Variables affecting establishment of schedule-induced attack on pictorial targets in White King pigeons.

White King pigeons exposed to food schedules before introduction of a colored photograph of a pigeon showed sustained schedule-induced attack on that image; additional birds given an early introduction to both the photograph and the schedule subsequently attacked the image at lower rates. Other pigeons attacked a second photograph of a pigeon regardless of whether it was introduced early or late. The late-introduction procedure was also effective in establishing attack on a projected image of a conspecific. The combined results showed that 14 of 17 White King pigeons given a late introduction to a pictorial target exhibited sustained attack against it and that a pigeon's initial reaction to a photograph of a conspecific when introduced early was a good predictor of subsequent schedule-induced attack on it.     

Pictorial target control of schedule-induced attack in White Carneaux pigeons

Three pigeons with a history of attacking a mirror target, and two of six pigeons with no prior exposure to targets, attacked a colored photograph of a conspecific during exposure to intermittent schedules of reinforcement for key pecking. Rate of attack on the photograph decreased when the reinforcement schedule was removed. The topography, temporal pattern, and locus of attack on the picture were comparable to schedule-induced attack on live, stuffed, and mirror targets. When silhouette, outline, and plain paper targets were used, schedule-induced attack was more sensitive to a change in target characteristics with a concurrent target-preference procedure than with an analogous successive-testing procedure. The combined results of the two testing procedures indicated that an “upright” white-on-black silhouette of a pigeon with or without an eye was more effective in controlling attack than was a comparable “inverted” silhouette, an outline of a pigeon, or a piece of colored paper.     

Modulation of aggression toward an intruder during the reproductive cycle of white king pigeons (columba livia)

For three of four mated male pigeons, intrusion of a still, two-dimensional image of a conspecific into the reproductive situation initiated aggressive behavior over months of daily testing with little habituation. There was a transient increase in aggressive contacts against the head of the target within two days prior to or following the occurrence of the second egg. Such an increase did not occur with extended daily testing of the male alone and was more closely synchronized to egg laying than time since the introduction of the female. The results are consistent with previous studies of reproductive and schedule-induced aggression in birds.     

Schedule-induced aggression in humans and animals: A comparative parametric review

Studies of animal and human aggression induced by schedules of reinforcement are reviewed. Parameters reviewed include: schedules, characteristics of induced aggression, subject variables, target variables, and the control or modification of schedule-induced aggression. With the exception of temporal locus of schedule-induced attack, the parameters identified with animal subjects are remarkably similar to those identified with human subjects.     

Attack-target attributes and pigeon aggression: Accessibility,vocalization, and body movements

Characteristics of the attack-target in laboratory tests appear to greatly influence the tendency for aggression to be initiated, maintained, and stopped. To address this question, accessibility, target movement, and vocalization among pigeon conspecifics were investigated in two different aggression tests: 1) paired aggression (PA); and 2) schedule-induced aggression (SI). In the PA test, dominant and subordinate roles formed quickly, and soon aggression episodes started to decline. In part, the subordinate's yielding behavior appeared to discourage attack. The significance of the target bird's behavior on attack frequency was borne out by the finding in the SI test that pigeons attacked a shielded, live, active target more often than a passive one. Furthermore, using a specially constructed stuffed pigeon in the SI test, results indicated that programmed, combined aggressive vocalization and body movement evoked most attacks with the VT schedule, vocalization alone evoked the next highest number of attacks, body movement alone the third most, and combined silence and immobility the least attacks. These findings provide a basis for explaining some past reported results associated with target features and a means for selecting other target features for future study.     

The assessment of individual “Aggressiveness” in pigeons by a variety of means

Pigeons attack or threaten animate and inanimate targets. The assessment of their aggressiveness was studied by exposing them in their home cages to three different stimuli: the experimenter's hand, a live pigeon, and a rear-projected conspecific image when the birds were exposed to intermittent access to food. A positive correlation between the hand test and live pigeon test was evident, but no relationship between either of these responses and the response to a pictorial image was observed. These results combined with other ethological observations cast doubts on the usefulness of schedule-induced responses to pictorial targets in the assessment of the individual aggressiveness in pigeons, but suggest that the hand test is an adequate and reliable procedure for such evaluations.     

