How special is sameness for pigeons and people?

Because of the importance of the sense of sameness for psychological science and because of the tenuous support for this notion in pigeons' matching-to-sample behavior, we experimentally explored the possibly special status of sameness for pigeons. Using photographs from three different natural categories (dogs, fish, and flowers) in a three-alternative matching-to-sample design, we obtained a reliable sameness advantage for pigeons only when the number of correct sample-comparison combinations could have contributed to a sameness advantage; otherwise, no sameness advantage emerged. However, human participants exhibited an immediate and dramatic sameness advantage under essentially the same training and testing conditions as had been given to pigeons. At least under these experimental circumstances, humans exhibit a sameness advantage that far eclipses that of pigeons.     

Do pigeons (Columba livia) perceive object unity?

Human infants perceive two rods moving in concert behind an occluder as one unitary rod. In four experiments we tested whether pigeons also perceive unity of objects. Pigeons were trained on a matching-to-sample task to discriminate between one unitary rod moving at a constant speed and two aligned rods moving together at the same speed. The latter stimulus was identical to the former except for a gap in the center. In experiment 1, we tested pigeons in probe trials in which a rectangle occluded the center of the sample rods, to see which comparison stimulus, the unitary rod or the aligned two rods, the subjects would match to the sample. Two of the three subjects pecked at the two rods significantly more often than at the unitary rod. In experiment 2, we trained the same pigeons to match the sample rods moving "in front of" the occluder. Pigeons persisted in matching two separate rods to the unitary rod moving in front of the occluder. In experiments 3 and 4, we used a parallelogram and an undulating shape as the occluder to alter the shape and the size of the portions above and below the occluder by the motion of the sample rods. Both subjects chose the two rods significantly more often than chance in experiment 3 and one of them did so in experiment 4. The results suggest that pigeons do not complete occluded portions even though the two elements move in concert. These negative results suggest that some alternative way of identifying objects may have evolved in pigeons. Accepted after revision: 2 May 2001 Electronic Publication     

Responding to a positive stimulus by “satiated” pigeons

Skinner (1938) found that rats given discrimination training (Phase I) and then reinforced to “satiation” for responses in the presence of the negative stimulus (S?) (Phase II), began to respond again when the positive stimulus (S+) was reintroduced (Phase III). Experiment I replicated Skinner's finding with pigeons, alternating S+ and S? presentations during Phase III. In Experiment II, Phase II was extended, and Phase III results were similar to those of Experiment I, demonstrating that the recovery of S+ responding could not be attributed to a lax Phase II satiation criterion. In Experiment III, a uniform schedule of reinforcement was maintained throughout the three phases, and results similar to those of Experiment I were found, indicating that renewed S+ responding was not due to the shift in schedule between phases. In Experiment IV, Phase I consisted of discrimination training with two positive stimuli (S+_s and S+_n), and Phase II consisted of reinforcement for responses in the presence of S? and S+_s. During Phase III, significantly more responding was found to S+_s and S+_n than to S?, but no difference in responding was found between the two positive stimuli. In Experiment V, Phase I consisted of simple discrimination training, and during Phase II, responses in the presence of both S? and a novel stimulus (So) were reinforced. During Phase III, significantly more responding was found to S+ than to either S? or So, with no difference found between S? and So responding. Renewed responding to S+ during Phase III in the present experiments is best explained by behavioral contrast developed during Phase I.     

Technologies to reliably transduce the topographical details of pigeons’ pecks

Pigeons’ pecking has long been a subject of interest in behavioral research, with the response typically being viewed as unitary. Recent experiments done with computer-controlled devices have revealed that this response is at least bipartite in character, with beak opening and response location (head transport) as its components. In addition, experimental work has demonstrated that these response components may be separately influenced and controlled by respondent and operant conditioning procedures. The detailed topographic analysis and technology that have emerged may provide a background for similar work with other behavioral systems.     

Does the use of natural stimuli facilitate amodal completion in pigeons?

