Eyelid squinting during food pecking in pigeons.

The visual control of pecking by pigeons (Columba livia) has latterly been thought to be restricted to the fixation stops interrupting their downward head movements because these stops prevent interference by motion blur. Pigeons were also assumed to close their eyes during the final head thrust of the peck. Here, we re-examined their pecking motions using high-speed video recordings and supplementary provisions that permitted a three-dimensional spatial analysis of the movement, including measurement of pupil diameter and eyelid slit width. The results confirm that pigeons do not close their eyes completely during the presumed optically ballistic phase of pecking. Instead, their eyelids are narrowed to a slit. The width of this slit is sensitive to both the ambient illumination level and the visual background against which seed targets have to be detected and grasped. There is also evidence of some interaction between pupil diameter and eyelid slit width. We surmise that besides being an eye-protecting reflex, the partial covering of the pupil with the eyelids may increase the depth of focus, enabling pigeons to obtain sharp retinal images of peck target items at very close range and during the beak-gape 'handling' of food items and occasional grit particles.

Systematic Analysis of Pigeons' Discrimination of Pixelated Stimuli: A Hierarchical Pattern Recognition System Is Not Identifiable.

Pigeons learned to discriminate two different patterns displayed with miniature light-emitting diode arrays. They were then tested with 84 interspersed, non-reinforced degraded pattern pairs. Choices ranged between 100% and 50% for one or other of the patterns. Stimuli consisting of few pixels yielded low choice scores whereas those consisting of many pixels yielded a broad range of scores. Those patterns with a high number of pixels coinciding with those of the rewarded training stimulus were preferred and those with a high number of pixels coinciding with the non-rewarded training pattern were avoided; a discrimination index based on this correlated 0.74 with the pattern choices. Pixels common to both training patterns had a minimal influence. A pixel-by-pixel analysis revealed that eight pixels of one pattern and six pixels of the other pattern played a prominent role in the pigeons' choices. These pixels were disposed in four and two clusters of neighbouring locations. A summary index calculated on this basis still only yielded a weak 0.73 correlation. The individual pigeons' data furthermore showed that these clusters were a mere averaging mirage. The pigeons' performance depends on deep learning in a midbrain-based multimillion synapse neuronal network. Pixelated visual patterns should be helpful when simulating perception of patterns with artificial networks.

Symmetry recognition by pigeons: Generalized or not?

This note looks into the reasons why earlier reports may have arrived at differing conclusions about pigeons' capacity to categorize bilaterally symmetric and asymmetric visual patterns. Attention is drawn to pigeons' comparatively superior visual flicker resolution and superior visual linear acuity by reporting results of two ad-hoc experiments. This circumstance turns out to constrain conclusions drawn by earlier symmetry-asymmetry studies that used computer-generated patterns displayed on cathode ray tube monitors as these suffered from pictorial distortions. Additionally one of the studies involved patterns of inconsistent symmetry at global and local levels. A smaller-scale experiment using slide-projected unequivocal symmetric and asymmetric patterns yielded results compatible with the supposition that pigeons are capable of a symmetry-asymmetry categorization. The possibility that an artfactual cue may have inadvertently accentuated this capability in an earlier own experiment is considered.

Sensitization to apomorphine in pigeons: a multifactorial conditioning process.

Apomorphine (apo), an unspecific direct dopamine agonist, elicits an intense and lasting pecking bout in pigeons. Apo yielded orderly dose-response functions, and repeated administrations led to sensitization. Strain and individual differences in sensitivity to apo were at least partly due to genetic factors. However, a strong cage-context dependency of the sensitization, which is indicative of conditioning, occurred in both pigeon strains studied. Apo-induced pecking and sensitization also occurred in total darkness. Pigeons could be conditioned to discriminate between an apo state and a non-apo state. A small dose of apo was effective as a conditioned stimulus when paired with a high dose as an unconditioned stimulus. The conditioned response (CR) was strongly specific to the context in which the sensitization to apo took place. The resistance to extinction of the CR could be increased through an oversensitization treatment. The incremental responses arising during the sensitization treatment and the CRs shown afterward by individual pigeons correlated significantly. The sensitization to apo in pigeons is well accounted for by a conditioning schema in which an interoceptive drug state is a conditional conditioned stimulus for the full expression of the incremental response. Variants of the scheme might also account for the sensitization of rodents to psychostimulants. A neural model that embodies the characteristics of the conditioning scheme has been proposed.

