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1.  The Temporal Dynamics of Implicit Processing of Non-Letter, Letter, and Word-Forms in the Human Visual Cortex 
The decoding of visually presented line segments into letters, and letters into words, is critical to fluent reading abilities. Here we investigate the temporal dynamics of visual orthographic processes, focusing specifically on right hemisphere contributions and interactions between the hemispheres involved in the implicit processing of visually presented words, consonants, false fonts, and symbolic strings. High-density EEG was recorded while participants detected infrequent, simple, perceptual targets (dot strings) embedded amongst a of character strings. Beginning at 130 ms, orthographic and non-orthographic stimuli were distinguished by a sequence of ERP effects over occipital recording sites. These early latency occipital effects were dominated by enhanced right-sided negative-polarity activation for non-orthographic stimuli that peaked at around 180 ms. This right-sided effect was followed by bilateral positive occipital activity for false-fonts, but not symbol strings. Moreover the size of components of this later positive occipital wave was inversely correlated with the right-sided ROcc180 wave, suggesting that subjects who had larger early right-sided activation for non-orthographic stimuli had less need for more extended bilateral (e.g., interhemispheric) processing of those stimuli shortly later. Additional early (130–150 ms) negative-polarity activity over left occipital cortex and longer-latency centrally distributed responses (>300 ms) were present, likely reflecting implicit activation of the previously reported ‘visual-word-form’ area and N400-related responses, respectively. Collectively, these results provide a close look at some relatively unexplored portions of the temporal flow of information processing in the brain related to the implicit processing of potentially linguistic information and provide valuable information about the interactions between hemispheres supporting visual orthographic processing.
PMCID: PMC2796900  PMID: 20046826
word reading; ERPs; visual cortex; visual orthography
2.  Are All Letters Really Processed Equally and in Parallel? Further Evidence of a Robust First Letter Advantage 
Acta psychologica  2013;144(2):10.1016/j.actpsy.2013.07.018.
This present study examined accuracy and response latency of letter processing as a function of position within a horizontal array. In a series of 4 Experiments, target-strings were briefly (33 ms for Experiment 1 to 3, 83 ms for Experiment 4) displayed and both forward and backward masked. Participants then made a two alternative forced choice. The two alternative responses differed just in one element of the string, and position of mismatch was systematically manipulated. In Experiment 1, words of different lengths (from 3 to 6 letters) were presented in separate blocks. Across different lengths, there was a robust advantage in performance when the alternative response was different for the letter occurring at the first position, compared to when the difference occurred at any other position. Experiment 2 replicated this finding with the same materials used in Experiment 1, but with words of different lengths randomly intermixed within blocks. Experiment 3 provided evidence of the first position advantage with legal nonwords and strings of consonants, but did not provide any first position advantage for non-alphabetic symbols. The lack of a first position advantage for symbols was replicated in Experiment 4, where target-strings were displayed for a longer duration (83 ms). Taken together these results suggest that the first position advantage is a phenomenon that occurs specifically and selectively for letters, independent of lexical constraints. We argue that the results are consistent with models that assume a processing advantage for coding letters in the first position, and are inconsistent with the commonly held assumption in visual word recognition models that letters are equally processed in parallel independent of letter position.
PMCID: PMC3859888  PMID: 24012723
letter-processing; serial position; visual word recognition
3.  Letter-transposition effects are not universal: The impact of transposing letters in Hebrew 
Journal of memory and language  2009;61(3):285-302.
We examined the effects of letter transposition in Hebrew in three masked-priming experiments. Hebrew, like English has an alphabetic orthography where sequential and contiguous letter strings represent phonemes. However, being a Semitic language it has a non-concatenated morphology that is based on root derivations. Experiment 1 showed that transposed-letter (TL) root primes inhibited responses to targets derived from the non-transposed root letters, and that this inhibition was unrelated to relative root frequency. Experiment 2 replicated this result and showed that if the transposed letters of the root created a nonsense-root that had no lexical representation, then no inhibition and no facilitation were obtained. Finally, Experiment 3 demonstrated that in contrast to English, French, or Spanish, TL nonword primes did not facilitate recognition of targets, and when the root letters embedded in them consisted of a legal root morpheme, they produced inhibition. These results suggest that lexical space in alphabetic orthographies may be structured very differently in different languages if their morphological structure diverges qualitatively. In Hebrew, lexical space is organized according to root families rather than simple orthographic structure, so that all words derived from the same root are interconnected or clustered together, independent of overall orthographic similarity.
PMCID: PMC2748959  PMID: 20161017
Morphology; Letter Transposition; Hebrew; Masked-Priming
4.  Evidence for Letter-Specific Position Coding Mechanisms 
PLoS ONE  2013;8(7):e68460.