Three experiments were carried out to investigate whether amodal completion in pigeons can be facilitated by the use of colour photographs instead of highly artificial stimuli such as geometrical shapes. Ten pigeons were trained in a go/no-go procedure to discriminate between photographs of complete and of incomplete pigeon figures. In the subsequent test, the birds classified pictures of partly occluded pigeons as though they were complete (experiment 1). However, we found evidence that classification was based on spurious stimulus features that paralleled the intended class rule of figural completeness versus incompleteness. In particular, classification was shown to be guided by white background gaps that separated the parts of the fragmented pigeon figures (experiment 2), as well as by cues related to overall Gestalt (experiment 3). In summary, the present results indicate that the use of more natural stimuli such as photographs instead of geometrical shapes is insufficient for providing amodal completion in pigeons. It is suggested that a combination of various cues, including, eg, 3-D information and common motion in addition to surface and contour properties, may be required to induce a perceptual bias favouring visual completion of occluded portions.     

Memorization and “feature selection” in the acquisition of natural concepts in pigeons

Pigeons were trained on a successive discrimination task using complex visual stimuli. In Experiment 1, each photographic slide that contained a person had a corresponding “matched background” slide, one that showed the same scene with the person removed. Birds trained on a human positive discrimination acquired the matched pairs problem, but birds trained on a human negative discrimination performed poorly. This suggests a feature-positive effect for complex stimulus categories. Memorization control groups that were trained on a human-irrelevant discrimination also performed poorly with matched slides. However, subsequent experiments demonstrated that these effects depended on the use of matched pairs of slides. The human-as-feature effect was not obtained when human positive and human negative groups were subsequently trained with non-matched slides (Experiment 2), and memorization control groups acquired a human-irrelevant discrimination when trained with nonmatched slides (Experiment 3). Additional tests conducted in Experiments 2 and 3 found that performance was not disrupted when either the reinforced or nonreinforced slides were replaced. This effect was obtained when the category was relevant to the discrimination (Experiment 2) and when the category was irrelevant to the discrimination (Experiment 3). Finally, Experiment 4 demonstrated that memorization of a set of slides is possible when slides are sufficiently dissimilar, (i.e., nonmatched) but performance is not as good when the category exemplars are irrelevant to the discrimination.     

Transitive inference by pigeons: Does the geometric presentation of the stimuli make a difference?

In studies of transitive inference (TI), nonhuman animals are typically trained with the following 5-term task: A+B?, B+C?, C+D?, D+E? where the letters stand for arbitrary stimuli and [+] indicates that choice is reinforced and [?] indicates that choice is not reinforced. A TI effect is found when, given the untrained test pair BD, subjects choose B. TI effects have been found in many nonhuman species. Although reinforcement history has been posited as an account of the TI effect, it has failed to account for a variety of conditions under which TI effects have been found. A more cognitive account of TI is that organisms are able to form a representation of the series (A > B > C > D > E). In support of this hypothesis, Roberts and Phelps (Psychol Sci 5:368–374, 1994) found that presentation of the pairs of stimuli in a linear arrangement facilitated TI performance by rats, whereas presentation of the pairs of stimuli in a circular arrangement did not. Using methods adapted from Roberts and Phelps, we trained pigeons on either a linear or a circular arrangement of stimuli with the 5-term task. Results indicated that on the BD test pair, pigeons trained with a circular arrangement did not differ from those trained with a linear arrangement. Furthermore, we found that memory for training pairs was variable and was highly correlated with degree of TI. The results suggest that regardless of how pigeons are able to represent the stimuli, choice was not affected by the spatial arrangement of the stimuli during training.     

“Insight” in pigeons: absence of means–end processing in displacement tests

The understanding of functional relations between action and consequence is a critical component of intelligence. To examine this linkage in pigeons, we investigated their understanding of the relations of the elements tested in an extension of Köhler’s box stacking task to this species. In the experiments, the pigeons had to move a spatially displaced box under an out-of-reach target. Experiment 1 successfully replicated and extended the previous finding showing that when separately trained to move a box and stand on it to peck the target, pigeons can synthesize these behaviors to solve the single-box displacement problem quickly on their first attempt. Experiment 2 tested whether pigeons, when given a simultaneous choice between two boxes with identical reinforcement histories, would selectively choose the box with the correct functional affordance (i.e., permitting standing) to solve the problem rather than a non-functional one. Their extensive, equivalent, and undirected behavior in moving both boxes during these tests suggests the pigeons did not possess a means–end understanding of the functional properties of the boxes. Instead, their results were consistent with an analysis of their earlier synthetic behavior as being due to the temporal and spatial relations of the physical elements in the task and their prior learned behaviors.     

