Time‐of‐Day Discriminative Learning: Contrasting the Use of Spatial Compared to Feature Information in Homing Pigeons (Columba livia)

This study examined time‐of‐day associative learning to either spatial or feature information in homing pigeons in an open‐field, laboratory setting. Homing pigeons are well known for their navigational abilities and generally have been shown to rely more heavily on spatial than nonspatial cues in recognizing a goal. However, during goal localization, homing pigeons also successfully use nonspatial, feature information. Homing pigeons were divided into two groups and were trained to locate two time‐of‐day dependent, food reward sites using either discriminative spatial or feature information. Because of the importance of the hippocampus in controlling avian memory, we hypothesized that homing pigeons trained with spatial cues would be superior in learning the time‐of‐day discrimination compared to the pigeons trained with feature cues. Indeed, homing pigeons that were trained with spatial information outperformed the pigeons trained with feature information in learning the time‐of‐day discrimination task.     

Effects of stimulus size and spatial organization on pigeons’ conditional same-different discrimination

In two experiments, we explored the effects of varying the size and the spatial organization of the stimuli in multi-item arrays on pigeons’ same-different discrimination behavior. The birds had previously learned to discriminate a simultaneously presented array of 16 identical (Same) visual items from an array of 16 nonidentical (Different) visual items, when the correct choice was conditional on the presence of another cue: the color of the background (Castro et al., in press). In Experiment 1, we trained pigeons with 7-item arrays and then tested them with arrays containing the same item, but in a variety of sizes. In Experiment 2, we tested the birds with the items grouped in novel locations: the top, the bottom, the left, or the right portions of the display area, which generated different vertical and horizontal alignments. Accuracy scores revealed virtually perfect stimulus generalization across various item sizes and spatial organizations. Reaction times revealed that the birds perceived different sizes of a single icon as the same stimulus (Experiment 1) and that the birds processed vertical arrangements faster than horizontal arrangements (Experiment 2). These results suggest that the pigeons noticed both physical and spatial changes in the stimuli (as shown by their reaction times), but that these changes did not disrupt the birds’ discriminating the sameness or differentness of the multi-item arrays (as shown by their accuracy scores).     

Avian detection and identification of perceptual organization in random noise

Recent research has suggested that pigeons may have difficulty globally integrating visual information in hierarchically arranged stimuli. To isolate and understand the mechanisms responsible for processing emergent perceptual structure, three pigeons were tested in a two alternative choice task that required the global integration of organized local information. They were reinforced for localizing, on randomized distractor backgrounds of black and white square elements, different types of structured targets (e.g., stripes, squares, checkerboards) arranged from these same elements. These hierarchical stimuli were tested at four different levels of spatial granularity (i.e., different element sizes). Experiment 1 found rapid acquisition for the vertical and horizontal stripes or square targets and somewhat slower learning with the checkerboard pattern. Experiment 2 demonstrated successful transfer to a novel target types (alternating bars and "diagonal" stripes). In both experiments, displays with the greatest spatial granularity (smallest elements and most repetitive structure) monotonically supported the best discrimination. These results indicate pigeons can perceive and discriminate emergent visual structure under the right circumstances and suggest they do so with a generalized rule for detecting patterns of non-random perceptual structure.     

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.     

What is geometric information and how do animals use it?

Unlike investigations of animals’ use of spatial cues such as landmarks, studies of sensitivity to the geometry of surfaces in an enclosure have proceeded mostly as an attempt to explain a laboratory finding with few direct tests of how animals use such a cue in nature. In this brief review, I discuss the current debate over whether global or local information from the enclosure drives the typical rotational error pattern in such studies. A consideration of the form and function of geometric cues in natural settings suggests that the natural boundaries for which arena walls are considered analogous might better be thought as landmarks. With a clearer picture of what geometric information is and how it might be used in nature, the generality of findings from laboratory studies of geometry enclosure can be better assessed.     

