The key mimetic features of hoverflies through avian eyes

Batesian mimicry occurs when a palatable species (the mimic) gains protection from predators by resembling an unpalatable or otherwise protected species (the model). While some mimetic species resemble their models closely, other species ('imperfect mimics') are thought to bear only a crude likeness. In an earlier study, pigeons (Columba livia) were trained to recognize wasp images in one experiment and non-mimetic (NM) fly images in another by rewarding the pigeons for pecking on the respective image types. These pigeons were subsequently presented with different images, including seemingly wasp-like hoverfly species, and the recorded peck rates on these images were used as a measure of the pigeons' perception of the hoverflies' mimetic similarity. To identify a candidate set of morphological features that the pigeons used when assessing this mimetic similarity, we first extracted a range of biometrical measurements from images originally presented to the pigeons. We then repeatedly optimized an empirical model in an attempt to match the recorded pigeon peck rates while using as few biometrical features as input as possible. Our models were able to fit the pigeon peck rates with considerable accuracy even while excluding many input features. Antennal length, a feature commonly used to discriminate between flies and wasps, was regularly retained as an input variable, but overall a different set of biometrical features was important for predicting the peck rates of pigeons rewarded for identifying wasps compared to those rewarded for identifying NM flies. In highlighting the importance of specific biometrical features in promoting mimicry and the irrelevance of others, our optimized models provide an explanation as to why certain species that appear to be poor mimics to humans are judged to be good mimics by birds.     

Eelectrophysiological correlates of pecking.

Electrophysiological correlates of pecking were studied in 50 freely moving pigeons. With the backward averaging technique average motor potentials (AMPs) were found in all subdivisions of the striatum. The AMPs appeared significantly earlier (40 msec before pecking) in the Wulst than in the neostriatum, ectostriatum, and archistriatum (18 to 28 msec before pecking). Their amplitude could be reduced by monocular occlusion. Average visual responses (AVRs) in the optic tectum were decreased at brief peck-flash intervals, reached maximum with 100 to 200 msec peck-flash delays and then again decreased at intervals corresponding to the occurrence of the next peck (300 msec). A head-mounted microdive system and a miniature FET probe were employed to record unit activity with glass microelectrodes. Out of 280 examined neurones, 83% were activated and 6% inhibited by pecking. The pre- and post-pecking histograms indicated that hyperstriatal neurones are activated earlier and for a longer time than the neostriatal units. It is concluded that rostral parts of striatum play an important role in the functional organisation of pecking.     

Effects of unsignaled delays of reinforcement on fixed-interval schedule performance

Key pecking of pigeons was maintained by a fixed-interval (FI) 61-s schedule. The effects of resetting and nonresetting unsignaled delays of reinforcement then were examined. The resetting delay was programmed as a differential-reinforcement-of-other-behavior schedule, and the nonresetting delay as a fixed-time schedule. Three delay durations (0.5, 1 and 10 s) were examined. Overall response rates were decreased by one and 10-s delays and increased by 0.5-s delays. Response patterns changed from positively accelerated to more linear when resetting or nonresetting 10-s delays were imposed, but remained predominantly positively accelerated when resetting and nonresetting 0.5- and 1-s delays were in effect. In general, temporal control, as measured by quarter-life values, changed less than overall response rates when delays of reinforcement were in effect. The response patterns controlled by FI schedules are more resilient to the nominally disruptive effects of delays of reinforcement than are corresponding overall response rates.     

Behavioral contrast of a complex operant

Two experiments were conducted in which pigeons were trained to perform a complex operant consisting of a peck to one key followed by a peck to another key. In the first experment this performance was reinforced on a variable-interval schedule and the birds were then subjected to a multiple schedule in which the variable-interval was alternated with extinction. The first two pigeons trained showed some deterioration of the cohesiveness of the response sequence. After measures were taken to correct this the pigeon gave evidence of behavioral contrast in a condition where sessions with discrimination were alternated rapidly with sessions with no discrimination (variable-interval only). Another two birds were trained on variable-interval followed by discrimination. One of these birds showed evidence of contrast but the results of the other were not clear.Experiment II was a three phase experiment in which animals were first trained to make the complex response. In the second phase they were extinguished by providing noncontingent reinforcement in each of two components. In the third phase reinforcement was removed from one component, providing for a stimulus-reinforcer contingency. In the third phase the complex response reappeared temporarily.The results of these two studies indicate that a complex response may show behavioral contrast and that it may be enhanced by stimulus-reinforcer contingencies if it has already been trained.     