Stimulus properties of conspecific behavior

Two experiments identified the conditions in which the behavior of one bird acquired discriminative control of the behavior of a second bird. The schedule-controlled behaviors of the “stimulus” bird were differentially correlated with the components of a multiple schedule according to which the pecking of an “experimental” bird produced food. In Experiment 1, three pairs of pigeons acquired a successive discrimination and two reversals with the conspecific stimuli. Experiment 2 included a control condition in which no systematic relationship existed between the conspecific stimuli and the component schedules. While differential responding during the components of the multiple schedule was again found when the conspecific stimuli were available, differential responding did not occur in the control condition. Test conditions included in the experiments indicated that (a) the differential responding was not dependent on the discriminative properties of reinforcement, (b) the pecking of the stimulus and experimental birds was temporally interrelated, (c) the visual conspecific stimuli were critical to the maintenance of the discrimination, and (d) the observed stimulus control immediately generalized to an unfamiliar conspecific.     

Effects of reinforcement amount on attack induced under a fixed-interval schedule in pigeons

Key pecking by pigeons was maintained on a chained fixed-interval 4-min (12-min for 1 subject) fixed-ratio 1 schedule of food presentation. Attacks toward a restrained and protected conspecific were recorded. In the first experiment, the amount of food presented per interval was manipulated across phases by varying the number of fixed ratios required in the terminal link of the chain. Measures of attack for all pigeons during the fixed-interval component increased monotonically as a function of food amount. In the second experiment, two different food amounts alternated within each experimental session under a multiple schedule. For both pigeons in this experiment, measures of attack were higher during the component that delivered the larger food amount per interval. The differences in levels of attack induced by the two food amounts in Experiment 2, however, were not as great as in Experiment 1; apparently this was because attack during the first interval of each component was controlled in part (P-5626) or entirely (P-7848) by the reinforcement amount delivered at the end of the previous component. Attack was also a function of the location of the interfood interval within the session. For both pigeons, attack tended to decrease throughout the session. The results of both experiments suggest that attack is an increasing function of reinforcement amount under fixed-interval schedules, but that this function may be influenced by the manner in which reinforcement amount is manipulated, by the duration of the interfood interval, and by the location of the interfood interval within the experimental session. In general, these results are compatible with theories of induced attack and other schedule-induced behavior that emphasize aversive after-effects of reinforcement presentation.     

Relation between level of food deprivation and rate of schedule-induced attack

The relation between food deprivation and schedule-induced attack was investigated in four White Carneaux pigeons. Attack toward a mirror target was induced by a schedule of reinforcement in which 3-sec food presentations occurred at alternate intervals of 15 and 120 sec (multiple fixed-time 15-sec fixed-time 120-sec schedule). A continuous tone was presented during the 15-sec periods; it was absent during the 120-sec periods. Each pigeon was tested at 65, 80, and 95% of its free-feeding weight in ascending, descending, and ascending orders, respectively. Two relations were apparent; an inverse relation between body weight and rate of attack, and a tendency for rate of attack to increase during the experiment. Reduction or elimination of attack when the mirror was covered with brown paper for some sessions indicated that the results were due neither to changes in activity that might covary with weight nor to habituation to the experimental situation.     

The role of intermittent food in the induction of attack in pigeons

The present experiments evaluated whether transitions in reinforcer probability are necessary to induce attack in pigeons. In Experiment I, three of six pigeons exposed to response-contingent constant-probability food schedules and a photograph of a conspecific as a target exhibited sustained postreinforcement attack on the target. The postreinforcement pattern of attack developed over the course of the experiment and was accompanied by a reduction in the rate of postreinforcement key pecking and an increase in the postreinforcement pause in key pecking. These effects on key pecking resulted in unprogrammed variations in the probability of reinforcement which may have been responsible for the induction of attack. In Experiment II, the attack-inducing properties of a constant-probability response-independent food schedule were compared to a periodic food schedule matched for overall rate of food delivery and to a no-food condition. In addition to attack, the spatial location of the subjects was monitored during each interfood interval. The periodic and aperiodic food schedules generated very different patterns of spatial location. Postfood attack was induced by both food schedules, although the constant-probability schedule induced attack in fewer birds. The no-food condition was not effective in inducing attack in any birds. These experiments indicate that intermittent food schedules without reductions in reinforcer probability are sufficient to induce attack in some pigeons, although not as effective as schedules with transitions in reinforcer probability.     

The effects of two response-elimination procedures on reinforced and induced aggression.

Pecks against a stuffed pigeon were reinforced according to a fixed-interval schedule for one group of pigeons and a variable-interval schedule for a second group. Red and green stimulus lights were alternately illuminated. Subsequently, food deliveries no longer occurred during one color (extinction). In the presence of the other color, food was presented only when no attack occurred for 30 sec. When attack produced food, all pigeons generally exhibited characteristic fixed-interval or variable-interval response patterns. Two birds in each group frequently exhibited postreinforcement schedule-induced aggression. Attack was reduced to low levels at approximately the same rate by extinction and differential reinforcement of other behavior. For birds that had previously exhibited schedule-induced aggression the initial reduction of attack during the second experimental phase was followed by induced attack immediately after food delivery in the differential-reinforcement-of-other-behavior component and upon onset of the extinction component, Either extinction or differential reinforcement of other behavior may eliminate reinforced aggression but may be relatively ineffective for reducing induced attack.     

