A Comparison of Homing Behavior in Feral and Homing Pigeons

The social system of 4 adult Chinese hamsters was analysed under semi-natural conditions in a large enclosure. In all trials the home ranges overlapped, but were patterned in a different way and patrolled in turn. ♂♂ were inferior and less aggressive than ♀♀ and had to change their nest site more frequently. The observed social organization is based on a dominance order influenced by home range site and by activity.     

Does Familiarity with the Release Site Influence the Initial Orientation of Homing Pigeons? Experiments with Clock-shifted Birds

Experiments were performed to test whether the familiarity with the release site plays a role in the initial orientation of homing pigeons. Repeated releases of 6 h clock-shifted birds from the same site during the shifting time produced an improvement of their initial orientation: the shift effect decreased progressively. Since in subsequent releases from unfamiliar sites the shift effect reappears, the course correction observed at the familiar site is attributable to local stimuli and not to a general recalibration of the sun compass.     

Superparamagnetic Magnetite in the Upper Beak Tissue of Homing Pigeons

Homing pigeons have been subject of various studies trying to detect magnetic material which might be involved in magnetic field perception. Here we focus on the upper-beak skin of homing pigeons, a region that has previously been shown to contain nerves sensitive to changes of the ambient magnetic field. We localized Fe³⁺ concentrations in the subcutis and identified the material by transmission electronmicroscopy (TEM) as aggregates of magnetite nanocrystals (with grain sizes between 1 and 5 nm). The particles form clusters of 1–3 m diameter, which are arranged in distinct coherent elongated structures, associated with nervous tissue and located between fat cells. Complementary low-temperature magnetic measurements confirm the microscopic observations of fine-grained superparamagnetic particles in the tissue. Neither electron-microscopic nor magnetic measurements revealed any single-domain magnetite in the upper-beak skin tissue.     

Attempts to Determine the Roles of Visual and Olfactory Inputs in Initial Orientation and Homing of Pigeons over Familiar Terrain

The role of familiar visual landmarks in pigeon homing is still unsettled. If they are involved, they must be thought to be utilized in parallel with olfactory signals. In order to recognize the effectiveness of either one of the input channels separately, vision and olfaction, it is therefore necessary to interfere with both of them. Pigeons were temporarily deprived of image vision by spectacles made of translucent white paper, producing a condition called V^- as compared with the largely unimpaired condition V⁺. Access to olfactory signals was temporarily prohibited by charcoal filters before release and nasal anaesthesia upon release, resulting in condition O^- versus unimpaired smelling of natural air in O⁺. Prior to the test releases, all the participating pigeons had been made familiar with the two or four test sites (and with other sites in the area) by training flights. According to decreasing levels of initial homeward orientation and homing performance, the four combinations of treatments are to be arranged in the following order: V⁺O⁺, V⁺O^-, V^-O⁺, V^-O^-. In the last type, no trace of initial homeward orientation remained, indicating that at least one of the two input channels need be functional to enable home-related orientation. Over well known terrain, either alone seems to be more or less sufficient. The results are indicative but not definitely conclusive. Effects of visual impairment on behavioural activities in general cannot clearly be separated from orientation-specific effects. Therefore, and because many pigeons refuse to fly in V^- condition or fail to provide useful data, we stopped applying this method.     

Homing in German Cockroaches, Blattella germanica (L.) (Insecta: Dictyoptera): Multi-Channelled Orientation Cues

Cockroaches use navigational cues to elaborate their return path to the shelter. Our experiments investigated how individuals weighted information to choose where to search for the shelter in situations where path integration, visual and olfactory cues were conflicting. We showed that homing relied on a complex set of environmental stimuli, each playing a particular part. Path integration cues give cockroaches an estimation of the position of their goal, visual landmarks guide them to that position from a distance, while olfactory cues indicate the end of the path. Cockroaches gave the greatest importance to the first cues they encountered along their return path. Nevertheless, visual cues placed beyond aggregation pheromone deposits reduced their arrest efficiency and induced search in the area near the visual cues.     

The Orientational Consequences of Flocking Behaviour in Homing Pigeons,Columba livia

Two experiments are described which investigate the orientational consequences of flocking in homing pigeons Columba livia. Previous experiments have shown that homing pigeons placed inside a clear-sided release box for 5 min before release from a familiar site have enhanced ground homing speed compared with those placed in an opaque-sided box. It is assumed that previewing the surrounding landscape allows for faster homing since a bird denied this information must accumulate the knowledge on release. In experiment 1, using the same technique developed in these experiments but releasing the birds in pairs we showed that within familiar areas, homing pigeons can exploit a partner that has acquired more information, allowing them to home more quickly. In experiment 2 we attempted to test three potential strategies which may occur during homing flights. The results do not conclusively distinguish between these three mechanisms but suggest that orientation of the pairs of birds is most likely to have resulted from a compromise of individual tendencies, or from following the best homer, but not from following a ‘governing leader’. The consequence of these mechanisms is discussed.     