Differential resurgence and response elimination

Resurgence refers to the transient recovery of previously reinforced, but presently not reinforced, responding when more recently reinforced responding is extinguished. The primary purpose of our research was to determine how differential resurgence results from the procedures used to eliminate that responding. There were three conditions in each of five experiments. In Condition 1, key pecking by pigeons was maintained under a two-component multiple variable-interval (VI) 30-s VI 30-s schedule. In Condition 2, this pecking was eliminated in different ways across components. In Condition 3, extinction was in effect for all responses, and resurgence of key pecking was compared across components. These three conditions were repeated for most pigeons, and the procedures used to eliminate responding in Condition 2 varied across experiments. In Experiment 1, there was greater resurgence, and an earlier onset of it, after a differential-reinforcement-of-other-behavior (DRO) schedule than after a VI schedule was correlated with pecking an alternative key. Experiments 2 and 3 showed that the differential resurgence in Experiment 1 probably was not due to conditional stimulus control or the periodicity of food delivery, respectively. In Experiment 4, there was no systematic difference in resurgence after either a DRO schedule or a VI schedule correlated with treadle pressing. In Experiment 5, there was greater resurgence, and/or an earlier onset of it, after a VI schedule correlated with treadle pressing than after a VI schedule correlated with pecking an alternative key. Taken together, the results showed that the reinforcement of an alternative key-peck response was the most effective means of reducing subsequent key-peck resurgence. The relation of these results to an understanding of resurgence is discussed.     

Effects of methamphetamine on response rate: a microstructural analysis

Key pecking in pigeons was maintained under a multiple random-interval (RI) 1-min, RI 4-min schedule of food presentation. Several doses (0.3-5.6 mg/kg) of methamphetamine were administered, and effects on overall response rates and on the microstructure of responding were characterized. In three of the four pigeons, methamphetamine dose-dependently decreased overall response rate in both components; in the fourth pigeon, intermediate doses increased response rates. Log-survivor analyses did not produce the clear "broken-stick" pattern previously reported with rats [Shull, R.L., Gaynor, S.T., Grimes, J.A., 2001. Response rate viewed as engagement bouts: effects of relative reinforcement and schedule type. J. Exp. Anal. Behav. 75, 247-274]. A fine-grained analysis of inter-response times (IRTs) revealed clear bands of responding around certain IRT durations. Methamphetamine tended to decrease the frequency of IRTs in the shorter bands and increase the frequency of IRTs across all bins greater than 2s. These results suggest that (a) survivor analyses may not extend to pigeon key pecking, (b) microstructural analyses can reveal order not evident with overall response rate, and (c) a detailed analysis of responding might prove more useful than summary measures in characterizing drug effects on behavior.     

Automatic imitation in budgerigars

A fully automated procedure, involving computer-controlled stimulus presentation and computer-recorded response measurement, was used for the first time to study imitation in non-human animals. After preliminary training to peck and step on a manipulandum, budgerigars were given a discrimination task in which they were rewarded with food for pecking during observation of pecking and for stepping during observation of stepping (Compatible group), or for pecking while observing stepping and for stepping while observing pecking (Incompatible group). The Incompatible group, which had to counter-imitate for food reward, showed weaker discrimination performance than the Compatible group. This suggests that, like humans, budgerigars are subject to 'automatic imitation'; they cannot inhibit online the tendency to imitate pecking and/or stepping, even when imitation of these behaviours interferes with the performance of an ongoing task. The difference between the two groups persisted over 10 test sessions, but the Incompatible group eventually acquired the discrimination, making more counter-imitative than imitative responses in the final sessions. These results are consistent with the associative sequence learning model, which suggests that, across species, the development of imitation and the mirror system depends on sensorimotor experience and phylogenetically ancient mechanisms of associative learning.     

