Motion as a natural category for pigeons: Generalization and a feature-positive effect

Three groups of pigeons were trained with a modified discriminative autoshaping procedure to discriminate video images of other pigeons on the basis of movement. Birds of all groups were shown the same video images of other pigeons, which were either moving or still. The group to whom food was presented only after moving images learned the discrimination very quickly. A second group, to whom food was given only after still images, and a pseudocategory group, to whom food was presented after arbitrarily chosen stimuli, showed no evidence of discrimination during acquisition training. Extinction conditions led to clear differences in peck rates to moving and still images in the second group but not in the pseudocategory group. The result is related to the feature-positive effect. Generalization tests showed that the discrimination performance was based on visual features of the stimuli but was invariant against changes of size, perspective, brightness, and color. Furthermore, discrimination was maintained when novel images of pigeons under different viewing angles and seven other types of motion categories were presented. It is argued that the discrimination is based not on a common motion feature but on motion concepts or high-order generalization across motion categories.     

Drug discrimination under a concurrent fixed-ratio fixed-ratio schedule.

Pigeons were trained to discriminate 5.0 mg/kg pentobarbital from saline under a two-key concurrent fixed-ratio 10 fixed-ratio 40 schedule of food presentation, in which the fixed-ratio component with the lower response requirement was programmed to reinforce responding on one key after drug administration (pentobarbital-biased key) and on the other key after saline administration (saline-biased key). After responding stabilized, pigeons averaged 98% of their responses on the pentobarbital-biased key during training sessions preceded by pentobarbital, and they averaged 90% of their responses on the saline-biased key during training sessions preceded by saline. In test sessions preceded by doses of pentobarbital, chlordiazepoxide, or ethanol, pigeons switched from responding on the saline-biased key at low doses to responding on the pentobarbital-biased key at higher doses (the dose-response curve was quantal). High doses of phencyclidine produced responding on both keys, whereas pigeons responded almost exclusively on the saline-biased key after all doses of methamphetamine. These and previous experiments using concurrent reinforcement schedules to study drug discrimination illustrate that the schedule of reinforcement is an important determinant of the shape of dose-effect curves in drug-discrimination experiments.     

Discrimination of methadone and cocaine by pigeons without explicit discrimination training.

Pigeons were trained to peck a key on a variable-interval 2-min schedule of food reinforcement. Prior to each session, either 2.0 mg/kg methadone (n = 3), 3.0 mg/kg cocaine (n = 4), or 5.6 mg/kg cocaine (n = 2) was administered. When each pigeon's rate of pecking was stable, a range of doses of the training drug and saline were administered prior to 20-min extinction sessions separated by at least four training sessions. Rate of pecking during these extinction tests was generally an increasing function of dose, with the lowest rates obtained following saline and low doses and the highest rates obtained following doses near the training doses. Dose functions from pigeons trained with 5.6 mg/kg cocaine were steeper than those from pigeons trained with 3.0 mg/kg cocaine. Pigeons trained with methadone or 3.0 mg/kg cocaine were then given discrimination training, in which food reinforcement followed drug administration and 20-min extinction sessions followed saline administration. Rates of pecking under these conditions quickly diverged until near-zero rates were obtained following saline and high rates were obtained following drug. Discrimination training steepened dose functions for the training drugs, and the effects of several other substituted drugs depended on the pharmacology of the training drug. The pigeons trained with 5.6 mg/kg cocaine were tested with d-amphetamine, methadone, and morphine prior to discrimination training. d-Amphetamine increased rates dose dependently, and methadone and morphine did not. The results suggest that discriminative control by methadone and cocaine was established without explicit discrimination training.     

