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Animals rely on movement to explore and exploit resources in their environment. While movement can provide energetic benefits, it also comes with energetic costs. This study examines how group phenotypic composition influences individual speed and energy expenditure during group travel in homing pigeons. We manipulated the composition of pigeon groups based on body mass and leadership rank. Our findings indicate that groups of 'leader' phenotypes show faster speeds and greater cohesion than 'follower' phenotype groups. Additionally, we show that groups of homogenous mass composition, whether all heavy or all light, were faster and expended less energy over the course of a whole flight than flocks composed of a mixture of heavy and light individuals. We highlight the importance of considering individual-level variation in social-level studies, and the interaction between individual and group-level traits in governing speed and the costs of travel., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

The ability of birds to fly through cluttered environments has inspired biologists interested in understanding its underlying mechanisms, and engineers interested in applying its underpinning principles. To analyse this problem empirically, we break it down into two distinct, but related, questions: How do birds select which gaps to aim for? And how do they steer through them? We answered these questions using a combined experimental and modelling approach, in which we released pigeons (Columbia livia domestica) inside a large hall with an open exit separated from the release point by a curtain creating two vertical gaps - one of which was obstructed by an obstacle. We tracked the birds using a high-speed motion capture system, and found that their gap choice seemed to be biased by their intrinsic handedness, rather than determined by extrinsic cues such as the size of the gap or its alignment with the destination. We modelled the pigeons' steering behaviour algorithmically by simulating their flight trajectories under a set of six candidate guidance laws, including those used previously to model target-oriented flight behaviours in birds. We found that their flights were best modelled by delayed proportional navigation commanding turning in proportion to the angular rate of the line-of-sight from the pigeon to the midpoint of the gap. Our results are consistent with this being a two-phase behaviour, in which the pigeon heads forward from the release point before steering towards the midpoint of whichever gap it chooses to aim for under closed-loop guidance. Our findings have implications for the sensorimotor mechanisms that underlie clutter negotiation in birds, uniting this with other kinds of target-oriented behaviours including aerial pursuit., Competing Interests: Competing interests N.P.-C.A. and G.K.T. declare no competing interests., (© 2023. Published by The Company of Biologists Ltd.)