Sex differences in mental rotation strategy.

When humans decide whether two visual stimuli are identical or mirror images of each other and one of the stimuli is rotated with respect to the other, the time discrimination takes usually increases as a rectilinear function of the orientation disparity. On the average, males perform this mental rotation at a faster angular speed than females. This experiment required the rotation of both mirror-image-different and non-mirror-different stimuli. The polygonal stimuli were presented in either spatially unfiltered, high-pass or low-pass filtered versions. All stimulus conditions produced mental rotation-type effects but with graded curvilinear trends. Women rotated faster than men under all conditions, an infrequent outcome in mental rotation studies. Overall, women yielded more convexly curvilinear response functions than men. For both sexes the curvilinearity was more pronounced under the non-mirror-different, low-pass stimulus condition than under the mirror different, high-pass stimulus condition. The results are considered as supporting the occurrence of two different mental rotation strategies and as suggesting that the women were predisposed to use efficiently an analytic feature rotation strategy, while the men were predisposed to employ efficiently a holistic pattern rotation strategy. It is argued that the overall design of this experiment promoted the application of an analytic strategy and thus conferred an advantage to the female participants.

Learning processes in matching and oddity: the oddity preference effect and sample reinforcement.

Eight pigeons learned either matching (to sample) or oddity (from sample) with or without reward for sample responding. The training stimuli were coarse-white, fine-black, or smooth-mauve gravels in pots with buried grain as the reinforcer. Oddity without sample reward was learned most rapidly, followed by matching with sample reward, oddity with sample reward, and matching without sample reward. Transfer was related to acquisition rate: The oddity group without sample reward showed full (equal to baseline) color and texture transfer; the matching group with sample reward showed partial texture transfer; other groups showed no transfer. Sample reward was shown to determine rate of acquisition of matching and oddity and the oddity preference effect. The results are discussed in terms of item-specific associations operating early in learning prior to any relational learning between sample and comparison stimuli.

Repeated apomorphine administration alters dopamine D1 and D2 receptor densities in pigeon basal telencephalon.

When pigeons are repeatedly administered a dose of apomorphine they show an increasing behavioral response, much as rodents do. In birds this expresses itself in an augmented pecking response. This sensitization is assumed to be largely due to a conditioning process. Here we present evidence that sensitization is accompanied by an alteration of the D(1) to D(2) dopamine receptor densities. An experimental group of pigeons was repeatedly injected with apomorphine, and a control group with saline. The basal forebrain tissue, known to be rich in dopamine receptors, was subjected to binding assays using tritiated specific D(1) and D(2) dopamine receptor antagonists. There was a trend towards an increase in D(1) and a significant decrease in D(2) receptor densities in apomorphine-treated birds compared to the saline-treated controls. We conclude that extended apomorphine treatment modifies the D(1) dopamine receptor density in the opposite manner to the D(2) dopamine receptor density.

Behavioral sensitization to apomorphine in pigeons (Columba livia): blockade by the D1 dopamine antagonist SCH-23390.

Repeated administration of apomorphine leads to a context-dependent pecking response sensitization. Previously sensitized pigeons (Columba livia) challenged with saline in the same context show a conditioned response (CR). The authors studied the effects of intrastriatal injections of the dopamine (D(1)) antagonist SCH-23390 on both the sensitized response and the CR. When coadministered with apomorphine, SCH-23390 inhibited the initial response to apomorphine, prevented the development of sensitization, and impaired the maintenance of an already developed sensitization. However, SCH-23390 had no effect on the retrieval of a previously established CR. It is concluded that the activation of D(1) receptors in the caudal avian striatum is necessary for the acquisition and maintenance of the sensitization, but not for the expression, of the CR.