Discrimination of moving video images of self by pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>)

The ability to recognize self has been known to be limited to some animal species, but previous research has focused almost exclusively on the animal's reaction to a mirror. Recent studies suggest that the temporal contingency between a subject's action and the corresponding visual scene reflected in a mirror plays an important role in self-recognition. To assess the roles of visual-proprioceptive contiguity in self-recognition, we explored whether pigeons are able to discriminate videos of themselves with various temporal properties. We trained five pigeons to respond to live video images of themselves (live self-movies) and not to video filmed during previous training sessions (pre-recorded self-movies). Pigeons learned to peck trial-unique live self-movies more frequently than pre-recorded self-movies. We conducted two generalization tests after pigeons learned to discriminate between the two conditions. First, discrimination acquired during training sessions was transferred to a test session involving live self-movies and new pre-recorded self-movies. Second, the same pigeons were tested in extinction procedure using delayed live self-movies and new pre-recorded self-movies. Although pigeons responded to delayed presentations of live self-movies more frequently than to new pre-recorded self-movies, the relative response rate to delayed presentation of live self-movies gradually decreased as the temporal discrepancy between pigeons' own behavior and the corresponding video increased. These results indicate that pigeons' discrimination of self-movies with various temporal properties was based on the temporal contiguity between their behavior and its visual feedback. The methodology used in the present experiment is an important step toward improving the experimental analysis of self-recognition in non-human animals.     

Choice as a dependent measure in autoshaping: sensitivity to frequency and duration of food presentation.

Previous investigations have shown that rate, latency, and percentage of trials with at least one response are somewhat insensitive measures of the strength of autoshaped responding. In the present studies, these measures were contrasted with the allocation of responding during simultaneous choice tests, a measure of response strength frequently used in operant paradigms. In two experiments, nine pigeons were exposed to a forward pairing autoshaping procedure. Training sessions consisted of the successive presentation of three stimuli, each followed by food on either 100%, 50%, or 0% of the trials. Choice testing involved the simultaneous presentation of the three stimuli. In Experiment I, all pigeons consistently directed their initial choice responses and the majority of subsequent responses to the stimulus always followed by food, despite the fact that during training sessions the response rates of most birds were highest in the presence of the stimulus followed by food on 50% of the trials. In Experiment II, rate, latency, and percentage of trials with at least one response did not change appreciably as a function of duration of feeder presentations. However, choice responding was lawfully affected by duration of feeder presentations. These data suggest that choice is perhaps a more sensitive measure of the strength of autoshaped responding than other, more commonly employed, indices.     

Acquisition of a same-different discrimination task by pigeons (Columba livia)

Four homing pigeons were trained to discriminate two figures simultaneously presented on an LCD screen. The figure was either a rectangle (A) or a square (B), and four combinations of the two figures, AA, AB, BA, BB, appeared in a pseudo-randomized order. The pigeons' task was to peck one of these figures based upon whether the two figures were identical or not. One pigeon successfully learned this discrimination, with proportions of correct responses above 90% in two consecutive sessions. Of the other birds, two performed above chance level but had difficulty meeting a learning criterion of above 80% in two consecutive sessions. All birds achieved this criterion when the combinations of figures presented were reduced to two. Results suggested that learning the present same-different discrimination is within the capacity of pigeons to a certain extent, although there exists considerable individual variation in the pigeons' skills to acquire complex discrimination.     

Effects of amphetamine-CNS depressant combinations and of other CNS stimulants in four-choice drug discriminations

Three pigeons were trained to discriminate among 5 mg/kg pentobarbital, 2 mg/kg amphetamine, a combination of these two drugs at these doses, and saline using a four-choice procedure (amphetamine-pentobarbital group). Three other pigeons were trained to discriminate among 5 mg/kg morphine, 2 mg/kg methamphetamine, a combination of these two drugs at these doses, and saline (methamphetamine-morphine group). After 10 to 13 months of training, the pigeons averaged more than 90% of their responses on the appropriate key during training sessions. In subsequent testing, dose-response curves were determined for the individual drugs, for a wide range of dose combinations of the training drugs, and for two drugs to which the pigeons had not been exposed previously (pseudoephedrine and nicotine). After low test doses of the training drugs, pigeons responded on the saline key. As the dose increased, responding on the key associated with that drug during training sessions increased. When training drugs were combined at doses that were not discriminable when given alone, responding occurred on the saline key. When a discriminable dose of one training drug was combined with a nondiscriminable dose of the other training drug, responding occurred on the key associated with the discriminable dose. When both drugs were given at discriminable doses, responding almost always occurred on the drug-combination key. The response-rate decreasing effects of pentobarbital and amphetamine were mutually antagonized when the drugs were combined, but the rate-decreasing effects of morphine and methamphetamine were not. After low doses of pseudoephedrine and nicotine, pigeons in both groups responded on the saline key. After higher doses of pseudoephedrine and nicotine, responding in the amphetamine-pentobarbital group occurred primarily on the amphetamine key. In the methamphetamine-morphine group, higher doses of pseudoephedrine and especially nicotine engendered more responding on the combination key than had occurred in the other group. The four-choice procedure can reveal subtle effects in the discrimination of individual drugs and drug combinations that are not apparent with procedures offering fewer response alternatives.     

