Cultural Diffusion of a Novel Food-finding Behaviour in Urban Pigeons: an Experimental Field Test

The cultural diffusion of a novel food-finding behaviour, piercing the paper cover of a box containing seed, was experimentally provoked in two flocks of pigeons (Columba livia): a closed aviary flock and an open urban flock. In the urban flock, piercing behaviour spread to 24 previously naive individuals. The rate of cultural diffusion was more rapid in the urban flock than in the aviary. In both situations, many individuals simply fed from the discoveries of birds that pierced, but this effect slowed diffusion less in the open urban flock because experienced piercers left the flock and naive birds entered it. Individual trial-and-error learning and natural shaping can be ruled out as mechanisms of diffusion in the present case.     

Development of the navigational map in homing pigeons: effects of flight experience on orientation performance

Experiments have shown that pigeons, Columba livia, are able to develop navigational abilities even if reared and kept confined in an aviary, provided that they are exposed to the natural winds. In contrast, pigeons reared in a wind-screened aviary do not learn to navigate. Pigeons kept confined in a screened aviary when young do not learn to navigate even if, once they become adult, the screens are removed and the birds are exposed to natural winds for a period sufficiently long for map learning. In all of these experiments, pigeons were kept confined until the day of test release. In the present study we found that pigeons reared in a wind-screened aviary developed navigational abilities if, once adult, they were allowed to perform spontaneous flights around the loft. Nevertheless, their navigational performance never reached the level of controls.Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Pigeon Homing: The Navigational Map Developed in Adulthood is Based on Olfactory Information

Homing pigeons raised in a shielded aviary and kept confined until the day of the test releases are not able to navigate even if, once adult, the screens are removed and the birds are exposed to natural winds for a sufficiently long period for map learning. However, pigeons raised in the same condition but, once adult, allowed to perform spontaneous flights around the loft are able to develop navigational abilities which, however, never reach the level of controls. In the present study, we show that the navigational map learned by the adult birds, which had the possibility to perform spontaneous flights, is based on olfactory information.     

Transmission of avian pox from starlings to Rothchild's mynahs

Several wild species of birds, including starlings (Sturnus vulgaris), house sparrows (Passer domesticus) and pigeons (Columba livia) gained access to an aviary housing Rothchild's mynahs (Leucospar rothchildii) and over 100 additional birds representing a variety of species. Six of approximately 15 mynahs became infected with avian pox and all of them died. None of the other birds in the aviary developed lesions. Pox virus was isolated from mynah facial lesions on chicken chorioallantoic membrane and in duck embryo fibroblast cell culture. It did not produce lesions in white Leghorn chickens, but did produce lesions in 4 of 11 wild starlings captured outside the aviary. Results indicated the agent was an indigenous starling pox capable of infecting and producing disease in mynah birds. Destruction of the captive starlings and isolation of the remaining mynahs immediately stopped the mortality.     

The ontogeny of the homing pigeon navigational map: evidence for a sensitive learning period

Homing pigeons can learn a navigational map by relying on the heterogeneous distribution of atmospheric odours in the environment. To test whether there might be a sensitive period for learning an olfactory-based navigational map, we maintained a group of young pigeons in an aviary screened from the winds until the age of three to four months post-fledging. Subsequently, the screens were removed and the pigeons were exposed to the winds and the environmental odours they carry for three months. One control group of pigeons was held in a similar aviary but exposed to the winds immediately upon Hedging, while another control group of pigeons was allowed free-flight. When the pigeons from the three groups were released from two distant release sites at about six months of age post-fledging, the two control groups were found to be equally good at orientating and returning home, while the experimental pigeons held in the shielded aviary for the first three months post-fledging were unable to orientate homeward and they were generally unsuccessful in returning home. This result supports the hypothesis that environmental experience during the first three months post-fledging is critical for some aspect of navigational map learning and that navigational map learning displays sensitive period-like properties.     

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.     

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.     

