Pigeons’ (Columba livia) hierarchical organization of local and global cues in touch screen tasks

Redundant encoding of local and global spatial cues is a common occurrence in many species. However, preferential use of the each type of cue seems to vary across species and tasks. In the current study, pigeons (Columba livia) were trained in three experiments on a touch screen task which included redundant local positional cues and global spatial cues. Specifically, pigeons were required to choose the middle out of three choice squares, such that the position within the array provided local information and the location on the screen provided global information. In Experiment 1, pigeons were trained and tested on vertically aligned arrays. In Experiment 2, pigeons were trained and tested on horizontally aligned arrays, and in Experiment 3, pigeons were trained and tested with vertical, horizontal and diagonally aligned arrays. The results indicate that preference for cue type depends upon the type of spatial information being encoded. Specifically, on vertical and diagonally aligned arrays, pigeons preferred global cues, whereas on horizontally aligned arrays, pigeons preferred local cues.     

Stable panoramic views facilitate snap-shot like memories for spatial reorientation in homing pigeons

Following spatial disorientation, animals can reorient themselves by relying on geometric cues (metric and sense) specified both by the macroscopic surface layout of an enclosed space and prominent visual landmarks in arrays. Whether spatial reorientation in arrays of landmarks is based on explicit representation of the geometric cues is a matter of debate. Here we trained homing pigeons (Columba livia) to locate a food-reward in a rectangular array of four identical or differently coloured pipes provided with four openings, only one of which allowed the birds to have access to the reward. Pigeons were trained either with a stable or a variable position of the opening on pipes, so that they could view the array either from the same or a variable perspective. Explicit mapping of configural geometry would predict successful reorientation irrespective of access condition. In contrast, we found that a stable view of the array facilitated spatial learning in homing pigeons, likely through the formation of snapshot-like memories.     

Brieftauben

Homing pigeons Homing pigeons are well known for their excellent homing abilities which allow them to return to their lofts from unknown releasing sites more than hundreds of kilometres away. Several orientation mechanisms – sun compass, earth's magnetic field, olfactory cues, visual cues – are known to be involved in homing performance as well as parameters such as motivation and experience. New technology give an insight in their homing behaviour and track preferences and it is shown that homing pigeons physiology and neurobiology seem to be functionally adapted to homing. Pigeons races are still common and it is shown how the pigeon breeder tries to maximize the success of his pigeons.     

The social transmission of spatial information in homing pigeons

We adapted a technique to explore the social transmission of spatial information in homing pigeons Columba livia. Five demonstrator pigeons were first trained to find a food goal within an indoor arena. This arena consisted of nine lidded cups laid out within a 12x12 grid on the floor. The task was to find the goal cup and flip the lid to obtain the food hidden within. Once the demonstrators had reached criterion the experiment proper began. During stage 1 of the experiment, 10 target birds, which had not previously been trained to find the goal, were introduced to the spatial task either in isolation or paired with a demonstrator. We measured how long they took to complete the task, the number of squares crossed on the grid, and the number of incorrect lids flipped. In stage 2, the target birds were introduced to the arena a second time, by themselves, and we compared the performance of the birds in the two treatments. The pigeons that had been introduced to the task with a demonstrator in stage 1 walked further and made more incorrect choices when searching for the food goal in stage 2 than the pigeons that were introduced to the task alone. This indicates that pigeons learn a spatial, food-finding task more effectively when performing the task alone than when accompanied by a knowledgeable conspecific. We discuss possible reasons for this in the light of previous experiments. Copyright 1999 The Association for the Study of Animal Behaviour.     

Comparison of visual and olfactory reversal learning in pigeons

Pigeons trained to discriminate between either two differentodors or two different colored lights acquired their discriminationsat the same rate. When the discrimination problems were reversedwithin a modality, however, the birds using visual cues acquirednew discriminations more rapidly than in original learning (positivetransfer), whereas the birds using olfactory cues acquired thediscrimination reversal less rapidly. On subsequent reversals,pigeons in the visual task condition developed a successivediscrimination reversal set with repeated reversals of the stimuli,while those in the olfactory condition did not. In a secondexperiment designed to assess the acquisition of redundant cues,birds received additional training with visual and olfactorycues in compound as discriminative stimuli, and were then testedwith only visual or olfactory cues. Birds previously trainedwith odor attended to visual cues in the compound, whereas birdspreviously trained to discriminate between lights did not attendto odor cues until they were presented alone. These resultsdemonstrate that the selection of stimuli may play a crucialrole in the performance of successive discrimination reversalsand suggest that, in contrast to rats, birds selectively attendto visual over olfactory cues in discrimination learning.     