Response-cost punishment via token loss with pigeons

Two experiments were conducted to investigate punishment via response-contingent removal of conditioned token reinforcers (response cost) with pigeons. In Experiment 1, key pecking was maintained on a two-component multiple second-order schedule of token delivery, with light emitting diodes (LEDs) serving as token reinforcers. In both components, responding produced tokens according to a random-interval 20-s schedule and exchange periods according to a variable-ratio schedule. During exchange periods, each token was exchangeable for 2.5-s access to grain. In one component, responses were conjointly punished according to fixed-ratio schedules of token removal. Response rates in this punishment component decreased to low levels while response rates in the alternate (no-punishment) component were unaffected. Responding was eliminated when it produced neither tokens nor exchange periods (Extinction), but was maintained at moderate levels when it produced tokens in the signaled absence of food reinforcement, suggesting that tokens served as effective conditioned reinforcers. In Experiment 2, the effect of the response-cost punishment contingency was separated from changes in the density of food reinforcement. This was accomplished by yoking either the number of food deliveries per component (Yoked Food) or the temporal placement of all stimulus events (tokens, exchanges, food deliveries) (Yoked Complete), from the punishment to the no-punishment component. Response rates decreased in both components, but decreased more rapidly and were generally maintained at lower levels in the punishment component than in the yoked component. In showing that the response-cost contingency had a suppressive effect on responding in addition to that produced by reductions in reinforcement density, the present results suggest that response-cost punishment shares important features with other forms of punishment.     

Categorical counting

Pigeons pecked on three keys, responses to one of which could be reinforced after a few pecks, to a second key after a somewhat larger number of pecks, and to a third key after the maximum pecking requirement. The values of the pecking requirements and the proportion of trials ending with reinforcement were varied. Transits among the keys were an orderly function of peck number, and showed approximately proportional changes with changes in the pecking requirements, consistent with Weber's law. Standard deviations of the switch points between successive keys increased more slowly within a condition than across conditions. Changes in reinforcement probability produced changes in the location of the psychometric functions that were consistent with models of timing. Analyses of the number of pecks emitted and the duration of the pecking sequences demonstrated that peck number was the primary determinant of choice, but that passage of time also played some role. We capture the basic results with a standard model of counting, which we qualify to account for the secondary experiments.     

Selective stimulus control in discrimination of lateral mirror images by pigeons

Four monocularly and two binocularly viewing pigeons were trained to peck a key when it displayed one stimulus (S+) but not to peck when it displayed another stimulus (S−). S+ and S− were a lateral mirror-image pair of two-coloured stimuli. When tested for transfer with the untrained eye open, two of the monocular birds pecked more during S− than S+, the other two continuing to favour S+. During generalization tests on the wavelength dimension all monocular birds pecked much more often during one S+ colour than during the other. The colour controlling pecking was that displayed on the side of the key facing the open eye during S+ presentations. Both binocular birds developed asymmetrical responses to the key, one favouring the left, the other the right side of the key. Generalization tests on the wavelength dimension showed selective control by the colour displayed on the favoured side of the key during S+ presentations. The results are interpreted as supporting the view that pigeons learn to discriminate lateral mirror images by developing asymmetrical observing responses that convert the left-right difference between the mirror images into a difference more easily discriminable.     

Some factors affecting pigeons' visual tracking behavior

A novel delayed conditional discrimination procedure was used successfully to investigate some factors affecting pigeons' tracking of visual stimuli. Trials began with the brief illumination of one of the keys (randomly selected) from the left- or right-hand column of a three by three matrix of pecking keys. Then the next key in the same row was briefly lit. Finally, the remaining key in the row was lit along with one (randomly chosen) of the remaining five keys from the left- or right-hand columns. A peck to the former but not the latter key was designated as correct and was rewarded. In Experiment 1 pigeons made correct choices on between 70 and 80 percent of trials, thereby demonstrating an ability to visually track objects. In Experiment 2 tracking accuracy was: a) reduced when either the first or second key in the sequence was omitted, b) improved when the sequence was repeated three times, c) reduced when duration of key illumination was reduced, and d) reduced by delays imposed between keys early but not late in the sequence. It was also found that tracking of vertical and horizontal sequences was approximately equal.     

Simple and multiple schedule responding and behavioral contrast when pigeons press treadles

Positive behavioral contrast has been observed when pigeons press treadles on multiple schedules for high rates of reinforcement, but not for low rates. Negative treadle-press contrast has been observed for low rates of reinforcement. Two experiments showed that differences between response rates emitted during simple and multiple schedules appear and fail to appear under similar conditions. The experiments showed that the rate of pressing during the less favorable component of a multiple schedule was less than the rate of pressing during a comparable simple schedule (negative contrast). The rate of treadle–pressing during the more favorable component was not greater than the rate of pressing during a comparable simple schedule, when the schedules provided a low rate of reinforcement (absence of positive contrast), but it was when the schedules provided a high rate of reinforcement (positive contrast). These results help to clarify the definition of behavioral contrast by showing that simple schedules may be appropriate baselines from which to define and measure contrast.     