The perceptual matching (same-different judgment) paradigm was used to investigate precision in position coding for strings of letters, digits, and symbols. Reference and target stimuli were 6 characters long and could be identical or differ either by transposing two characters or substituting two characters. The distance separating the two characters was manipulated such that they could either be contiguous, separated by one intervening character, or separated by two intervening characters. Effects of type of character and distance were measured in terms of the difference between the transposition and substitution conditions (transposition cost). Error rates revealed that transposition costs were greater for letters than for digits, which in turn were greater than for symbols. Furthermore, letter stimuli showed a gradual decrease in transposition cost as the distance between the letters increased, whereas the only significant difference for digit and symbol stimuli arose between contiguous and non-contiguous changes, with no effect of distance on the non-contiguous changes. The results are taken as further evidence for letter-specific position coding mechanisms.
PMCID: PMC3699680  PMID: 23844204
5.  Mechanisms underlying the effects of unsignaled delayed reinforcement on key pecking of pigeons under variable-interval schedules. 
Three experiments were conducted to test an interpretation of the response-rate-reducing effects of unsignaled nonresetting delays to reinforcement in pigeons. According to this interpretation, rates of key pecking decrease under these conditions because key pecks alternate with hopper-observing behavior. In Experiment 1, 4 pigeons pecked a food key that raised the hopper provided that pecks on a different variable-interval-schedule key met the requirements of a variable-interval 60-s schedule. The stimuli associated with the availability of the hopper (i.e., houselight and keylight off, food key illuminated, feedback following food-key pecks) were gradually removed across phases while the dependent relation between hopper availability and variable-interval-schedule key pecks was maintained. Rates of pecking the variable-interval-schedule key decreased to low levels and rates of food-key pecks increased when variable-interval-schedule key pecks did not produce hopper-correlated stimuli. In Experiment 2, pigeons initially pecked a single key under a variable-interval 60-s schedule. Then the dependent relation between hopper presentation and key pecks was eliminated by arranging a variable-time 60-s schedule. When rates of pecking had decreased to low levels, conditions were changed so that pecks during the final 5 s of each interval changed the keylight color from green to amber. When pecking produced these hopper-correlated stimuli, pecking occurred at high rates, despite the absence of a peck-food dependency. When peck-produced changes in keylight color were uncorrelated with food, rates of pecking fell to low levels. In Experiment 3, details (obtained delays, interresponse-time distributions, eating times) of the transition from high to low response rates produced by the introduction of a 3-s unsignaled delay were tracked from session to session in 3 pigeons that had been initially trained to peck under a conventional variable-interval 60-s schedule. Decreases in response rates soon after the transition to delayed reinforcement were accompanied by decreases in eating times and alterations in interresponse-time distributions. As response rates decreased and became stable, eating times increased and their variability decreased. These findings support an interpretation of the effects of delayed reinforcement that emphasizes the importance of hopper-observing behavior.
PMCID: PMC1284652  PMID: 9540229
6.  Effects of stimulus frequency and reinforcement variables on reaction time 
Pigeons pecked at one of two black forms, “+” or “O,” either of which could appear alone on a white computer monitor screen. In baseline series of sessions, each form appeared equally often, and two pecks at it produced food reinforcement on 10% of trials. Test series varied the relative probability or duration of reinforcement or frequency of appearance of the targets. Peck reaction times, measured from target onset to the first peck, were found to vary as a function of reinforcement probability but not as a function of relative target frequency or of reinforcement duration. Reaction times to the two targets remained approximately equal as long as the probability of reinforcement, per trial, was equal for the targets, even if the relative frequency of the targets differed by as much as 19 to 1. The results address issues raised in visual search experiments and indicate that attentional priming is unimportant when targets are easy to detect. The results also suggest that equalizing reinforcement probability per trial for all targets removes differential reinforcement as an important variable. That reaction time was sensitive to the probability but not the duration of reinforcement raises interesting questions about the processes reflected in reaction time compared with rate as a response measure.
PMCID: PMC1323068  PMID: 16812648
visual search; attention; priming; contrast; reaction time; reinforcement probability; reinforcement duration; peck; pigeon
Behavioural pharmacology  2008;19(4):347-352.
The present experiment was concerned with the role of environment in the production and form of apomorphine-induced pecking of pigeons. Previous literature has suggested that the pecking occurs even when pigeons are placed in complete darkness, but there are no systematic or quantitative reports of such pecking. Six pigeons were tested with doses of 0.1, 0.3, and 1.0 mg/kg apomorphine. Tests were made in conditions of white and infrared light. The apparatus employed novel force transduction measures that provided for both the detection of a peck as well as its peak forcefulness. At the lowest dose tested, apomorphine elicited pecking when the pigeon was placed in white light, but not when the dose was examined under infrared lighting. As the dose increased, however, pecking was observed regardless of lighting condition. No differences were found in forcefulness of pecking as a function of lighting condition or dose. Though response output was seemingly unaffected by the lighting condition at higher doses, videotaped analysis revealed important changes in the formal characteristics of pecking. In white light, apomorphine elicited pecking at stimuli in the chamber (e.g., screw heads or the pigeon’s own toes), whereas in infrared light pecking was directed at the floor directly in front of the pigeon. Such differences may be attributable to shifts in control to other stimulus modalities when vision in limited. Additionally, apomorphine may have direct effects on retinal dopamine function modulating the expression of pecking in the dark.