Pigeon's recognition of cartoons: effects of fragmentation, scrambling, and deletion of elements

J. Cerella [Pattern Recognit. 12 (1980) 1] and more recently S. Watanabe [Behav. Proc. 53 (2001) 3] demonstrated that pigeons showed no decrement in recognizing cartoons that were spatially scrambled, indicating that pigeons' discriminative responding is controlled by local features alone. In contrast Kirkpatrick-Steger et al. [J. Exp. Psychol. Anim. Behav. Proc. 24 (1998) 34] used line drawings as stimuli and demonstrated the importance of spatial organization for picture recognition by pigeons, confirming related findings reported in their previous studies. The present study revisited the recognition of cartoons by pigeons. In Experiment 1, pigeons were trained to discriminate cartoon people on a variety of background scenes. Subsequent tests revealed that discriminative performances with both familiar and novel instances decreased as the objects and object-like parts were progressively fragmented, indicating that search for the targeted cartoon people in the stimulus array might have enhanced the pigeons to attend to global aspects of cartoon people. Experiment 2 used line drawings of cartoon faces as stimuli and examined effects of scrambling and deletion of components. A set of components (eyes and eyebrows) exerted strong control over behavior and scrambling only moderately suppressed responding. The results suggest that pigeons use both global and local aspects, with different mixtures of these types of information depending on the particular perceptual context.     

Negotiating a noisy, information-rich environment in search of cryptic prey: olfactory predators need patchiness in prey cues

1. Olfactory predator search processes differ fundamentally to those based on vision, particularly when odour cues are deposited rather than airborne or emanating from a point source. When searching for visually cryptic prey that may have moved some distance from a deposited odour cue, cue context and spatial variability are the most likely sources of information about prey location available to an olfactory predator. 2. We tested whether the house mouse (Mus domesticus), a model olfactory predator, would use cue context and spatial variability when searching for buried food items; specifically, we tested the effect of varying cue patchiness, odour strength, and cue-prey association on mouse foraging success. 3. Within mouse- and predator-proof enclosures, we created grids of 100 sand-filled Petri dishes and buried peanut pieces in a set number of these patches to represent visually cryptic 'prey'. By adding peanut oil to selected dishes, we varied the spatial distribution of prey odour relative to the distribution of prey patches in each grid, to reflect different levels of cue patchiness (Experiment 1), odour strength (Experiment 2) and cue-prey association (Experiment 3). We measured the overnight foraging success of individual mice (percentage of searched patches containing prey), as well as their foraging activity (percentage of patches searched), and prey survival (percentage of unsearched prey patches). 4. Mouse foraging success was highest where odour cues were patchy rather than uniform (Experiment 1), and where cues were tightly associated with prey location, rather than randomly or uniformly distributed (Experiment 3). However, when cues at prey patches were ten times stronger than a uniformly distributed weak background odour, mice did not improve their foraging success over that experienced when cues were of uniform strength and distribution (Experiment 2). 5. These results suggest that spatial variability and cue context are important means by which olfactory predators can use deposited odour cues to locate visually cryptic prey. They also indicate that chemical crypsis can disrupt these search processes as effectively as background matching in visually based predator-prey systems.     

Microhabitat use affects goby (Gobiidae) cue choice in spatial learning task

This study investigated whether spatial learning ability and cue use of gobies (Gobiidae) from two contrasting habitats differed in a spatial task. Gobies were collected from the spatially complex rock pools and dynamic, homogenous sandy shores. Fishes were trained to locate a shelter under the simulated threat of predation and it was determined whether they used local or extra‐maze (global) and geometric cues to do so. It was hypothesized that fishes from rock pools would outperform fishes from sandy shores in their ability to relocate shelter and the two groups would differ in their cue use. It was found that rock‐pool species learnt the location of the correct shelter much faster, made fewer errors and used a combination of all available cues to locate the shelter, while sand species relied significantly more on extra‐maze and geometric cues for orientation. The results reported here support the hypothesis that fishes living in complex habitats have enhanced capacity for spatial learning and are more likely to rely on local landmarks as directional cues than fishes living in mundane habitats where local cues such as visual landmarks are unreliable.     