Food-attraction conditioning in the snail,Helix pomatia

Adult pulmonate snails (Helix pomatia) were released equidistant between two types of food, carrot and potato, respectively. Naive snails moved in different directions and did not locate either food above chance, although both foods were readily eaten upon direct contact. After a single carrot feeding episode, 75% of the carrot-fed snails moved directly towards the carrot and ate it. Conversely, potato-fed snails located the potato in 67% of the cases. Snails that were fed apple or lettuce behaved like naive animals, with the majority of animals (75% in both cases) locating neither the carrot nor the potato.The ability of snails to locate this particular food after a single feeding episode was maintained for at least 11 days, provided that the snails were not exposed to other foods in the interim. If the animals were fed a different food (but still tested for food-finding ability to the initially conditioned food) their orientation preference decreased gradually over a period of 5 days.Although the snails' orientation is based upon olfactory cues, exposure to food odor alone is not sufficient to enable food-finding; additional feeding related stimuli are necessary.These findings indicate that Helix do not possess a predisposition for the foods tested, and further suggest that processes underlying food-finding and food selection are strongly influenced by learning experiences. The conditioning phenomenon underlying food-finding behavior has been called Food-Attraction Conditioning, and appears to be a crucial link between the ecologies of learning and foraging behaviors. The accessibility of the snail's nervous system should permit neuronal analysis of the mechanisms underlying such a unique and complex learning phenomenon.     

The social transmission of a food-finding technique in pigeons: what is learned?

Experimentally-naive pigeons (Columba livia) were exposed to varying amounts of socially transmitted information needed for solving a food-finding problem. Observer pigeons that saw a trained bird (model) piercing the paper covering a food box and eating were able to solve the problem faster and more efficiently than pigeons that only saw the model eating but not piercing. This result held whether observer performance was simultaneous or deferred with respect to the model's demonstration. Pigeons that saw the model piercing but not eating showed almost no tendency to copy. These results suggest that copying was dependent upon observer recognition of the fact that the model was getting a food reward, and that pigeons were capable of learning aspects of the piercing technique by observation rather than by trial and error.     

Accurate route demonstration by experienced homing pigeons does not improve subsequent homing performance in naive conspecifics.

We describe an experiment that uses the grouping tendencies and navigational abilities of the homing pigeon (Columba livia) to investigate the possibility of socially mediated information transfer in a field setting. By varying the composition of paired-release types, we allowed some naive birds to receive an accurate demonstration of the home route whilst others were paired with similarly naive conspecifics. After this 'paired phase', we predicted that if any learning of spatial information occurred then naive members of the former pairs would outperform their untutored conspecifics when re-released individually during the subsequent 'single phase' of the experiment. This prediction was not confirmed. Neither homing speed nor initial orientation was superior in individually released tutored versus untutored birds, despite the fact that both performance measures were better in the earlier 'paired phase' with experienced demonstrators. Our results suggest that although naive homing pigeons clearly interact with their experienced partners, they are unable to transfer any individually useful spatial information to subsequent homing flights.     

Sun compass orientation in seed-caching scrub jays (Aphelocoma coerulescens)

Summary In their natural environment, scrub jays harvest pinyon pine seeds and store them in subterranean caches. In our tests, the birds performed this behavior in an octagonal outdoor aviary with sand-filled cups inserted in the floor. For caching, only 12 such cups in a 90° sector were available, while for the recovery session 4 to 6 days later all 48 cups in the entire aviary were open. In control tests, the birds concentrated their search in the sector where the seeds had been cached. When the internal clock of the birds was shifted 6 h between caching and recovery, they preferentially probed in the adjacent 90° sector. This indicated that they used sun compass information to relocate their caches, largely ignoring visual cues from surrounding landmarks.The dominant role of the sun compass which has a parallel in the orientation of homing pigeons, may reflect a general tendency to prefer compass information in spatial orientation tasks; it is in agreement with the model that birds generally have a directionally oriented view of space.Abbreviations OR Original caches - SH shifted caches     