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.     

An analysis of confusion errors in many-to-one matching with temporal and nontemporal samples

In experiment 1, pigeons were trained to match temporal (2, 8, and 10 s of houselight) and location (feeder light, left key, center key illumination) samples to color comparison stimuli. Red choices were correct following the 2-s and feeder light, orange choices were correct following the 8-s and center key, and green choices were correct following the 10-s and left key. Samples that were harder to discriminate (8- vs 10-s, and left vs center key) were mapped onto comparisons that were easy to discriminate (orange vs green), while samples that were easier to discriminate (2- vs 8-s, and feeder light vs left key) were mapped onto comparisons that were hard to discriminate(red vs orange). The pattern of errors for temporal and location samples indicated that these samples were not represented by a common code even though they were associated with the same comparison stimuli. In experiment 2, the same pigeons were trained with visual samples in which samples that were hard to discriminate (triangle vs circle) were mapped onto comparisons that were easy to discriminate (orange vs green), while samples that were easy to discriminate(plus vs triangle) were mapped onto comparisons that were hard to discriminate (red vs orange). Following acquisition of the visual discrimination, the temporal samples were re-introduced and many-to-one training was continued. During delay testing, the pattern of errors for temporal and visual samples was equivalent and consistent with the hypothesis that visual samples were being coded in terms of the duration appropriate for the temporal sample with which it shared a common comparison response. Data from no-sample test sessions ruled out a simple response bias explanation of the data. The properties of common codes for temporal and nontemporal events can be somewhat flexible and more complicated than previously envisaged. Received: 21 January 1998 / Accepted after revision: 5 April 1998     

Discrimination of intentional and random motion paths by pigeons

Twelve pigeons (Columba livia) were trained on a go/no-go schedule to discriminate between two kinds of movement patterns of dots, which to human observers appear to be "intentional" and "non-intentional" movements. In experiment 1, the intentional motion stimulus contained one dot (a "wolf") that moved systematically towards another dot as though stalking it, and three distractors ("sheep"). The non-intentional motion stimulus consisted of four distractors but no stalker. Birds showed some improvement of discrimination as the sessions progressed, but high levels of discrimination were not reached. In experiment 2, the same birds were tested with different stimuli. The same parameters were used but the number of intentionally moving dots in the intentional motion stimulus was altered, so that three wolves stalked one sheep. Despite the enhanced difference of movement patterns, the birds did not show any further improvement in discrimination. However, birds for which the non-intentional stimulus was associated with reward showed a decline in discrimination. These results indicated that pigeons can discriminate between stimuli that do and do not contain an element that human observer see as moving intentionally. However, as no feature-positive effect was found in experiment 1, it is assumed that pigeons did not perceive or discriminate these stimuli on the basis that the intentional stimuli contained a feature that the non-intentional stimuli lacked, though the convergence seen in experiment 2 may have been an effective feature for the pigeons. Pigeons seem to be able to recognise some form of multiple simultaneously goal-directed motions, compared to random motions, as a distinctive feature, but do not seem to use simple "intentional" motion paths of two geometrical figures, embedded in random motions, as a feature whose presence or absence differentiates motion displays. Electronic Publication     

A pellet feeder for the birds

An alternative means of delivering food to pigeons in operant conditioning research is described. The feeder allows greater control of the amount of food delivered and reduces the amount of time necessary for the pigeon to collect the food. It is possible to extend the length of experimental sessions due to the reduction of food intake. Data obtained using the pellet feeder indicated that the control of responding is comparable to that observed with the standard grain-magazine feeder.     