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.     

Experience Based Use of Landmark and Vector Based Orientation During Homing by the Ant Formica cunicularia (Hymenoptera: Formicidae)

Homing paths of Formica cunicularia foragers from an artificial food reward were analyzed on a familiar terrain and in displacement experiments on a familiar and an unfamiliar terrain. Foragers were tested either when relatively new to a foraging route (untrained group) or after a day’s experience with it (trained group). Untrained foragers followed direct homing paths to the nest site when tested in the familiar terrain but followed tortuous paths when displaced to the unfamiliar terrain. Trained foragers behaved similarly to untrained ones when tested from the food reward to the nest site in the familiar terrain but their behavior changed in displacements. Irrespective of the familiarity of the displacement site, these foragers followed paths taking them to the expected nest sites. The results showed that foragers did not rely on chemical cues for homing and revealed that untrained foragers disregarded path integration and were directed to the nest site when it is in their visual panorama. On the contrary, trained foragers may have relied on path integration on familiar and unfamiliar terrain. The results also demonstrated that experience greatly affected the preferential use of visual and vector based cues by foragers during homing.     

Hippocampal ablated homing pigeons show a persistent impairment in the time taken to return home

Summary Homing pigeons were subjected to either hippocampal or control anterior forebrain ablations to examine what role the hippocampus and related structures may play in homing behavior. One year after surgery, the test birds were released from five locations where they had never been before. Both groups were successful in orienting homeward from the release sites, indicating that the hippocampus is not necessary for the neural regulation of a pigeon's map and compass system. Nonetheless, hippocampal ablated pigeons were significantly poorer with respect to the time required to return home, indicating a homing performance impairment. Alternative hypotheses are discussed to explain this result, the most parsimonious being impaired ability on the part of the hippocampal ablated birds to direct a course homeward.     

Frequency shift discrimination: Can homing pigeons locate infrasounds by Doppler shifts?

Summary Homing pigeons can detect small shifts in sound frequency at 1, 2, 5, 10, and 20 Hz. Their thresholds range from a 1% shift at 20 Hz to a 7% shift at 1 Hz. The frequency shifts used were designed to simulate the natural Doppler shifts resulting from changes in flight path. Their ability was sufficiently sensitive to make it feasible that natural Doppler shifts can be detected. Further tests indicated that the birds can perform true frequency discrimination when all subjective amplitude cues are removed. These laboratory tests are further steps in a series designed to find out if homing pigeons use infrasounds as cues for orientation and navigation.We thank our colleague, Prof. Wm. T. Keeton for his encouragement and support and John Willis for technical assistance. This research was supported by Grants No. BNS 77-24903 and BNS 78-13016 from the National Science Foundation, and by Hatch funds.Present address: Department of Biology, University of Pittsburgh, and Section of Birds, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania 15260, USA     

Not just passengers: pigeons,Columba livia,can learn homing routes while flying with a more experienced conspecific

For animals that travel in groups, the directional choices of conspecifics are potentially a rich source of information for spatial learning. In this study, we investigate how the opportunity to follow a locally experienced demonstrator affects route learning by pigeons over repeated homing flights. This test of social influences on navigation takes advantage of the individually distinctive routes that pigeons establish when trained alone. We found that pigeons learn routes just as effectively while flying with a partner as control pigeons do while flying alone. However, rather than learning the exact route of the demonstrator, the paired routes shifted over repeated flights, which suggests that the birds with less local experience also took an active role in the navigational task. The efficiency of the original routes was a key factor in how far they shifted, with less efficient routes undergoing the greatest changes. In this context, inefficient routes are unlikely to be maintained through repeated rounds of social transmission, and instead more efficient routes are achieved because of the interaction between social learning and information pooling.     