Influence of physical activity on tibial bone material properties in laying hens

Laying hens develop a type of osteoporosis that arises from a loss of structural bone, resulting in high incidence of fractures. In this study, a comparison of bone material properties was made for lines of hens created by divergent selection to have high and low bone strength and housed in either individual cages, with restricted mobility, or in an aviary system, with opportunity for increased mobility. Improvement of bone biomechanics in the high line hens and in aviary housing was mainly due to increased bone mass, thicker cortical bone and more medullary bone. However, bone material properties such as cortical and medullary bone mineral composition and crystallinity as well as collagen maturity did not differ between lines. However, bone material properties of birds from the different type of housing were markedly different. The cortical bone in aviary birds had a lower degree of mineralization and bone mineral was less mature and less organized than in caged birds. These differences can be explained by increased bone turnover rates due to the higher physical activity of aviary birds that stimulates bone formation and bone remodeling. Multivariate statistical analyses shows that both cortical and medullary bone contribute to breaking strengthThe cortical thickness was the single most important contributor while its degree of mineralization and porosity had a smaller contribution. Bone properties had poorer correlations with mechanical properties in cage birds than in aviary birds presumably due to the greater number of structural defects of cortical bone in cage birds.     

Echoic memory in pigeons

It is unknown whether birds are able to retain the memory of purely sensory auditory information such as white noise over an extended period of time. In a Pavlovian heart rate conditioning paradigm, four pigeons were trained to associate a mild electric shock with periodic random waveforms, and no shock with aperiodic noise. Periodic waveform detection requires echoic memory, i.e., the online retention of a waveform pattern over a limited time. Starting with 40ms, the waveform period was increased after successful learning until no significant stimulus discrimination could be found. Significant discrimination was achieved at periods of up to 2560ms. This is the first demonstration that echoic memory performance in birds is clearly superior to cats and gerbils, and comparable to naive human performance.     

Fecal corticosterone, body mass, and caching rates of Carolina chickadees (Poecile carolinensis) from disturbed and undisturbed sites

We tested for hormonal and behavioral differences between Carolina chickadees (Poecile carolinensis) taken from a disturbed (recently logged) forest, an undisturbed forest, or a residential site. We measured fecal corticosterone and body mass levels in the field, and fecal corticosterone, body mass, and caching behavior in an aviary experiment. In the field, birds from the disturbed forest exhibited significantly higher fecal corticosterone levels than birds from either the undisturbed forest or from the residential site. Birds from the disturbed forest also exhibited lower body mass than those from the undisturbed forest but higher body mass than those from the residential site. Our aviary results suggest that these physiological differences between field sites are the result of short-term responses to ecological factors: neither body mass nor fecal corticosterone levels varied between birds captured at different sites. Aviary sample sizes were sufficient to detect seasonal variation in fecal corticosterone (lowest in summer), body mass (highest in spring), and rate of gain in body mass (highest in winter). Under "closed-economy" aviary conditions (all food available from a feeder in the aviary), there were no site differences in the percent of seeds taken from the feeder that were cached. However, under "open-economy" conditions (food occasionally available ad libitum), significantly fewer seeds were cached by birds from the disturbed forest compared to the undisturbed or residential sites. On average, there was only a two-fold difference in population levels of fecal corticosterone. This difference is about the same as an increase in fecal corticosterone induced by a 2-h increase in food deprivation and cannot be considered to be an acute stress response to disturbance.     

Does pigeon homing depend on stimuli perceived during displacement?

Summary In order to test whether stimuli perceived during passive displacement are important for the subsequent homing, pigeons were transported in an apparatus designed to prevent them from receiving relevant information: The experimental birds were continuously rotated quite rapidly (1.5 cps, radial acceleration about 4 g); in addition, they were exposed to an artificial magnetic field and supplied with bottled air. Control birds were transported in open-air cages on top of the van with free view to all sides.Five pairs of releases from equal distances in opposite directions were conducted. Experienced birds were released at distances of about 15, 90, and 300 km from the loft, inexperienced birds at distances of about 180km. In each pair of releases both groups of pigeons were significantly homeward oriented. Neither in initial orientation nor in homing performance nor in the distribution of recoveries were the experimental birds inferior to the controls or in any perceptable way different from them.It is concluded that homing of passively displaced pigeons is not primarily based on information gathered during the outward journey.Abbreviations EP experimental pigeon(s) - CP control pigeon(s) The possibility to maintain our pigeon loft in a building that belongs to the Zoological Institute (Prof. M. Lindauer) of the University of Würzburg is gratefully acknowledged.     