Pineal Body and Magnetic Sensitivity: Homing in Pinealectomized Pigeons under Overcast Skies

Since birds use the earth's magnetic field for compass orientation when astronomical cues are lacking and it has recently been suggested that the pineal body is part of their magnetic compass, test releases have been performed in overcast conditions with pigeons deprived of the pineal body. On the whole, both experimental and control birds were capable of homeward orientation, though the bearings of experimental were rather more scattered. No differences in homing speed or success were recorded. Thus, the pineal body does not appear to play an important role in the homing of pigeons.     

The Effects of Zinc Sulphate Anosmia on Homing Pigeons,Columba livia,in a Homing and a Non-homing Experiment

There is debate over whether homing pigeons, Columba livia, use olfactory information as part of their navigational map. Antagonists of the theory argue that homing deficits noted in anosmic pigeons may be due to a non-specific impairment in general information processing. In Experiment I, we present data from a modest investigation describing the typical navigational deficits that occur following zinc sulphate-mediated anosmia. Our results are consistent with previous experiments that noted impairments in homing performance from unfamiliar locations of anosmic pigeons. Experiment II is a critical experiment that involved a spatial working memory paradigm; this paradigm consisted of testing zinc sulphate-treated birds in a forced-choice alternation task in a T-maze. This experiment allowed us to determine whether anosmic pigeons were impaired in memory performance, a robust measure of general information processing. There were no differences between the last day of training and a subsequent-test day when pigeons received an intranasal injection of zinc sulphate. This experiment suggests that zinc sulphate anosmia does not impair general information processing, supporting the hypothesis that homing pigeons use olfactory cues when homing from unfamiliar locations.     

Spatial Memory of Homing Pigeons,Columba livia,Tested in an Outdoor Aviary

Five homing pigeons were trained to dig up hidden seeds buried in one of eight sand-filled cups in an octagonal aviary. All birds located the correct cup with an accuracy well above chance, irrespective of whether the retention time was between 15 min and 2 1/2 h, or whether it was between 1 d and 4 d. Even after 10 mo, the birds remembered the correct cup with great accuracy. This indicates well-developed spatial memory capacities of long duration. The method used in this study, based on training pigeons to dig for hidden seeds in combination with an arbitrary standard task, seems highly suitable to test pigeons for their spatial abilities.     

Homing in Harvester Termites: Evidence of Magnetic Orientation

Colonies of Trinervitermes geminatus (Wasman) build a network of forked foraging trails. Homing workers choose, with few exceptions, those tracks at the forks of the pheromone trail network that lead direct to the nest. Assessments show that these workers are also able to distinguish between the homeward and outward direction along an unbranched trail. Experiments involving the alteration of the local natural magnetic field by means of a weak permanent magnet provide evidence that the termites recognise the home direction by means of magnetoreception.     

Pigeon Homing: Does Early Experience Determine what Cues are Used to Navigate?

To test the hypothesis that early experience might determine the nature of the cues used to navigate, homing pigeons were made anosmic by nerve section before they could experience the natural odours in the region of the loft. They were allowed to make free flights and trained by flock releases with intact controls. Next they were tested and compared with intact controls as well as birds made temporarily anosmic just before the experiment. Initial orientation and homing performance of the experimental birds were very poor and showed that the pigeons were unable to acquire an alternative mechanism of navigation.     

Collective Decision-Making in Homing Pigeons: Larger Flocks Take Longer to Decide but Do Not Make Better Decisions

Social animals routinely are challenged to make consensus decisions about movement directions and routes. However, the underlying mechanisms facilitating such decision-making processes are still poorly known. A prominent question is how group members participate in group decisions. We addressed this question by examining how flocks of homing pigeons (Columba livia) decide their homing direction. We released newly formed flocks varying in size and determined the time taken to choose a homing direction (decision-making period) and the accuracy of that choice. We found that the decision-making period increases exponentially with flock size, which is consistent with a participatory decision-making process. We additionally found that there is no effect of flock size on the accuracy of the decisions made, which does not match with current theory for democratic choices of flight directions. Our combined results are better explained by a participatory choice of leaders that subsequently undertake the flock directional decisions. However, this decision-making model would only entirely fit with our results if leaders were chosen based on traits other than their navigational experience. Our study provides rare empirical evidence elucidating decision-making processes in freely moving groups of animals.     

Localisation of the Putative Magnetoreceptive Protein Cryptochrome 1b in the Retinae of Migratory Birds and Homing Pigeons

Cryptochromes are ubiquitously expressed in various animal tissues including the retina. Some cryptochromes are involved in regulating circadian activity. Cryptochrome proteins have also been suggested to mediate the primary mechanism in light-dependent magnetic compass orientation in birds. Cryptochrome 1b (Cry1b) exhibits a unique carboxy terminus exclusively found in birds so far, which might be indicative for a specialised function. Cryptochrome 1a (Cry1a) is so far the only cryptochrome protein that has been localised to specific cell types within the retina of migratory birds. Here we show that Cry1b, an alternative splice variant of Cry1a, is also expressed in the retina of migratory birds, but it is primarily located in other cell types than Cry1a. This could suggest different functions for the two splice products. Using diagnostic bird-specific antibodies (that allow for a precise discrimination between both proteins), we show that Cry1b protein is found in the retinae of migratory European robins (Erithacus rubecula), migratory Northern Wheatears (Oenanthe oenanthe) and pigeons (Columba livia). In all three species, retinal Cry1b is localised in cell types which have been discussed as potentially well suited locations for magnetoreception: Cry1b is observed in the cytosol of ganglion cells, displaced ganglion cells, and in photoreceptor inner segments. The cytosolic rather than nucleic location of Cry1b in the retina reported here speaks against a circadian clock regulatory function of Cry1b and it allows for the possible involvement of Cry1b in a radical-pair-based magnetoreception mechanism.     