A method for automatically recording topographical differences in pigeons’ keypecking for food and water reinforcers

The topography of pigeons’ keypecking responses for food reinforcers is visibly different from the keypeck topography for water reinforcers. Previous methods of recording topographies have been complex and not without problems. A simple inexpensive modification of a standard pecking key permits the recording of two components of the topography in sufficient detail to allow the detection of these topographical differences.     

Do birds (pigeons and bantams) know how confident they are of their perceptual decisions?

Rhesus monkeys are known to recognize confidence about their immediate perceptual and cognitive decisions by using a betting procedure (Son and Kornell in The missing link in cognition: origins of self-reflective consciousness. Oxford University Press, New York, pp 296–320, 2005; Kornell et al. in Psychol Sci 18:64–71, 2007). In this report, we examined whether this ability is shared in two avian species (pigeons and bantams) in order to know how widespread this metacognitive ability is among animals. We trained pigeons and bantams to search for a differently colored disk (target) among others (distracters) displayed on a touch-sensitive monitor. In test, the subjects were required to choose one of two confidence icons, “risk” and “safe”, after the visual search. A peck at the “risk” icon after a correct response in the visual search (i.e., a peck at the target) was reinforced by food and light, while that after an incorrect response (i.e., a peck at a distracter) resulted in a timeout. A peck at the “safe” icon was always reinforced by food and light, or by light only, regardless of the visual search result. The percentages of “safe” choices after incorrect responses were higher than after correct responses in all six pigeons and two of three bantams. This behavior generalized to novel stimuli in some subjects, and even to a novel line-classification task in a pigeon. These results suggest that these two distantly related avian species have in common a metacognitive ability that allows them to recognize confidence about their immediate perceptual decisions.     

A relational differential outcomes effect: pigeons can classify outcomes as “good” and “better”

In a conditional discrimination each of two sample stimuli indicates which of two comparison stimuli is correct. When correct choice following each conditional stimulus is followed by a different outcome (one kind of food following one, a different kind of food following the other) it often facilitates acquisition and improves memory. In transfer designs, in which two different conditional discriminations are followed by the same two differential outcomes, outcome expectation can be shown to be sufficient for comparison choice. That is, the samples from one conditional discrimination are matched to comparisons from the other conditional discrimination based on the common outcomes alone. In the present study we asked if for pigeons the relative value of the differential outcomes (higher versus lower value) can serve as the basis for comparison choice, independent of other characteristics of the outcomes and of differential sample responding. That is, would different outcomes that could be described as “good” and “better” form two stimulus classes. For one conditional discrimination, the differential outcomes involved differential probability of reinforcement for choice of the correct comparison stimulus (0.80 vs. 0.20 for correct choice of the two comparisons, respectively). For the other conditional discrimination, the differential outcomes involved differential responding to the two comparison stimuli (5 pecks vs. 20 pecks to the correct comparisons, respectively). On test trials, when conditional stimuli from the two conditional discriminations were interchanged and the relative value of the differential outcomes could serve as the only basis for comparison choice, we found positive transfer. The results indicate that relational attributes of outcomes can serve as effective cues for comparison choice.     

Perception of the standard and the reversed Müller-Lyer figures in pigeons (Columba livia) and humans (Homo sapiens)

The authors compared perception of the standard and reversed Müller-Lyer figures between pigeons (Columbia livia) and humans (Homo sapiens). In Experiment 1, pigeons learned to classify 6 lengths of target lines into "long" and "short" categories by pecking 2 keys on the monitor, ignoring the 2 brackets so placed that they would not induce an illusion. In the test that followed, all 3 birds chose the "long" key more frequently for the standard Müller-Lyer figures with inward-pointing brackets (><) than for the figures with outward-pointing brackets (<>). The subjects' responses were accountable by neither overall lengths of the figures nor horizontal gaps between the 2 brackets. For the reversed figures, effects of the brackets were absent. These results suggested that the pigeons perceived the standard Müller-Lyer illusion but not the reversed one. Experiment 2 confirmed that humans perceived both types of the illusion. Pigeons and humans may perceive the same illusory figures in different ways.     