Sensitization to apomorphine, effects of dizocilpine NMDA receptor blockades.

The dopamine agonist apomorphine (apo) elicits bouts of stereotyped pecking in pigeons, a response which increases with successive apo injections. This sensitization is strongly context-specific and has been suggested to arise through a Pavlovian conditioning to both external and internal cues. We hypothetized that this learning involves dopamino-glutamatergic interactions and investigated the issue by inducing NMDA glutamate receptor blockades with the antagonist dizocilpine (diz). A first experiment examined the effects that four different doses (ranging between 0.05 and 0.12 mg/kg) of diz co-administered with a standard dose of 0.5 mg/kg of apo had on the development of the incremented response and on the later expression of the conditioned pecking response. Both responses were impaired by doses of around 0.10 mg/kg diz. A second experiment assessed whether either a diz treatment or a diz plus apo co-treatment affected the development of a subsequent sensitization to apo. The first treatment had no effect on the latter sensitization. A part sensitization that arose with the second treatment did not transfer to the final sensitization. The last experiment examined whether the administration of diz had an immediate effect on the incremented responding to apo and on the conditioned response shown by already sensitized pigeons. No effect was apparent with the first treatment, but there was a marked response inhibition with the second treatment. It is concluded that NMDA glutamate receptors play an important role in apo-induced sensitization in pigeons which is compatible with the Pavlovian conditioning account of sensitization.

Right or wrong, familiar or novel in pictorial list discrimination learning.

The interaction between nonassociative learning (presentation frequencies) and associative learning (reinforcement rates) in stimulus discrimination performance was investigated. Subjects were taught to discriminate lists of visual pattern pairs. When they chose the stimulus designated as right they were symbolically rewarded and when they chose the stimulus designated as wrong they were symbolically penalised. Subjects first learned one list and then another list. For a "right" group the pairs of the second list consisted of right stimuli from the first list and of novel wrong stimuli. For a "wrong" group it was the other way round. The right group transferred some discriminatory performance from the first to the second list while the control and wrong groups initially only performed near chance with the second list. When the first list involved wrong stimuli presented twice as frequently as right stimuli, the wrong group exhibited a better transfer than the right group. In a final experiment subjects learned lists which consisted of frequent right stimuli paired with scarce wrong stimuli and frequent wrong stimuli paired with scarce right stimuli. In later test trials these stimuli were shown in new combinations and additionally combined with novel stimuli. Subjects preferred to choose the most rewarded stimuli and to avoid the most penalised stimuli when the test pairs included at least one frequent stimulus. With scarce/scarce or scarce/novel stimulus combinations they performed less well or even chose randomly. A simple mathematical model that ascribes stimulus choices to a Cartesian combination of stimulus frequency and stimulus value succeeds in matching all these results with satisfactory precision.

Ontogeny has a phylogeny: background to adjunctive behaviors in pigeons and budgerigars.

Animals coping with operant conditioning tasks often show behaviors that are not recorded by keys, levers and similar response transducers. Nevertheless, these adjunctive behaviors should not be disposed of by classifying them as incidental. Often they are found to be at least partially influenced by the experimentally programmed contingencies, and under certain conditions they can in turn influence conditioned behaviors. Here we describe the occurrence and characteristics of two such behaviors, stimulus grasping in operantly key-pecking pigeons and intra-delay stereotypies in a delayed matching-to-sample task with budgerigars. It is argued that for a proper account of these behaviors it is necessary to refer to a behavioral systems approach that appeals to longer ranging ontogenetic and phylogenetic histories than is usually considered in the psychological literature. The gaping towards on-key stimuli by pigeons is attributed to the hypothesis that operantly conditioned key-pecks probably relate to a grasp-pecking response that is normally executed towards non-edible items covering food. The intra-delay behaviors shown by the budgerigars are assumed to have originated from stress-induced displacement responses that adventitiously came under the influence of differential reinforcement contingencies. Finally, we discuss what kinds of evidence are needed to put these hypothetical explanations on a more certain footing.

Behavioural consequences of nucleus accumbens dopaminergic stimulation and glutamatergic blocking in pigeons.