The forest or the trees: preference for global over local image processing is reversed by prior experience in honeybees

Traditional models of insect vision have assumed that insects are only capable of low-level analysis of local cues and are incapable of global, holistic perception. However, recent studies on honeybee (Apis mellifera) vision have refuted this view by showing that this insect also processes complex visual information by using spatial configurations or relational rules. In the light of these findings, we asked whether bees prioritize global configurations or local cues by setting these two levels of image analysis in competition. We trained individual free-flying honeybees to discriminate hierarchical visual stimuli within a Y-maze and tested bees with novel stimuli in which local and/or global cues were manipulated. We demonstrate that even when local information is accessible, bees prefer global information, thus relying mainly on the object''s spatial configuration rather than on elemental, local information. This preference can be reversed if bees are pre-trained to discriminate isolated local cues. In this case, bees prefer the hierarchical stimuli with the local elements previously primed even if they build an incorrect global configuration. Pre-training with local cues induces a generic attentional bias towards any local elements as local information is prioritized in the test, even if the local cues used in the test are different from the pre-trained ones. Our results thus underline the plasticity of visual processing in insects and provide new insights for the comparative analysis of visual recognition in humans and animals.     

Use of water flow direction to provide spatial information in a small-scale orientation task

Experiments were designed to investigate whether three-spined sticklebacks Gasterosteus aculeatus can use direction of water flow as an orientation cue. The fish had to learn the location of a food patch in a channel where water flow direction was the only reliable indicator of the food patch position. Fish from two ponds and two rivers were trained and tested in the spatial task to determine whether river three-spined sticklebacks are more adept at using water flow as a spatial cue than fish from ponds. All fish were able to use water flow to locate the food patch but one of the two river groups was significantly faster at learning the patch location. When the task was reversed so that fish that had formerly been trained to swim downstream now had to learn to swim upstream and vice versa both river groups learned the reversed task faster than the two pond groups. In a second experiment, to investigate whether fish from ponds or rivers vary in the type of spatial cue that they prefer to use, fish from one pond and one river were given a choice between two different types of spatial cue: flow direction or visual landmarks. A test trial in which these two cues were put into conflict revealed that the river population showed a strong preference for flow direction whilst the pond population preferred to use visual landmarks.     

Orientation in pied flycatchers: the relative importance of magnetic and visual information at dusk

We investigated the orientation of juvenile pied flycatchers, Ficedula hypoleuca, during autumn migration in south Sweden using orientation cage experiments, to study the relative importance of visual and magnetic information at sunset. We performed cage tests under 12 experimental conditions that manipulated the geomagnetic and visual sunset cues available for orientation: natural clear skies in the local or a vertical magnetic field; simulated total overcast in the local or a vertical magnetic field; natural pattern of skylight polarization and directional information from stars screened off, with the sun's position as normal or shifted 120 degrees anticlockwise with mirrors; reduced polarization in the local or a vertical magnetic field; directions of polarization (e-vector) NE/SW and NW/SE, respectively, in the local or a vertical magnetic field. The pied flycatchers were significantly oriented towards slightly south of west when they could use a combination of skylight and geomagnetic cues. The mean orientation was significantly shifted along with the deflection of the sunset position by mirrors. Reduced polarization had no significant effect on orientation either in the local, or in a vertical, magnetic field. The birds tended to orient parallel with the axis of polarization, but only when the artificial e-vector was aligned NW/SE. The mean orientation under simulated total overcast in a vertical, and in the local, magnetic field was not significantly different from random. It is difficult to rank either cue as dominant over the other and we conclude that both visual and magnetic cues seem to be important for the birds' orientation when caught and tested during active migration. Copyright 1999 The Association for the Study of Animal Behaviour.     

Orientation of attention in the visual space]

The display was composed of four boxes, horizontally aligned above the fixation point. In Experiment I, each box was cued by a digit shown at fixation. In Experiment II there were only two numeric cues, signalling the inner or the outer boxes, depending on the experimental condition. The subject was instructed to orient attention to the cued box, and to respond to the imperative stimulus as fast as possible, wherever it appeared. By using four time interval (SOAs), we tried to determine the route covered by attention movements. In Experiment I, with the shortest SOA (100 msec), it was shown that attention does not reach the cued box through a direct path. Rather it moves first on the inner boxes, thereafter focusing on the cued location. The same results were obtained in Experiment II, where the cue directed attention to the inner boxes. When the external boxes were cued, however, this trend was not observed.     