Free-choice preference under uncertainty

Preferences in pigeons for free choice over forced choice under uncertain contingencies were compared with the one under certain contingencies in multiple concurrent-chain schedules of reinforcement. The uncertain condition examined the preference for two alternatives over one alternative when reinforcement probability at the end of the terminal link equalled 0.5, and with all keys in each terminal link lit green (two mixed fixed-interval extinction keys versus one mixed fixed-interval extinction key). Key and schedule arrangement in the certain condition was the same except that a peck on any terminal-link key after the FI interval always produced food. When naive pigeons were first exposed to uncertain contingencies, preference for two lit keys over one lit key was observed, and the preference was confirmed by sequential reversals of the terminal-link contingencies. However, no consistent preference was observed when uncertain condition followed the certain condition. Under certain contingencies, unlike the earlier experiments, very small and inconsistent preferences for free choice were demonstrated. A possible reason for the different preferences in the uncertain conditions was that pigeons may lessen their sensitivity to the circumstances with uncertainty by any history or carry-over effect of the prior contingency.     

Some characteristics of spatial associative memory in the pigeon, Columba livia

Willson and Wilkie (1993) developed a novel procedure to assess pigeons' memory for the spatial location of food. Only one of four locations provided food each daily session. Each location consisted of an illuminated pecking key and grain feeder. Over different days different locations, randomly selected, provided food during a 16-min session. The pigeons tended to revisit the location at which food was found on the previous day thereby demonstrating memory for food-spatial location associations over 24 h. Three experiments were conducted to further investigate this phenomenon. In Experiment 1 the session duration was varied between 4 and 32 min. Longer sessions had no detectable effect on their ability to remember the rewarded location 24 h later, a result that suggests that only brief encounters with food at a particular location are necessary for recall. In Experiment 2 the necessity of an active search for the day's rewarded location was removed; a 5-min period in which only the rewarded key was lit preceded the regular 16-min session. Pecks to the lit key in this 5-min period produced grain on the standard schedule. This manipulation facilitated the pigeons' discovery of food but did not affect their ability to remember the rewarded location, suggesting that the process of search and discovery is not essential to the associative memory process. In Experiment 3, food was available during the complete session (non-depleting condition) or was available only during the first half of the session (depleting condition). No detectable differences in the birds' memory of yesterday's profitable location were found. This suggests that non-depletion of food is not a necessary condition for day-to-day recall of food location. Taken together these findings enlarge our understanding of the spatial associative memory process.     

Within-trial contrast: The effect of probability of reinforcement in training

There is evidence that pigeons prefer conditioned reinforcers that are preceded by greater effort over those that are preceded by less effort (an effect that has been attributed to within-trial contrast). In past research the probability of reinforcement for correct choice of the conditioned reinforcer has been 100%, however, the high level of reinforcement for both alternatives in training may result in a performance ceiling when choice between those alternatives is provided on test trials. In the present study we tested this hypothesis by including a group for which the probability of reinforcement in training was only 50%. Pigeons were trained on two simultaneous discriminations, one that was preceded by a 30 peck requirement the other by a single peck requirement. On test trials, we found a significant preference for the S+ that required the greater effort in training for pigeons trained with 100% and a small but nonsignificant effect for pigeons trained with 50% reinforcement. Although the hypothesis that the within-trial contrast effect was constrained by a performance ceiling was not confirmed, we did find a reliable within-trial contrast effect with 100% reinforcement.     

Interactions among unit-price, fixed-ratio value, and dosing regimen in determining effects of repeated cocaine administration