Upon systemic administration of apomorphine, a potent dopamine agonist, pigeons show a bout of pecking behaviour. When the drug is repeatedly administered a sensitization takes place that is associated with pronounced discrimination learning. Here we show that intra-cerebral injections of apomorphine in the periphery of the nucleus accumbens of pigeons also elicit pecking. We additionally show that injections of 5-amino-phosphonohepatnoic acid, a NMDA-glutamate receptor blocker, into the Acc impairs the performance of a learned visual discrimination incorporating pecking as a choice response. We conclude that, as it is the case in mammals, the control mechanisms of learned sensory-motor behaviour in birds involves dopaminergic and glutamatergic synaptic transmission within the nucleus accumbens area.

Conditioning the pecking motions of pigeons.

The spatio-temporal courses of head and neck motions of pigeons while pecking at small grains are described. Single and serial pecks are distinguished but the inter- and intraindividual variability of the peck kinetics is stressed. Pigeons were then trained with instrumental conditioning procedures to speed-up their pecking. A partial reinforcement schedule where pigeons had to peck repeatedly before receiving reward led to a mild shortening of inter-peck intervals at lower reinforcement rates but surprisingly, a lengthening at higher rates. A schedule where short inter-peck intervals were differentially rewarded yielded a pronounced abbreviation of the inter-peck intervals, but this was achieved by a reduction of the movement path rather than an increase in motion velocity. A schedule whereby increased approach velocities were differentially rewarded yielded marked movement accelerations. When pigeons were rewarded for diminished approach speeds they also showed significant movement decelerations. Finally, it is shown that pigeons could learn to reliably abort their peck approach movement when a visual stimulus signalling a penalty was occasionally presented during the approach movement. The proportion of successful peck interruptions decreased as these interruption signals occurred later during the approach phase. It is concluded that the pecking of pigeons is neither an innately fixed nor a visually ballistic movement. It is instead a multiply controlled and flexibly adaptable response pattern.

A new approach to the formation of equivalence classes in pigeons.

Four pigeons were given simultaneous discrimination training with visual patterns arbitrarily divided into two sets, with the stimuli in one set designated A1, B1, C1, and D1 and those in the other set designated A2, B2, C2, and D2. In sequentially introduced training phases, the pigeons were exposed to a series of reversals to establish AB and then CD equivalences. In subsequent testing sessions, a subset of stimuli from one set served as positive stimuli and those from the other set as negative stimuli on training trials, and transfer of the reinforced relation to other members of the sets was tested with nonreinforced probe trials. The pigeons were trained further on AC and BD equivalences and then were tested for the emergence of untrained AD and BC equivalences. Two of the 4 pigeons exhibited the emergence of one of these untrained equivalences, evidence for the emergence of transitive relations. This finding suggests that the pigeons established three-member functional equivalence classes by incorporating separately trained multiple equivalence relations. Repeated reversal training and probe testing enabled us to explore the formation and expansion of functional equivalence classes in pigeons.

Discrimination of shape reflections and shape orientations by Columba livia.

By using a free-operant instrumental discrimination procedure, it was demonstrated that pigeons find two-dimensional mirror-image visual forms more difficult to distinguish than otherwise similar forms. Variations in orientation of the discriminanda exacerbated the relative confusability of mirror images. No significant difference was found in the pigeons' performance whether the birds were discriminating vertically or horizontally reflected mirror-image pairs. Mirror images of shapes were also shown to be less discriminable than upside-down versions of shapes. The similarity of mirror-image patterns is discussed in relation to the generalized recognition of bilaterally symmetrical forms by pigeons. Pigeons found an orientation discrimination task involving a 45 degree tilt comparatively hard. A second experiment with a discrete-trial conditional paradigm confirmed that discriminations of shape orientations can be difficult for these birds. The addition of shape cues improved the performance on the orientation discrimination task, more so when arbitrary shapes were employed than when mirror images were used, which indicates again that the latter were more difficult to discriminate than the former. The relative insensitivity to shape orientations is ascribed to normal ecological demands on pigeons.