The visual control of pecking by pigeons ( Columba livia ) has latterly been thought to be restricted to the fixation stops interrupting their downward head movements because these stops prevent interference by motion blur. Pigeons were also assumed to close their eyes during the final head thrust of the peck. Here, we re-examined their pecking motions using high-speed video recordings and supplementary provisions that permitted a three-dimensional spatial analysis of the movement, including measurement of pupil diameter and eyelid slit width. The results confirm that pigeons do not close their eyes completely during the presumed optically ballistic phase of pecking. Instead, their eyelids are narrowed to a slit. The width of this slit is sensitive to both the ambient illumination level and the visual background against which seed targets have to be detected and grasped. There is also evidence of some interaction between pupil diameter and eyelid slit width. We surmise that besides being an eye-protecting reflex, the partial covering of the pupil with the eyelids may increase the depth of focus, enabling pigeons to obtain sharp retinal images of peck target items at very close range and during the beak-gape 'handling' of food items and occasional grit particles., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Pecking in birds is analogous to reaching and grasping movements in primates. Earlier studies on visuomotor control in birds, which were conducted mostly in pigeons, suggested that avian pecking is controlled feedforwardly, and is out of the control of visual guidance during movement. However, recent studies using crows suggested a role of vision in pecking control during movement. To unveil what visuomotor mechanisms underlie the flexibility of pecking in crows, we examined whether pigeons and crows adjust their pecking to the visual distortion induced by prisms. Because prisms induce visual shifts of object positions, birds were required to adjust their movements. Pecking kinematics were examined before and after attaching prisms in front of the birds' eyes. Analysis of lateral deviation caused by the prisms showed that crows rapidly adjusted their pecking trajectories, but pigeons did so slowly. Angular displacement also increased in pigeons after attachment of the prism, but decreased in crows. These responses to prisms were consistent among individuals in pigeons but varied in crows, though the adjustment of pecking commonly succeeded in crows. These results suggest that pecking in pigeons predominantly involves feedforward control and that the movement is determined depending on the visual information available before the initiation of pecking. In contrast, the results from crows suggest that their pecking trajectories are corrected during the movement, supporting on-line visual control. Our findings provide the first evidence to suggest the on-line visual control of pecking in birds., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Gaze behavior offers valuable insights into attention and cognition. However, technological limitations have prevented the examination of animals' gaze behavior in natural, information-rich contexts; for example, during navigation through complex environments. Therefore, we developed a lightweight custom-made logger equipped with an inertial measurement unit (IMU) and GPS to simultaneously track the head movements and flight trajectories of free-flying homing pigeons. Pigeons have a limited range of eye movement, and their eye moves in coordination with their head in a saccadic manner (similar to primate eye saccades). This allows head movement to act as a proxy for visual scanning behavior. Our IMU sensor recorded the 3D movement of the birds' heads in high resolution, allowing us to reliably detect distinct saccade signals. The birds moved their head far more than necessary for maneuvering flight, suggesting that they actively scanned the environment. This movement was predominantly horizontal (yaw) and sideways (roll), allowing them to scan the environment with their lateral visual field. They decreased their head movement when they flew solo over prominent landmarks (major roads and a railway line) and also when they flew in pairs (especially when flying side by side, with the partner maintained in their lateral visual field). Thus, a decrease in head movement indicates a change in birds' focus of attention. We conclude that pigeons use their head gaze in a task-related manner and that tracking flying birds' head movement is a promising method for examining their visual attention during natural tasks., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

The complex and elaborate vocalizations uttered by many of the 10,000 extant bird species are considered a major driver in their evolutionary success, warranting study of the underlying mechanisms of vocal production. Additionally, birdsong has developed into a highly productive model system for vocal imitation learning and motor control, where, in contrast to humans, we have experimental access to the entire neuromechanical control loop. In human voice production, complex laryngeal geometry, vocal fold tissue properties, airflow and laryngeal musculature all interact to ultimately control vocal fold kinematics. Quantifying vocal fold kinematics is thus critical to understanding neuromechanical control of voiced sound production, but in vivo imaging of vocal fold kinematics in birds is experimentally challenging. Here, we adapted and tested electroglottography (EGG) as a novel tool for examining vocal fold kinematics in the avian vocal organ, the syrinx. We furthermore imaged and quantified syringeal kinematics in the pigeon ( Columba livia ) syrinx with unprecedented detail. Our results show that EGG signals predict (1) the relative amount of contact between the avian equivalent of vocal folds and (2) essential parameters describing vibratory kinematics, such as fundamental frequency, and timing of syringeal opening and closing events. As such, EGG provides novel opportunities for measuring syringeal vibratory kinematic parameters in vivo Furthermore, the opportunity for imaging syringeal vibratory kinematics from multiple planar views (horizontal and coronal) simultaneously promotes birds as an excellent model system for studying kinematics and control of voiced sound production in general, including in humans and other mammals., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Poor environmental conditions experienced during early development can have negative long-term consequences on fitness. Animals can compensate for negative developmental effects through phenotypic plasticity by diverting resources from non-vital to vital traits such as spatial memory to enhance foraging efficiency. We tested in young feral pigeons ( Columba livia ) how diets of different nutritional value during development affect the capacity to retrieve food hidden in a spatially complex environment, a process we refer to as 'spatial memory'. Parents were fed with either high- or low-quality food from egg laying until young fledged, after which all young pigeons received the same high-quality diet until memory performance was tested at 6 months of age. The pigeons were trained to learn a food location out of 18 possible locations in one session, and then their memory of this location was tested 24 h later. Birds reared with the low-quality diet made fewer errors in the memory test. These results demonstrate that food quality during development has long-lasting effects on memory, with a moderate nutritional deficit improving spatial memory performance in a foraging context. It might be that under poor feeding conditions resources are redirected from non-vital to vital traits, or pigeons raised with low-quality food might be better in using environmental cues such as the position of the sun to find where food was hidden., (© 2017. Published by The Company of Biologists Ltd.)