Does Rearing Laying Hens in Aviaries Adversely Affect Long-Term Welfare following Transfer to Furnished Cages?

This study tests the hypothesis that hens that are reared in aviaries but produce in furnished cages experience poorer welfare in production than hens reared in caged systems. This hypothesis is based on the suggestion that the spatial restriction associated with the transfer from aviaries to cages results in frustration or stress for the aviary reared birds. To assess the difference in welfare between aviary and cage reared hens in production, non-beak trimmed white leghorn birds from both rearing backgrounds were filmed at a commercial farm that used furnished cage housing. The videos were taken at 19 and 21 weeks of age, following the birds'' transition to the production environment at 16 weeks. Videos were analysed in terms of the performance of aversion-related behaviour in undisturbed birds, comfort behaviour in undisturbed birds, and alert behaviour directed to a novel object in the home cage. A decrease in the performance of the former behaviour and increase in the performance of the latter two behaviours indicates improved welfare. The results showed that aviary reared birds performed more alert behaviour near to the object than did cage reared birds at 19 but not at 21 weeks of age (P = 0.03). Blood glucose concentrations did not differ between the treatments (P>0.10). There was a significant difference in mortality between treatments (P = 0.000), with more death in aviary reared birds (5.52%) compared to cage birds (2.48%). The higher mortality of aviary-reared birds indicates a negative effect of aviary rearing on bird welfare, whereas the higher duration of alert behavior suggests a positive effect of aviary rearing.     

Passive Immunization of Pigeons Against Trichomoniasis

SYNOPSIS Nonimmune homing pigeons Columba livia were infected with the Jones' Barn strain of Trichomonas gallinae and subsequently transfused with plasma from acute or chronically infected pigeons harboring one of 3 different strains of T. gallinae. The transfusions were either a single 2 ml dose given one day after inoculation or three 1 ml doses given 0, 5, and 10 days after inoculation. Plasma from pigeons harboring any of the 3 strains was capable of passively immunizing nonimmune birds. All birds which were immunized with plasma from infected pigeons survived until killed at the end of the test period and no visceral lesions were found on necropsy but trichomonads were present in the oropharynx. All controls (untreated or transfused with normal plasma) died of visceral trichomoniasis.
Immune plasma produced some lysis of trichomonads in vitro, and inhibition of motility and vacuolization occurred in some of the non-lysed organisms. The overall lytic activity in vitro affected less than 10% of the suspended trichomonads.     

The role of a social factor in exploration of a novel environment in great tits (Parus major) under conditions of limited space

Sequential tests in an open field (OF) and in an aviary were performed to study the exploration behavior of great tits, Parus major Each aviary of 2×2×2 m included a standard set of objects: trees, feeder, and shelters. Twenty-four aviaries included a single individual, and 49 aviaries included two heterosexual individuals. During the 15-min observation, a number of explored objects, the latent exploration period of each object (the time period from the start of observation to the first visit of each object), and the time period to the onset of feeding were recorded. The exploration rate of each individual was characterized by the sum of all latent periods (CLP). No association between the cumulative locomotor activities in OF and in an aviary was found. Probably, in the complex environment of an aviary, the frequency of directed actions increases, competing with the movement activity. Faster individuals in OF showed a shorter CLP; i.e., they were fast explorers. The exploration rate of a novel environment in the presence of socid partner was unrelated to exploration scores in OF. Interrelations of birds combined in an aviary were dependent on combinations of personality characteristics identified in OF.     

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.     

Aspects of risk-aversion in foraging white-crowned sparrows

Two series of aviary experiments with white-crowned sparrows (Zonotrichia leucophrys) are reported. The first series attempts to mimic foraging choices between microhabitats. The second series examines foraging choices made within a single artificial patch, a design intended to resemble dietary selection. In all experiments subjects chose between a constant food reward and a variable reward, the mean of which equalled the constant reward. Each experiment allowed the bird to feed at an overall rate greater than that minimally required for daily energy balance. In both series of experiments, the birds usually preferred the constant reward. Results of the first series do not differ significantly from previously reported preferences of dark-eyed juncos (Junco hyemalis), birds weighing 10 g less than the white-crowned sparrows.