Previous research has shown tolerance to cocaine that was dependent on fixed-ratio (FR) parameter size in the context of a multiple FR schedule of food reinforcement. Completion of the FR requirement in these studies resulted in the same magnitude of reinforcement, regardless of ratio size. The cost-to-benefits ratio (unit-price) was therefore not equated across the different FR components. The current study examined the effects of repeated administration of cocaine to pigeons when unit-price under FR schedules was either the same or different. Additionally, the role of a chronic variable-dosing versus chronic fixed-dosing procedure was examined when unit-price was equated across different ratio values. Pigeons were trained to key peck in daily sessions under a three-component multiple FR schedule of food presentation, according to which either 10, 30, or 100 pecks were required for each delivery of food. In Experiment 1, completion of the FR 10, FR 30, and FR 100 resulted in 1.5, 4.5, and 15.0 s access to food, respectively. That is, the response requirement was correlated with access to food time so that unit-price (pecks per second of access to food) was equated across components. After assessing acute effects of a range of doses of cocaine, drug administration occurred daily before each session, with the dose varying from day to day. Tolerance, the magnitude of which was unrelated to the peck requirement, developed under the repeated-dosing regimen. In order to assess whether having equal unit-price was responsible for producing similar levels of tolerance across components, daily drug administration continued in Experiment 2 using the variable-dose regimen, but the amount of food presented each time was fixed at 4.5 s access to food, yielding different unit-prices under the three pecking requirements. Subsequently, the conditions of Experiment 1 were reinstated, i.e., unit-price was equated. Making unit-price different or the same had little influence on effects of cocaine. In Experiment 3, a fixed dose of cocaine was administered before each session while programmed unit-price remained the same across components. Under these conditions, tolerance became peck-requirement related. Specifically, tolerance was most prevalent under the smaller requirements and less robust or absent when the largest requirement was in effect. Differences in unit-price, therefore, were not related to degree of tolerance, but work requirement was. Differences in effects of cocaine across responses requirements, however, were observed only when each session was preceded by the same dose, not when dose varied from session to session.     

Extinction as discrimination: the molar view

The traditional molecular view of behavior explains extinction as the dissipation or inhibition of strength, formerly built up by contiguous reinforcement. In obstinate opposition to this explanation was the partial-reinforcement extinction effect: a partially reinforced response extinguishes more slowly than a continuously reinforced response. It suggests instead that extinction is discrimination. Four pigeons were exposed to daily sessions in which a variable period of food delivery, produced by pecking on a variable-interval schedule, was followed by extinction. The rate of food delivery was varied over a wide range across conditions. Varying the amount of food per delivery inversely with rate of delivery kept response rate from varying excessively. The results confirmed and extended the partial-reinforcement effect; persistence of pecking and time to extinction were inversely related to rate of obtaining food. The results support the molar view of extinction, not as loss of strength of a particular discrete response, but as a transition from one allocation of time among activities to another. Although molecular theories dismiss discrimination due to repeated training and extinction as an impurity or complication, repeated cycles of availability and privation are probably typical of the environment in which most vertebrate species evolved.     

A further study of choice and percentage reinforcement

In Experiment I four pigeons were trained in a concurrent chains procedure with fixed-ratio schedules (FR1) in the initial components and fixed-time schedules in the terminal components. Pecking one of the keys when both keys were white initiated a fixed time schedule on that key. A peck to the left key produced three stripes on the key. At the termination of the fixed-time component food always occurred. Pecking the other key produced either a circle or a triangle. If a circle appeared, reinforcement occurred. If a triangle appeared a brief timeout was given. Initially the stripes appeared on the left key and the circle and triangle on the left. This was reversed during the course of the experiment. In addition, sessions were conducted in which both circle and triangle sometimes preceded reinforcement and sometimes timeout. For most birds under most conditions there was a preference for the key that produced the circle and triangle. When these were uncorrelated with reinforcement and time out three of the birds preferred the key producing 100% reinforcement.

In Experiment II three factors were varied and VI 20 sec schedules were used in the initial links instead of FR1. The results showed that pigeons preferred the 50% condition more 1) the greater the duration of the terminal links, 2) the smaller the value on the initial link VI schedules and 3) the less the probability of food in the terminal link with stripes on the key.     

Reaction to frustration in high and low feather pecking laying hens

Reaction to frustration of high (HFP) and low feather pecking (LFP) laying hens was investigated. From a HFP- and a LFP-line five birds with a HFP- and five birds with a LFP-phenotype were selected. Birds from the HFP-line were expected to show more key pecking and covered feeder pecking during frustration than birds from the LFP-line. When a bunch of feathers was presented, birds with a HFP-phenotype were expected to redirect their pecks at the bunch. Birds were trained to peck a key for a food reward in an automated Skinnerbox and subjected to two sessions: a control session, where food was available, and a frustration session, where the feeder was covered with Perspex. These two sessions were repeated in the presence of a bunch of feathers. Unexpectedly, birds from the LFP-line had a stronger reaction to frustration than birds from the HFP-line, expressed in pecking behaviour. When a bunch of feathers was offered, birds with a HFP-phenotype did not show more bunch pecking during frustration than birds with a LFP-phenotype.