Some effects of punishment shock intensity upon discriminative responding

Three pigeons received visual discrimination training under both multiple variable-ratio extinction and variable-interval extinction schedules. All birds developed nearly perfect discrimination. When punishment for every tenth response during food reinforcement was presented, responding decreased as shock intensity increased. At the same time, responding during extinction, which was not punished, increased at intermediate punishment intensities, but returned to low levels under severe punishment. A second procedure, in which punishment and no-punishment sessions alternated unsystematically, was employed with two of the birds. The results under this procedure essentially replicated the data obtained as punishment shock intensity increased gradually.     

Effects of search cost on foraging and feeding: a three-component chain analysis

An experiment determined whether pigeons minimize number of key pecks per food delivery and maintain their baseline intake of food while key pecking on a three-component chain schedule. Pigeons at either 80% or 100% body weight obtained all their food during baseline and contingency sessions. During baseline sessions, pecks on the left and center keys had no consequences; each peck on the right key activated the feeder. During contingency sessions, pigeons key pecked on a three-component chain schedule simulating components of a foraging chain. In the search component either 3, 9 or 15 key pecks (varied parametrically across blocks of sessions) on the left key produced a stimulus on the middle key, indicating an encounter with either the low-cost prey (3 key pecks) or an equally probable high-cost prey (21 key pecks). In the procurement component the pigeon pecked either: (a) the left key once, thus returning to the search component, or (b) the middle key either 3 or 21 times, which activated the right response key. In the handling component one peck on the right key operated the feeder. The pigeons always procured the low-cost prey and minimized the number of key pecks per hopper by procuring the high-cost prey when the search-cost ratio was high (15 key pecks) but not when it was low (3 key pecks). All pigeons maintained their baselines of eating during contingency sessions by key pecking more frequently and eating more efficiently. The 80% body-weight birds produced higher overall rates of key pecking and eating. These results have implications for ecological theories of optimal foraging and for psychological theories of learned performance.     

Applying bubbles to localize features that control pigeons' visual discrimination behavior

The authors trained pigeons to discriminate images of human faces that displayed: (a) a happy or a neutral expression or (b) a man or a woman. After training the pigeons, the authors used a new procedure called Bubbles to pinpoint the features of the faces that were used to make these discriminations. Bubbles revealed that the features used to discriminate happy from neutral faces were different from those used to discriminate male from female faces. Furthermore, the features that pigeons used to make each of these discriminations overlapped those used by human observers in a companion study (F. Gosselin & P.G. Schyns, 2001). These results show that the Bubbles technique can be effectively applied to nonhuman animals to isolate the functional features of complex visual stimuli.     

Airflow as a discriminative stimulus

In one experiment, pigeons were taught to discriminate airflow by having availability of reinforcement signalled by its presence and extinction signalled by its absence. After they reached criterion, some were trained on a discrimination reversal. Others were trained on an intradimensional discrimination with a low airflow velocity associated with reinforcement and a higher airflow velocity associated with extinction. All discriminations were learned rapidly, indicating that airflow velocity can function as a discriminative stimulus. In the second and third experiments, naive pigeons were trained to discriminate the presence of a compound stimulus (one of three tonal intensities paired with one of three airflow velocities) from its absence. These pigeons were subsequently given a component stimulus test during extinction on four stimulus values; the two training values, the tone alone, and the airflow alone. High or moderate velocity airflow controlled more responding than any of the three tone intensities. However, low velocity airflow controlled more responding only when a low intensity tone was employed.     