Attack and defense in conspecific fighting in tree shrews (Tupaia belangeri)

Principles of conspecific defense have been analyzed for rodents, in which specific target sites for biting by attackers on defenders serve as an important determinant of the actions involved in both attacker and defender behavior. In an effort to determine the generality of these principles, attack and defensive behaviors and target sites for biting attack were evaluated in a nonrodent species, the tree shrew (Tupaia belangeri). Brief daily and repeated conspecific dyadic encounters between adult, socially experienced males (dominants, attackers), and adult, socially naive males (subordinates, defenders) that had been transferred into the territory of the dominants, produced a polarization of attack and defense. The dominant males showed chase, chase attack, jump attack, and biting behaviors, while the subordinates displayed flight and freezing. The vast majority of bites, as well as wounds and bruises, were on the subordinates’ backs. These patterns are very similar to those previously found in rats and mice and suggest that the organization of fighting, with targets of biting (or other painful) attack serving as an important determinant of both attacker (dominant) and defender (subordinate) behavior, may show considerable generality across nonrodent as well as rodent species. Although relatively few wounds were found after 28 days of repeated and daily encounters, the subordinate tree shrews show a variety of behavioral, neuroendocrine, and central nervous changes, indicating that they are stressed by these encounters per se. Aggr. Behav. 27:139–148, 2001. © 2001 Wiley‐Liss, Inc.     

Conspecific aggression in the laboratory rat.

Attacks by dominant colony males of a domesticated rat strain on conspecific strangers introduced into the colony include both a full range of threat displays and actual biting attack. These dominant males attack and bite both anesthetized and unanesthetized strangers, but threat displays and bites to anesthetized rats are limited. Lesions resulting from these bites are nonrandomly distributed, with most damage to the head and upper back and very few bites to ventral surfaces. When introduced strangers were left in the colonies for several days, 66% were killed, indicating that domesticated rats are capable of lethal attacks on conspecifics without external provocation.     

Effects of the cholinomimetic drug arecoline upon aggression: Intra- vs. inter-specific allocation of attack

Systemic injections of cholinomimetic drugs have been reported to induce both rage and predatory attack in several species. In order to assess the relative contribution of each of these two behavioral patterns in the control of cholinergically induced attack, a group of adult female cats was chemically stimulated in the simultaneous presence of both a prey object and a conspecific attack object. In this choice situation stimulated cats initially tended to engage in rage attack. When a second group of subjects was tested in a successive choice situation a significantly greater number of attacks occurred against conspecifics. The results suggest that cholmergic stimulation initially induces affective attack, with somewhat less frequent incidents of predation.     

Rank order in pairs of communally nursing female mice (Mus musculus domesticus) and Maternal Aggression Towards Conspecific Intruders of Differing Sex

This study examined social behavior between pairs of unfamiliar lactating females, with litters of the same age, at different periods after parturition (3, 7, and 17 days). Tests were generally followed by the formation of communal rearing nests, and subsequent maternal attack on intruders of differing sex was assessed. In all three intervals lactating females showed ritualized attack with formation of clear dominance-subordination relationships before combining litters into communal nests. The dominant females in 90% of cases started to retrieve alien pups into their nests. Agonistic behavior and communal nest formation were most rapid when pups were around 3 days old. Maternal attack on conspecific intruders was mainly displayed by the dominant lactating females. Male and female intruders were equally attacked (in terms of frequency and intensity of attack), but there was less such aggression when pups were around 17 days of age. Nevertheless the topography of biting attack employed against female and male conspecific intruders was different. Females were attacked using a strategy avoiding bites to the head and ventral surface (indicative of “offensive” behavior) whereas males were severely bitten on vulnerable body regions (indicative of “defensive” behavior).     

Responding maintained by the opportunity to attack during an interval food reinforcement schedule

Pigeons responded in a two-key situation. Responses on the right key (food key) were reinforced with food presentation on a response-initiated fixed-interval schedule, (i.e., first response after a fixed period of time was reinforced); responses on the left key (target key) were reinforced on a fixed-ratio schedule (i.e., every nth response was reinforced) with the presentation of a target bird that could be attacked. When the interval value of the food reinforcement schedule was varied from 1 min to 5 min, both the rate of attack responding on the target bird and the rate of responding on the target key were a function of the interval value. Responding on the target key was not maintained by the stimulus change associated with target availability, and was successively extinguished and reconditioned by removing and returning the target bird to the restraining box. When food was delivered independently of behavior, responding on the target key either remained unchanged or decreased, but was not eliminated. Responding on the target key was not maintained in the absence of an intermittent schedule of food presentation.     