The complexity of low speed maneuvering flight is apparent from the combination of two critical aspects of this behavior: high power and precise control. To understand how such control is achieved, we examined the underlying kinematics and resulting aerodynamic mechanisms of low speed turning flight in the pigeon (Columba livia). Three birds were trained to perform 90 deg level turns in a stereotypical fashion and detailed three-dimensional (3D) kinematics were recorded at high speeds. Applying the angular momentum principle, we used mechanical modeling based on time-varying 3D inertia properties of individual sections of the pigeon's body to separate angular accelerations of the torso based on aerodynamics from those based on inertial effects. Directly measured angular accelerations of the torso were predicted by aerodynamic torques, justifying inferences of aerodynamic torque generation based on inside wing versus outside wing kinematics. Surprisingly, contralateral asymmetries in wing speed did not appear to underlie the 90 deg aerial turns, nor did contralateral differences in wing area, angle of attack, wingbeat amplitude or timing. Instead, torso angular accelerations into the turn were associated with the outside wing sweeping more anteriorly compared with a more laterally directed inside wing. In addition to moving through a relatively more retracted path, the inside wing was also more strongly pronated about its long axis compared with the outside wing, offsetting any difference in aerodynamic angle of attack that might arise from the observed asymmetry in wing trajectories. Therefore, to generate roll and pitch torques into the turn, pigeons simply reorient their wing trajectories toward the desired flight direction. As a result, by acting above the center of mass, the net aerodynamic force produced by the wings is directed inward, generating the necessary torques for turning., (© 2015. Published by The Company of Biologists Ltd.)

The gravity vector theory postulates that birds determine their position to set a home course by comparing the memorized gravity vector at the home loft with the local gravity vector at the release site, and that they should adjust their flight course to the gravity anomalies encountered. As gravity anomalies are often intermingled with geomagnetic anomalies, we released experienced pigeons from the center of a strong circular gravity anomaly (25 km diameter) not associated with magnetic anomalies and from a geophysical control site, equidistant from the home loft (91 km). After crossing the border zone of the anomaly--expected to be most critical for pigeon navigation--they dispersed significantly more than control birds, except for those having met a gravity anomaly en route. These data increase the credibility of the gravity vector hypothesis., (© 2014. Published by The Company of Biologists Ltd.)

We analysed the tracks of clock-shifted pigeons from six releases to determine how they cope with the conflict between their sun compass and other navigational cues. Time-lag embedding was used to calculate the short-term correlation dimension, a parameter that reflects the complexity of the navigational system, and with it, the number of factors involved. Initially, while pigeons were still at the release site, the short-term correlation dimension was low; it increased as the birds left the site, indicating that the birds were now actively navigating. Clock-shifted pigeons showed more scatter than the control birds, and their short-term correlation dimension became significantly smaller than that of the controls, remaining lower until the experimental birds reached their loft. This difference was small, but consistent, and suggests a different rating and ranking of the navigational cues. Clock-shifted pigeons do not seem to simply ignore the information from their manipulated sun compass altogether, but appear to merely downgrade it in favour of other cues, like their magnetic compass. This is supported by the observation that the final part of the tracks still showed a small deviation in the expected direction, indicating an effect of clock-shifting until the end of the homing flight., (© 2014. Published by The Company of Biologists Ltd.)