Pigeon concept formation: successive and simultaneous acquisition

Pigeons were trained to discriminate the presence of one or more human forms in displays projected on a panel above the response key. This task was mastered, although imperfectly, with successive and with simultaneous presentations of positive and negative instances. The course of acquisition of the discrimination was similar for the two training procedures. Animals were able to transfer the discrimination from the successive to the simultaneous situation. Various tests were carried out to control for artifactual cues on which the discrimination might have been based. The discrimination was maintained when new displays were presented, when reinforcement was omitted, and when displays were inverted 180°. Animals were also able to discriminate between pairs of displays that were identical, except that one member of the pair contained a human form. The research extends the techniques used by Herrnstein and Loveland (1964), and confirms their finding that pigeons can master the concept of “person-present” in a visual display.     

Pigeons concurrently categorize photographs at both basic and superordinate levels

We studied categorization in pigeons, using carefully controlled photographs. Within daily sessions, 4 pigeons had to classify each of 32 photographs into either its proper basic-level category (cars, chairs, flowers, or people; four-key forced choice procedure) or its proper superordinate-level category (natural or artificial; two-key forced choice procedure). The pigeons successfully classified the same stimuli at both levels. Overall, the pigeons learned the basic discrimination more quickly than the superordinate discrimination, but this difference was reliable only for artificial stimuli (cars and chairs), not for natural stimuli (flowers and people). The pigeons also exhibited reliable discrimination transfer to novel photographs, attesting to the open-endedness of these basic and superordinate categories.     

Time-space learning in homing pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>): orientation to an artificial light source

Time-space learning reflects an ability to represent in memory event-stimulus properties together with the place and time of the event; a capacity well developed in birds. Homing pigeons were trained in an indoor octagonal arena to locate one food goal in the morning and a different food goal in the late afternoon. The goals differed with respect to their angular/directional relationship to an artificial light source located outside the arena. Further, the angular difference in reward position approximated the displacement of the sun's azimuth that would occur during the same time period. The experimental birds quickly learned the task, demonstrating the apparent ease with which birds can adopt an artificial light source to discriminate among alternative spatial responses at different times of the day. However, a novel midday probe session following successful learning revealed that the light source was interpreted as a stable landmark and not as a surrogate sun that would support compass orientation. Probe sessions following a phase shift of the light–dark cycle revealed that the mechanism employed to make the temporal discrimination was prevailingly based on an endogenous circadian rhythm and not an interval timing mechanism.     

A new learning paradigm elicits fast visual discrimination in pigeons

The authors describe a novel multistimulus, multiple-matching learning paradigm for pigeons, which they believe not only simulates pigeons' natural foraging behavior but also more realistically shows their visual discrimination ability. A touch screen thin film transistor panel shows a stimulus configuration consisting of a 2-dimensional array of bitmap images that changes from trial to trial. The pigeon's task during each trial is to peck at images that are defined as positive by the experimenter. An experiment exploring the ability of pigeons to discriminate between achromatic male and female human faces demonstrated the effectiveness of the procedure. In contrast to 12 pigeons that show little improvement over 60 hr of go/no-go discrimination training, 14 multiple-matching pigeons mastered the problem within a few hours.     

The pigeon's discrimination of movement patterns (Lissajous figures) and contour-dependent rotational invariance

The ability of pigeons to discriminate complex motion patterns was investigated with the aid of moving Lissajous figures. The pigeons successfully learned to differentiate two successively presented cyclic trajectories of a single moving dot. This suggests that they can recognize a movement Gestalt when information about shape is minimal. They also quickly learned a new discrimination between moving-outline stimuli with repetitively changing contour patterns. Contrasting results were obtained when the dot or outline stimuli were axis-rotated through 90 degrees. Rotational invariance of pattern discrimination was clearly demonstrated only when moving contours were visible. Nevertheless, pigeons could discriminate the axis-orientation of a moving-dot or moving-outline pattern when trained to do so. Discrimination did not seem to depend on single parameters of motion but rather on the recognition of a temporally integrated movement Gestalt. The visual system of pigeons, as well as that of humans, may be well adapted to recognize the types of oscillatory movements that represent components of the motor behaviour shown by many living organisms.