regression, as if the photoperiod was shorter than it actually was, and both resulted in daily cycles in body temperature that reflected cycles under shorter photoperiods. This implies that the daily cycle in body temperature is possibly a common pathway through which nonphotoperiodic cues may operate.

Development of avian embryos requires thermal energy, usually from parents. Parents may, however, trade off catering for embryonic requirements against their own need to forage through intermittent incubation. This dynamically adjusted behaviour can be affected by properties of the nest. Here, we experimentally show a novel mechanism by which parents, through incorporation of aromatic herbs into nests, effectively modify their incubation behaviour to the benefit of their offspring. Our study species, the European starling, includes in its nest aromatic herbs which promote offspring fitness. We provided wild starlings with artificial nests including or excluding the typically selected fresh herbs and found strong support for our prediction of facilitated incubation. Herb effects were not explained by thermal changes of the nests per se , but by modified parental behaviours. Egg temperatures and nest attendance were higher in herb than herbless nests, egg temperatures dropped less frequently below critical thresholds and parents started their active day earlier. These effects were dynamic over time and particularly strong during early incubation. Incubation period was shorter in herb nests, and nestlings were heavier one week after hatching. Aromatic herbs hence influenced incubation in beneficial ways for offspring, possibly through pharmacological effects on incubating parents., (© 2018 The Author(s).)

Environmental resources are proposed to fine-tune the timing of breeding, yet how they may do so remains unclear. In female European starlings ( Sturnus vulgaris ), nest cavities are limited resources that are necessary for breeding. Females that explore nest cavities, compared with those that do not, readily perform sexually motivated behaviors. We assigned female starlings to aviaries with: (1) no nest boxes, (2) nest boxes, or (3) nest boxes, plants, flowing water, insects and berries to test the hypothesis that environmental resources alter neural systems to stimulate mating behavior. Compared with other females, females that were housed with and explored nest boxes had higher estradiol, higher preproenkephalin (PENK) mRNA and lower levels of D1 and D2 dopamine receptor mRNA in the medial preoptic area (mPOA); a region in which opioids and dopamine modify female sexual behaviors and sexual motivation. Additionally, in the mPOA, PENK and tyrosine hydroxylase mRNA positively predicted, whereas estrogen receptor beta mRNA negatively predicted, nest box exploration. In the ventromedial hypothalamus (a region in which estradiol acts to stimulate sexual behavior), estrogen receptor alpha mRNA was highest in females that had access to but did not explore nest cavities. It is likely that seasonal increases in estradiol modify mRNA in the mPOA to facilitate nest cavity exploration. It is also possible that nest cavity exploration further alters gene expression in the mPOA, functioning to coordinate mating with resource availability. Thus, nest cavity exploration may be a form of self-stimulation that alters neural systems to fine-tune sexual behavior., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The peregrine falcon Falco peregrinus is renowned for attacking its prey from high altitude in a fast controlled dive called a stoop. Many other raptors employ a similar mode of attack, but the functional benefits of stooping remain obscure. Here we investigate whether, when, and why stooping promotes catch success, using a three-dimensional, agent-based modeling approach to simulate attacks of falcons on aerial prey. We simulate avian flapping and gliding flight using an analytical quasi-steady model of the aerodynamic forces and moments, parametrized by empirical measurements of flight morphology. The model-birds' flight control inputs are commanded by their guidance system, comprising a phenomenological model of its vision, guidance, and control. To intercept its prey, model-falcons use the same guidance law as missiles (pure proportional navigation); this assumption is corroborated by empirical data on peregrine falcons hunting lures. We parametrically vary the falcon's starting position relative to its prey, together with the feedback gain of its guidance loop, under differing assumptions regarding its errors and delay in vision and control, and for three different patterns of prey motion. We find that, when the prey maneuvers erratically, high-altitude stoops increase catch success compared to low-altitude attacks, but only if the falcon's guidance law is appropriately tuned, and only given a high degree of precision in vision and control. Remarkably, the optimal tuning of the guidance law in our simulations coincides closely with what has been observed empirically in peregrines. High-altitude stoops are shown to be beneficial because their high airspeed enables production of higher aerodynamic forces for maneuvering, and facilitates higher roll agility as the wings are tucked, each of which is essential to catching maneuvering prey at realistic response delays.

Animals communicate in distinct social contexts to convey information specific to those contexts, such as sexual or agonistic motivation. In seasonally-breeding male songbirds, seasonal changes in day length and increases in testosterone stimulate sexually-motivated song directed at females for courtship and reproduction. Dopamine and testosterone may act in the same brain regions to stimulate sexually-motivated singing. The neuropeptide neurotensin, acting at the neurotensin receptor 1 (NTR1), can strongly influence dopamine transmission. The goal of this study was to gain insight into the degree to which seasonal changes in physiology modify interactions between neurotensin and dopamine to adjust context-appropriate communication. Male European starlings were examined in physiological conditions that stimulate season-typical forms of communication: late summer/early fall non-breeding condition (low testosterone; birds sing infrequently), late fall non-breeding condition (low testosterone; birds produce non-sexually motivated song), and spring breeding condition (high testosterone; males produce sexually-motivated song). Double fluorescent immunolabeling was performed to detect co-localization patterns between tyrosine hydroxylase (TH; the rate-limiting enzyme in dopamine synthesis) and NTR1 in brain regions implicated in motivation and song production (the ventral tegmental area, medial preoptic nucleus, periaqueductal gray, and lateral septum). Co-localization between TH and NTR1 was present in the ventral tegmental area for all physiological conditions, and the number of co-localized cells did not differ across conditions. Immunolabeling for TH and NTR1 was also present in the other examined regions, although no co-localization was seen. These results support the hypothesis that interactions between NTR1 and dopamine in the ventral tegmental area may modulate vocalizations, but suggest that testosterone- or photoperiod-induced changes in NTR1/TH co-localization do not underlie seasonally-appropriate adjustment of communication., (Copyright © 2018 Elsevier B.V. All rights reserved.)

The avian dorsal telencephalon has two vast territories, the nidopallium and the mesopallium, both of which have been shown to contribute substantially to higher cognitive functions. From their connections, these territories have been proposed as equivalent to mammalian neocortical layers 2 and 3, various neocortical association areas, or the amygdala, but whether these are analogies or homologies by descent is unknown. We investigated the molecular profiles of the mesopallium and the nidopallium with RNA-seq. Gene expression experiments established that the mesopallium, but not the nidopallium, shares a transcription factor network with the intratelencephalic class of neocortical neurons, which are found in neocortical layers 2, 3, 5, and 6. Experiments in alligators demonstrated that these neurons are also abundant in the crocodilian cortex and form a large mesopallium-like structure in the dorsal ventricular ridge. Together with previous work, these molecular findings indicate a homology by descent for neuronal cell types of the avian dorsal telencephalon with the major excitatory cell types of mammalian neocortical circuits: the layer 4 input neurons, the deep layer output neurons, and the multi-layer intratelencephalic association neurons. These data raise the interesting possibility that avian and primate lineages evolved higher cognitive abilities independently through parallel expansions of homologous cell populations., (Copyright © 2018 Elsevier Ltd. All rights reserved.)