Homing pigeons (Columba livia) have been the central model of avian navigation research for many decades, but only more recently has research extended into understanding their mechanisms of orientation in the familiar area. The discovery (facilitated by GPS tracking) that pigeons gradually acquire with experience individually idiosyncratic routes home to which they remain faithful on repeated releases, even if displaced off-route, has helped uncover the fundamental role of familiar visual landmarks in the avian familiar area map. We evaluate the robustness and generality of the route-following phenomenon by examining extant studies in depth, including the single published counter-example, providing a detailed comparison of route efficiencies, flight corridor widths and fidelity. We combine this analysis with a review of inferences that can be drawn from other experimental approaches to understanding the nature of familiar area orientation in pigeons, including experiments on landmark recognition, and response to clock-shift, to build the first detailed picture of how bird orientation develops with experience of the familiar area. We articulate alternative hypotheses for how guidance might be controlled during route following, concluding that although much remains unknown, the details of route following strongly support a pilotage interpretation. Predictable patterns of efficiency increase, but limited to the local route, typical corridor widths of 100-200 m, high-fidelity pinch-points on route, attraction to landscape edges, and a robustness to clock-shift procedures, all demonstrate that birds can associatively acquire a map of their familiar area guided (at least partially) by direct visual control from memorised local landscape features.

Homing pigeons (Columba livia) are believed to adopt a map-and-compass strategy to find their way home. Surprisingly, to date a clear demonstration of the use of a cognitive map in free-flight experiments is missing. In this study, we investigated whether homing pigeons use a mental map in which - at an unknown release site - their own position, the home loft and a food loft are represented simultaneously. In order to test this, homing pigeons were trained to fly to a 25-30 km distant food loft. A total of 131 hungry and satiated pigeons were then released from an unfamiliar site equidistant from the food loft and the home loft. Their vanishing bearings and homing times were assessed conventionally at four sites, and also their flight tracks from one release site by means of GPS loggers. The vanishing bearings of fed and hungry birds differed significantly at all release sites and a highly significant proportion of hungry birds flew to the food loft, while the fed birds headed home. The GPS experiment revealed a number of pigeons flying very precisely to the food loft, others correcting their flight direction after topography-induced detours. This implies that the pigeons knew their geographical position in relation to the targets, and chose a flight direction according to their locally manipulated needs - clearly the essence of a cognitive navigational map.

The cues by which homing pigeons are able to return to a home loft after displacement to unfamiliar release sites remain debated. A number of experiments in which migratory birds have been treated with a magnetic pulse have produced a disruption in their orientation, which argues that a ferrimagnetic sense is used for navigation in birds. One previous experiment has also indicated an effect of magnetic pulses on homing pigeon navigation, although with inconsistent results. Previous studies have shown that some magnetic-related information is transmitted by the trigeminal nerve to the brain in some bird species, including the homing pigeon. The function of this information is still unclear. It has been suggested that this information is important for navigation. Previous studies with trigeminal nerve lesioned homing pigeons have clearly shown that the lack of trigeminally mediated information, even if magnetic, is not crucial for homing performance. However, this result does not completely exclude the possibility that other ferrimagnetic receptors in the homing pigeon play a role in navigation. Additionally, recent studies on homing pigeons suggested the existence of a ferrimagnetic sense in a novel location presumably located in the inner ear (lagena). In the present study, we tested whether any ferrimagnetic magnetoreceptors, irrespective of their location in the bird's head, are involved in pigeons' homing. To do this, we treated homing pigeons with a strong magnetic pulse before release, tracked birds with GPS loggers and analyzed whether this treatment affected homing performance. In the single previous magnetic pulse experiment on homing pigeons, only initial orientation at a release site was considered and the results were inconsistent. We observed no effect of the magnetic pulse at any of the sites used on initial orientation, homing performance, tortuosity or track efficiency, which does not support a role for the ferrimagnetic sense in homing pigeon navigation, at least not in this geographic area, where magnetic field variations are in the region of 200 nT intensity and 0.8 deg inclination.