Orientation Behavior with and Without Visual Cues in Newly Hatched and Adult Praying Mantis

In the praying mantis, vision plays a major role even in newly hatched nymphs, which have compound eyes that are not yet fully developed. This study examines how this factor affects the visual orientation behavior of freely mobile Mantis religiosa. Mantises from three age groups (nymphs newly hatched to 2 h old, three-day-old nymphs, and three- to four-month-old female and male adults) were placed in a completely unprotected open area, either with or without visual cues in the surroundings. As a visual cue, five vertical rods with high-, medium- or low-luminance contrast, having a vertical extension of 45° and overall horizontal extension of 40°, were presented at a distance of 300 mm, simulating a group of plant stems with differing contrasts. The mantis search behavior, probability of reaction to the visual cues, distance at which the first target-related reaction occurred, and target approach behavior were investigated. It was found that the search behavior differed in the different age groups. With high-contrast visual cues, newly hatched nymphs performed similarly to adults, but this was not the case for medium- and low-contrast visual cues. Visual performance was greatly improved 3 days after hatching, presumably due to the complete hardening of the cuticle. Nevertheless, despite differences in visual acuity, it was found that even newly hatched nymphs used visual orientation mechanisms similar to those of adult mantises, including fixation, scanning and peering.     

Initial Orientation of Deer Mice (Peromyscus maniculatus) released on Snow in Homing Experiments

Im Winter 1966–67 wurden 19 Peromyscus maniculatus ein- oder mehrmals auf 100–500 m außerhalb ihrer Heimbezirke versetzt. Sie wurden bei Nacht in einem flachen mit Schnee bedeckten Acker freigelassen; die menschlichen Spuren waren symmetrisch um die Freilassungsplätze verteilt. Unter diesen Bedingungen verließen die Mäuse, die im Heimfinden erfolgreich waren, den Freilassungsort in die Heimrichtung ± 90°. Dieser Befund stützt die Hypothese, daß die von Lindenlaub (1955, 1960) und Bovet (1960) nachgewiesene potentielle Anfangsorientierung von verfrachteten Mäusen zum Heimplatz durch eine Orientierung zu asymmetrisch um die Tiere verteilten Deckungen (z. B. Mäuselöcher und Wechsel) in den ?üblichen” Feldsituationen vorübergehend maskiert oder aufgehoben wird.     

Use of a Magnetic Compass for Nocturnal Homing Orientation in the Palmate Newt, Lissotriton helveticus

Previous studies have shown that migrating palmate newts (Lissotriton helveticus) can rely on acoustic cues for orientation to breeding ponds. Nonetheless, although acoustic cues are reliable over relatively short distances, they are unlikely to account for the long‐distance homing demonstrated in several other species of newts. Most individuals of L. helveticus migrate only a few hundred meters (Diego‐Rasilla, F. J. & Luengo, R. M. 2007: Acoustic orientation in the palmate newt, Lissotriton helveticus. Behav. Ecol. Sociobiol. 61, 1329—1335), raising the possibility that this species may only utilize short‐distance cues (Joly, P. & Miaud, C. 1993: How does a newt find its pond? The role of chemical cues in migrating newts (Triturus alpestris). Ethol. Ecol. Evol. 5, 447—455; Russell, A. P., Bauer, A. M. & Johnson, M. K. 2005: Migration of amphibians and reptiles: an overview of patterns and orientation mechanisms in relation to life history strategies. In: Migration of Organisms (Elewa, M. T., ed). Springer‐Verlag, Berlin Heidelberg, pp. 151—203; Sinsch, U. 2006: Orientation and navigation in Amphibia. Mar. Freshw. Behav. Phy. 39, 65—71). Therefore, experiments were carried out to investigate the use of the geomagnetic field in the nocturnal homing orientation of L. helveticus. Tests were carried out at night in an outdoor circular arena, under total overcast sky that prevented access to celestial compass cues. Individual newts were tested in one of four symmetrical alignments of an earth‐strength magnetic field. We studied the orientation behaviour of newts from two breeding ponds located 9.05 km west‐southwest and 19 km east‐northeast of the testing site. The distribution of magnetic bearings from both groups of newts exhibited significant orientation in the homeward direction. These findings indicate that palmate newts are capable of long‐distance homing and are able to orient in the homeward direction at night using the magnetic compass as the sole source of directional (i.e., compass) information.