The role of the head in configural body processing: Behavioural and electrophysiological evidence from the inversion and scrambling effect

The present study aimed to further explore the role of the head for configural body processing by comparing complete bodies with headless bodies and faceless heads (Experiment 1). A second aim was to further explore the role of the eye region in configural face processing (Experiment 2). Due to that, we conducted a second experiment with complete faces, eyeless faces, and eyes. In addition, we used two effects to manipulate configural processing: the effect of stimulus inversion and scrambling. The current data clearly show an inversion effect for intact bodies presented with head and faces including the eye region. Thus, the head and the eye region seem to be central for configural processes that are manipulated by the effect of stimulus inversion. Furthermore, the behavioural and electrophysiological body inversion effect depends on the intact configuration of bodies and is associated with the N170 as the face inversion effect depends on the intact face configuration. Hence, configural body processing depends not only on the presence of the head but rather on a complete representation of human bodies that includes the body and the head. Furthermore, configural face processing relies on intact and complete face representations that include faces and eyes.     

The effect of social learning on avoidance of aposematic prey in juvenile great tits (<Emphasis Type="Italic">Parus major</Emphasis>)

Social learning plays an important role in acquiring new foraging skills and food preferences in many bird species but its potential role in learning to avoid aposematic prey has never been studied. We tested the effect of social learning on the acquisition of avoidance of aposematic insect prey (firebug Pyrrhocoris apterus; Heteroptera) in juvenile, hand-reared great tits (Parus major). Behaviour towards aposematic prey was compared between two groups of birds: (1) the observers that were, prior to encounter with firebugs, allowed to watch the experienced conspecific demonstrator repeatedly refuse to attack the prey, and (2) the control birds that lacked this opportunity. Observing an experienced demonstrator was not sufficient for learning complete avoidance, because birds from both groups attacked at least the first firebug they had encountered in avoidance training. However, the opportunity to observe the avoidance behaviour of another bird significantly increased the rate of subsequent individual learning of observers in comparison with control birds. Social learning also decreased mortality of firebugs killed by the birds during the avoidance learning. Socially enhanced learning to avoid aposematic prey might be a mechanism important especially for naive juvenile birds learning from their parents, but it could also enhance learning in adults from their more experienced flock mates. Because social learning of avoidance may also lead to decreased mortality of aposematic prey, its effect should be taken into account in scenarios considering evolution and maintenance of prey warning signals.     

Assessment of attack and drinking in White King pigeons on response-independent food schedules.

Four White King pigeons in Experiment I were exposed to a fixed-time 90-second food schedule with successive access to water and a conspecific target. Drinking per session was sporadic and minimal, while attack per session occurred during most interfood intervals for all animals. Analysis of the temporal distribution of attack showed that the typical postreinforcement pattern of attack developed over the course of the experiment. In Experiment II, the same animals were exposed to a series of fixed-time schedules ranging from 30 to 360 seconds with successive access to water and target. Time engaged in drinking and the number of interfood intervals with drinking were less than that of attack. Food and no-food baselines, which have been typically used to assess schedules-induced drinking and attack, respectively, were used to evaluate the effect of the schedule on attack and water ingestion. Relative to the no-food baseline, both attack and drinking were enhanced by the schedule in all birds. Relative to the food baseline, drinking was slightly suppressed in three birds and attack was enhanced in all. For all animals, the food baseline resulted in more attack and drinking than the no-food baseline.     

The effects of signalled and unsignalled lick-dependent delays on the development of schedule-induced drinking in rats

Food pellets were programmed to be delivered to rats every 60 sec (Fixed Time 60-sec schedule), and the development of schedule-induced drinking was measured in terms of the amount of water consumed and the number of licks per inter-pellet interval. For some rats (masters) 10-sec delays in food delivery were dependent on licks. Yoked-control rats received food at the same time as their masters and independently of their own behaviour. In Experiment 1, in which the delays were signalled by a blackout, the master rats began to drink, but this schedule-induced behaviour then decreased to levels lower than those shown by the yoked controls. When the signalled delays were discontinued, the drinking of the master rats recovered. In Experiment 2, in which the delays were not signalled, the master rats did not develop as much schedule-induced drinking as the yoked controls, and discontinuing the delays led to only small increases in drinking. These results support the view that schedule-induced drinking is subject to control by its consequences.