Magnetic versus celestial cues: cue-conflict experiments with migrating silvereyes at dusk

To assess the relative importance of celestial and magnetic cues for orientation at dusk, Australian silvereyes, Zosterops l. lateralis, were subjected to artificial magnetic fields under the natural evening sky, beginning 30 min before sunset. Control birds tested in the local geomagnetic field preferred their normal south-southwesterly migratory direction. Birds tested in a magnetic field with north deflected counterclockwise to 240°WSW showed northeasterly tendencies from the first test onward. Birds subjected to a corresponding clockwise deflection to 120°ESE, in contrast, first showed southerly directions, but from the 7th test onward shifted towards the northwest. Hence, both experimental groups followed the shift in magnetic north, one immediately, the other after a delay. When the birds were later tested in a vertical magnetic field without directional information, the two experimental groups continued in the direction they had preferred in the artificial magnetic fields, presumably by celestial cues alone. This indicates that they had not simply ignored celestial cues, but had recalibrated them according to the altered magnetic fields. The reasons for the initial difference between the two experimental groups remain unclear. Delayed responses to deflections of magnetic north have also been observed in previous studies. They appear to be the main reason why studies that expose birds only once to a cue-conflict situation often seem to indicate a dominance of celestial cues, whereas studies exposing the birds repeatedly usually indicate a dominance of magnetic cues. Accepted: 17 September 1997     

Disturbance cues as a source of risk assessment information under natural conditions

1. Disturbance cues are released by stressed or disturbed prey prior to a predator attack and convey useful risk assessment information regarding local threats. While studies have shown that disturbance cues may be important early on within the predation sequence (prior to an attack), their role in predator–prey interactions remains relatively overlooked by ecologists. Critically, experimental studies examining disturbance cues, especially among prey fishes, have been conducted primarily under laboratory or semi-natural conditions.
2. Here, we tested the prediction that disturbance cues function as sources of risk assessment information in situ. We exposed Trinidadian guppies, in two natural populations differing in predation risk, to a model predator paired with stream water or the disturbance cue collected from guppies from either a high- or low-predation risk population.
3. We found that the predator inspection response of guppies to disturbance cues depends on the level of risk of both the focal and the cue source population. Guppies from both populations exhibited increased latencies to inspect, lower inspection rates and reduced inspecting group sizes towards the model paired with conspecific disturbance cues versus a stream water control. Interestingly, guppies of both populations showed evidence of higher perceived predation risk towards the disturbance cues collected from high-predation risk donors compared to low-predation risk donors.
4. Our results support the hypothesis that disturbance cues function as a source of information used by prey fish in the assessment of predation risk and provide the first evidence of disturbance cue function under fully natural conditions.

     

Interaction between positional but not between non-positional cues in human predictive learning

Four experiments with human subjects examined the cue-interaction effects using a computer-controlled predictive learning task. In Phase 1, subjects learned that cue P was consistently associated with the occurrence of an outcome (P+), whereas cue N was never followed by the outcome (N−). In Phase 2, two neutral cues, R and I, were compounded with P and N, respectively. Each compound was followed by the outcome (PR+ and NI+). Thus, cue R was compounded with the already predictive cue P, whereas cue I was compounded with the non-predictive cue N. In each phase, subjects rated the contingency between the different cues and the outcome. In experiments 1 and 2, the spatial position of the cues was fixed, whereas it was variable in experiments 3, 4a and 4b. Verbal cues were used in experiments 1–3, whereas the cues consisted of geometrical figures in experiments 4a and 4b. Evidence for cue interaction, as indicated by giving cue I a higher contingency rating than cue R after or during Phase 2, was only found under the conditions of experiments 1 and 2. The results indicate that the use of positional cues facilitates the occurrence of cue-interaction effects. Possible reasons for this finding are discussed.     