空气污染对信鸽比赛个体归巢速度的影响

空气污染作为一种有害的环境因素,对人类及动物的生理、心理均有影响。在鸟类中,信鸽(Columba livia)是研究空气污染影响的理想模型。为探究空气污染的行为学效应,通过收集并筛选2018和2019年成都市信鸽协会春秋两个季节举办的64场赛事共285羽参赛5场及以上的信鸽不同空距等级下的归巢速度,利用混合线性模型分析了6种大气污染物(CO、NO₂、O₃、PM_2.5、PM₁₀、SO₂)及气温、季节、天气等环境因子对信鸽归巢速度的影响。结果表明,PM₁₀、SO₂、NO₂、O₃浓度与归巢速度间有显著正相关关系,即这些空气污染物浓度越高,信鸽归巢速度越快;而CO浓度与归巢速度显著负相关,即CO浓度越高,信鸽归巢速度越慢。通过模型评估及比较,发现O3模型是归巢速度的最佳拟合模型,其次是NO2模型。本研究表明,信鸽的归巢速度的确受到空气污染的影响,但潜在的影响机制如归巢动机假说和嗅觉强化假说仍需进一步研究。     

Pigeons identify individual humans but show no sign of recognizing them in photographs

Photographs, especially of humans, are widely used as stimuli in behavioural research with pigeons. Despite their abundant use, it is not clear to what extent pigeons perceive photographs as representing three-dimensional objects. To address this question, we trained 16 pigeons to identify individual, real-life humans. This discrimination depended primarily on visual cues from the heads of the persons. Subsequently, the pigeons were shown photographs of these individuals to test for transfer to a two-dimensional representation. Successful identification of a three-dimensional person did not facilitate learning of the corresponding photographs. These results demonstrate limitations of cross-recognition of complex objects and their photographs in pigeons.     

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.     

Defining Natural Categories in Acoustic Signals: Comparison of Three Methods Applied to ‘Chick-a-dee’ Call Notes

Chicks were trained to discriminate between two identical boxes on the basis of their position. Subsequently, the colour of parts of the positive (reinforced) box was changed and chicks were retrained. Results showed that chicks were more or less impaired during retraining depending on the spatial distribution of the changed stimuli. Chicks behaved as if a figure (a disc or a spot of dots) painted on a box was irrelevant to them, whereas they did respond to changes in the colour of a uniformly coloured box or of scattered dots painted on a box. Similar results were obtained in simultaneous discrimination learning tasks involving addition of cues (e.g. colour plus position). Addition of cues facilitated learning using boxes the same colour all over or with painted scattered dots, but not using boxes with a disc or a spot of dots. Furthermore, addition of shape and position information had different outcomes depending on the use of three-dimensional objects or of painted figures: learning facilitation occurred only using three-dimensional objects. Results are interpreted in terms of an “object hypothesis”, and the validity and usefulness of traditional terms such as cues is questioned.     

No evidence that consistent auditory cues facilitate learning of spatial relations among locations

Human participants searched in a dynamic three-dimensional computer-generated virtual-environment open-field search task for four hidden goal locations arranged in a diamond configuration located in a 5×5 matrix of raised bins. Participants were randomly assigned to one of two groups: Consistent or Inconsistent. All participants experienced 30 trials in which four goal locations maintained the same spatial relations to each other (i.e., a diamond pattern), but this diamond pattern moved to random locations within the 5×5 matrix from trial-to-trial. For participants in the Consistent group, each goal location within the pattern always provided a unique and consistent auditory cue throughout the experimental session. For participants in the Inconsistent group, the same distinct auditory cues were provided for each goal location; however, the locations of these auditory cues within the pattern itself were randomized from trial-to-trial throughout the experimental session. Results indicated that participants in both groups learned the spatial configuration of goal locations, but the presence of consistent auditory cues did not facilitate the learning of spatial relations among locations.     

Migratory‐stage sea lamprey Petromyzon marinus stop responding to conspecific damage‐released alarm cues after 4 h of continuous exposure in laboratory conditions

This study investigated the length of avoidance response of migratory‐stage sea lamprey Petromyzon marinus exposed continuously to conspecific damage‐released alarm cues for varying lengths of time in laboratory stream channels. Ten replicate groups of P. marinus, separated by sex, were exposed to either deionized water control or to P. marinus extract for 0, 2 or 4 h continuously. Petromyzon marinus maintained their avoidance response to the conspecific damage‐released alarm cue after continuous exposure to the alarm cue for 0 and 2 h but not 4 h. Beyond being one of the first studies in regards to sensory–olfactory adaptation–acclimation of fishes to alarm cues of any kind, these results have important implications for use of conspecific alarm cues in P. marinus control. For example, continuous application of conspecific alarm cue during the day, when P. marinus are inactive and hiding, may result in sensory adaptation to the odour by nightfall when they migrate upstream.     