Failure of acute and chronic administration of Δ9-tetrahydrocannabinol to affect the repeated acquisition of serial position responses in pigeons

Pigeons were trained to acquire a new four-response position sequence each day by pecking three response keys in a predetermined order. The key color varied after each correct response prior to food delivery. Acute administration of Γ⁹-tetrahydrocannabinol (Δ⁹-THC) up to a dose that completely eliminated responding, had no effect on total acquisition errors, or on within session patterns of error elimination. Chronic administration of Δ⁹-THC (3–10 mg/kg/day), either before or after the session for 4–7 weeks, also did not affect these error measures, although rates of responding were markedly suppressed and at times no responding occurred Discontinuation of Δ⁹-THC administration for periods of 4–6 weeks also was without effect on errors. These experiments suggest that neither acute nor chronic Δ⁹-THC produce specific effects on the repeated acquisition of serial position responses in pigeons.     

“work ethic” in pigeons: Reward value is directly related to the effort or time required to obtain the reward

Stimuli associated with less effort or with shorter delays to reinforcement are generally preferred over those associated with greater effort or longer delays to reinforcement. However, the opposite appears to be true of stimuli thatfollow greater effort or longer delays. In training, a simple simultaneous discrimination followed a single peck to an initial stimulus (S+_FR1 S−_FR1) and a different simple simultaneous discrimination followed 20 pecks to the initial stimulus (S+_FR20 S−_FR20). On test trials, pigeons preferred S+_FR20 over S+_FR1 and S−_FR20 over S−_FR1. These data support the view that the state of the animal immediately prior to presentation of the discrimination affects the value of the reinforcement that follows it. This contrast effect is analogous to effects that when they occur in humans have been attributed to more complex cognitive and social factors.     

Do ‘literate’ pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>) show mirror-word generalization?

Many children pass through a mirror stage in reading, where they write individual letters or digits in mirror and find it difficult to correctly utilize letters that are mirror images of one another (e.g., b and d). This phenomenon is thought to reflect the fact that the brain does not naturally discriminate left from right. Indeed, it has been argued that reading acquisition involves the inhibition of this default process. In the current study, we tested the ability of literate pigeons, which had learned to discriminate between 30 and 62 words from 7832 nonwords, to discriminate between words and their mirror counterparts. Subjects were sensitive to the left–right orientation of the individual letters, but not the order of letters within a word. This finding may reflect the fact that, in the absence of human-unique top-down processes, the inhibition of mirror generalization may be limited.     

Reorientation in a two-dimensional environment: II. Do pigeons (Columba livia) encode the featural and geometric properties of a two-dimensional schematic of a room?

Pigeons (Columba livia) searched for a hidden target area in images showing a schematic rectangular environment. The absolute position of the goal varied across trials but was constant relative to distinctive featural cues and geometric properties of the environment. Pigeons learned to use both of these properties to locate the goal. Transformation tests showed that pigeons could use either the color or shape of the features, but performance was better with color cues present. Pigeons could also use a single featural cue at an incorrect corner to distinguish between the correct corner and the geometrically equivalent corner; this indicates that they did not simply use the feature at the correct corner as a beacon. Interestingly, pigeons that were trained with features spontaneously encoded geometry. The encoded geometric information withstood vertical translations but not orientation transformations.     

A visual familiarity account of evidence for orthographic processing in pigeons <Emphasis Type="Italic">(Columbia livia)</Emphasis>: a reply to Scarf,Corballis, Güntürkün,and Colombo (2017)

Scarf et al. (Proc Natl Acad Sci 113(40):11272–11276, 2016) demonstrated that pigeons, as with baboons (Grainger et al. in Science 336(6078):245–248, 2012; Ziegler in Psychol Sci.  https://doi.org/10.1177/0956797612474322, 2013), can be trained to display several behavioural hallmarks of human orthographic processing. But, Vokey and Jamieson (Psychol Sci 25(4):991–996, 2014) demonstrated that a standard, autoassociative neural network model of memory applied to pixel maps of the words and nonwords reproduces all of those results. In a subsequent report, Scarf et al. (Anim Cognit 20(5):999–1002, 2017) demonstrated that pigeons can reproduce one more marker of human orthographic processing: the ability to discriminate visually presented four-letter words from their mirror-reversed counterparts (e.g. “LEFT” vs. “ Open image in new window ”). The current report shows that the model of Vokey and Jamieson (2014) reproduces the results of Scarf et al. (2017) and reinforces the original argument: the recent results thought to support a conclusion of orthographic processing in pigeons and baboons are consistent with but do not force that conclusion.