Memory for spatial and object-specific cues in food-storing and non-storing birds

Two storer/non-storer pairs of species, marsh tit (Parus palustris)/blue tit (P. caeruleus) and jay (Garrulus glandarius)/jackdaw (Corvus monedula) were compared on a one-trial associative memory task. In phase I of a trial birds searched for a reward in one of four feeders which differed in their trial-unique spatial location and object-specific cues. Following a retention interval, the birds had to return to the same feeder to obtain a further reward. In control trials the array of feeders was unaltered, whilst in dissociation tests it was transformed to separate spatial location and object-specific cues.In control trials there was no difference in performance between species. In dissociation tests, the two storing species went first to the correct spatial location and second to the correct object-specific cues, whereas the two non-storing species went first with equal probability to the correct spatial and local object cues.Monocular occlusion was used to investigate differences between the two eye-systems. In control trials there was no effect of occlusion. In dissociation trials, all 4 species preferentially returned to the feeder with the correct object-specific cue when the left eye had been covered in phase I and to the feeder in the correct spatial position when the right eye had been covered in phase I.These results suggest that (a) food-storing birds differ from non-storers in responding preferentially to spatial information and (b) in storers and non-storers the right eye system shows a preference for object-specific cues and the left eye system for spatial cues.     

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.     

When to approach novel prey cues? Social learning strategies in frog-eating bats

Animals can use different sources of information when making decisions. Foraging animals often have access to both self-acquired and socially acquired information about prey. The fringe-lipped bat, Trachops cirrhosus, hunts frogs by approaching the calls that frogs produce to attract mates. We examined how the reliability of self-acquired prey cues affects social learning of novel prey cues. We trained bats to associate an artificial acoustic cue (mobile phone ringtone) with food rewards. Bats were assigned to treatments in which the trained cue was either an unreliable indicator of reward (rewarded 50% of the presentations) or a reliable indicator (rewarded 100% of the presentations), and they were exposed to a conspecific tutor foraging on a reliable (rewarded 100%) novel cue or to the novel cue with no tutor. Bats whose trained cue was unreliable and who had a tutor were significantly more likely to preferentially approach the novel cue when compared with bats whose trained cue was reliable, and to bats that had no tutor. Reliability of self-acquired prey cues therefore affects social learning of novel prey cues by frog-eating bats. Examining when animals use social information to learn about novel prey is key to understanding the social transmission of foraging innovations.     

The visual system of the honeybee (Apis mellifera): the maximum length of the orientation detector

Bees were trained to discriminate between two or more black bars and similar bars at right angles, presented on a vertical surface. The positions of the bars were shifted every 5 min to prevent their locations being used as cues. The experiments exploit the fact that bees do not discriminate the global orientation of a straight line of small black squares that are individually resolved, because the local responses to equal lengths of edges at right angles cancel out, and each square has no residual orientation cue. The experiments measure the resolution of this effect by control of the width of the gaps between the squares. At the limit the unit orientation detectors cannot span the gaps. Training with vertical or with horizontal bars in separate experiments, and testing with vertical or horizontal lines of squares, shows that the vertical gaps in horizontal rows are detected with better resolution than horizontal gaps in vertical rows. The results show that unit orientation detectors span not more than 3 ommatidia.     

Behavioural dissociation between exogenous and endogenous temporal orienting of attention

BACKGROUND: In the current study we compared the effects of temporal orienting of attention based on predictions carried by the intrinsic temporal structure of events (rhythm) and by instructive symbolic cues; and tested the degree of cognitive, strategic control that could be exerted over each type of temporal expectation. The experiments tested whether the distinction between exogenous and endogenous orienting made in spatial attention may extend to the temporal domain. TASK DESIGN AND MAIN RESULTS: In this task, a ball moved across the screen in discrete steps and disappeared temporarily under an occluding band. Participants were required to make a perceptual discrimination on the target upon its reappearance. The regularity of the speed (rhythmic cue) or colour (symbolic cue) of the moving stimulus could predict the exact time at which a target would reappear after a brief occlusion (valid trials) or provide no temporal information (neutral trials). The predictive nature of rhythmic and symbolic cues was manipulated factorially in a symmetrical and orthogonal fashion. To test for the effects of strategic control over temporal orienting based on rhythmic or symbolic cues, participants were instructed either to "attend-to-speed" (rhythm) or "attend-to-colour". Our results indicated that both rhythmic and symbolic (colour) cues speeded reaction times in an independent fashion. However, whilst the rhythmic cueing effects were impervious to instruction, the effects of symbolic cues were contingent on the instruction to attend to colour. FINAL CONCLUSIONS: Taken together, our results provide evidence for the existence of qualitatively separable types of temporal orienting of attention, akin to exogenous and endogenous mechanisms.