The Role of “Outward-Journey Information” in Homing Experiments with Pigeons: New Data on Ontogeny of Navigation and General Survey

Potentially, homing from distant areas can be based on two different principles of navigation: (1) A path-integration mechanism records and integrates an animal's motions during the outward trip; it is independent of location-specific stimuli. (2) Site localization, by contrast, is performed by deducing the animal's position in relation to home from such stimuli. Hence the first mechanism entirely depends on an uninterrupted flow of “outward-journey information”. The second mechanism may but need not be independent of stimuli recorded during the outward journey. Homing of pigeons is evidently based on site localization. Empirical findings do not support the idea that in experiments using passive displacement path integration is involved in addition or alternatively. Also, there is no reason to assume that very young pigeons transitionally, for only few weeks, apply such a method (as has been concluded by Wiltschko & Wiltschko 1982, 1985, etc.). It is shown that very young pigeons require local olfactory signals for initial homeward orientation as do older birds (Fig. 1). They are not generally better at homeward orientation than older inexperienced pigeons and show similar deviations from home and preferences for a particular compass direction (Table 1, Fig. 2). Olfactory signals appear to be gathered, as good as conditions allow, during any stage of a homing experiment. No fundamental difference can be recognized between olfactory “outward-journey information”, “release-site information”, etc. Signals received at different times and sites before release may contribute by varying proportions to the initial-orientation patterns observed under varying circumstances.     

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.     

When Does Previewing the Landscape Affect Pigeon Homing?

The importance of visual landmarks during homing in pigeons (Columba livia) remains a contentious issue. Three experiments which explore the role of visual landmarks at release sites are reported here. The effects of releasing homing pigeons after a 5-minute period in either a clear or an opaque sided release box were investigated. In the clear sided box pigeons were able to observe local surroundings at a release site, but this view was obstructed in the opaque sided box. In experiment 1 pigeons were released from familiar locations close to home (between 2 and 5.6 km): being unable to view landmarks prior to release significantly slowed homing speeds. In experiment 2 pigeons were released at familiar locations further from home (between 8.4 and 10 km): being unable to view landmarks prior to release did not significantly affect homing speeds. In experiment 3 pigeons were trained to home from distant release sites but were tested at closer, unfamiliar sites located on the likely homing routes used by the pigeons in training. No significant difference in homing speeds were observed when pigeons were released from either the clear or opaque sided box. The significance of these results for understanding the role of visual landmarks within a pigeon's familiar area is discussed.     

Multi-modal Orientation Cues in Homing Pigeons

How homing pigeons displaced into unfamiliar territory findtheir way home has been the subject of extensive experimentationand debate. One reason for the controversy is that pigeons seemto use multiple cues. Clock-shifting experiments show that experiencedpigeons use the sun as a preferred compass; when it is not availablethey rely on magnetic cues. That pigeons can home successfullywhile wearing frosted lenses suggests that landmarks, whilenot an essential navigational cue, are important in the finalstages. The sensory basis of the "map" or position finding systemis probably equally or even more complicated. When conditionsaround the loft are suitable, pigeons may use olfactory cuesto find their way or might use some feature of the earth's magneticfield for their navigation. The Wiltschkos (1989) showed thatpigeons raised without free access to ambient odors are notdisoriented when anosmic while their siblings raised with freeaccess to the prevailing wind were disoriented. Similarly, siblingpigeons from two lofts in Lincoln, Massachusetts. were welloriented or totally disoriented when released at magnetic anomaliesunder sunny skies depending upon which of the two lofts theyhad been reared in. All of these experiments and many more suggestthat pigeons use multiple and redundant cues to find their wayhome. Further, there is the suggestion that which cues theyadopt may well be influenced by the characteristics of the areaaround the home loft in which they were reared.