PMCID: PMC3198829  PMID: 18622183
Apomorphine; Stereotypy; Infrared light; White light; Force; Pigeon; Peck
8.  Behavioral research in pigeons with ARENA: An automated remote environmental navigation apparatus 
Behavioural processes  2009;81(1):105-113.
Three experiments established the effectiveness of an Automated Remote Environmental Navigation Apparatus (ARENA) developed in our lab to study behavioral processes in pigeons. The technology utilizes one or more wireless modules, each capable of presenting colored lights as visual stimuli to signal reward and of detecting subject peck responses. In Experiment 1, subjects were instrumentally shaped to peck at a single ARENA module following an unsuccessful autoshaping procedure. In Experiment 2, pigeons were trained with a simultaneous discrimination procedure during which two modules were illuminated different colors; pecks to one color (S+) were reinforced while pecks to the other color (S−) were not. Pigeons learned to preferentially peck the module displaying the S+. In Experiment 3, two modules were lit the same color concurrently from a set of six colors in a conditional discrimination task. For three of the colors pecks to the module in one location (e.g., upper quadrant) were reinforced while for the remaining colors pecks at the other module (e.g., lower quadrant) were reinforced. After learning this discrimination, the color-reinforced location assignments were reversed. Pigeons successfully acquired the reversal. ARENA is an automated system for open-field studies and a more ecologically valid alternative to the touchscreen.
PMCID: PMC2680771  PMID: 19429204
pigeon; autoshaping; conditional discrimination; open field; touchscreen; simultaneous discrimination
9.  Short-term memory in the pigeon: relative recency1 
Three pigeons pecked for food in an experiment in which each trial consisted of two phases. The first phase consisted of a pattern of three successively illuminated, randomly selected left or right keys. A subject was required to peck each of the lighted keys as they appeared. Thus, in the first phase, a subject emitted a pattern of three left- or right-key pecks. Over trials, all eight possible patterns appeared. A time interval separated the first phase from the second phase, which began with presentation of a randomly selected one of three cues. A reinforcer was delivered in the second phase if a subject pecked the side key that had appeared in the first phase in an ordinal position corresponding to the cue presented in the second phase. That is, the three cues probed a pigeon's memory for the side key it had pecked first, second, or third, in the first phase of a trial. The results show that a pigeon can remember for more than 4 sec the order in which it has just seen and pecked two lighted keys: a pigeon can remember the temporal organization or pattern of events in its recent environment. Consequently, the functional stimulus present when a reinforcer is delivered may include a subject's short-term memory for the temporal organization of recent events, such as the pattern of its own recent behavior. This possibility is consistent with a molecular analysis of operant behavior focusing on local patterns of behavior.
PMCID: PMC1333414  PMID: 16811895
short-term memory; delayed stimulus control; relative recency; behavioral patterns; pigeons
10.  Increasing the variability of response sequences in pigeons by adjusting the frequency of switching between two keys. 
Three experiments compared the amounts of behavioral variability generated with two reinforcement rules. In Experiments 1 and 2 pigeons received food whenever they generated a sequence of eight pecks, distributed over two keys, provided that the sequence contained a certain number of change-overs between the keys. Although no variability was required-the birds could obtain all reinforcers by repeating the same sequence-the pigeons emitted a large number of different sequences. In Experiment 3 pigeons received food whenever they generated a sequence that had not occurred during the last 25 trials. After prolonged training, the birds showed more sequence variability than in the first two experiments. The analysis of the internal structure of the response sequences revealed that, in general, (a) the location of the first peck was highly stereotyped; (b) as the trial advanced, the probability of switching to the initially preferred key decreased whereas the probability of switching to the other key increased; and (c) a first-order Markov chain model with transition probabilities given by a logistic function accounted well for the internal structure of the birds' response sequences. These findings suggest that, to a large extent, the variability of response sequences is an indirect effect of adjustments in changeover frequency.
PMCID: PMC1284614  PMID: 9241860
11.  A comparison of the key-peck and treadle-press operants in the pigeon: differential-reinforcement-of-low-rate schedule of reinforcement1 
Key pecking and treadle pressing in pigeons were compared under concurrent (key-treadle) and single-operant differential-reinforcement-of-low-rate schedules of food reinforcement ranging from 5 to 60 sec (concurrent procedure) or 5 to 120 sec (single-operant procedure). Under both procedures, the two operants followed the same general law: decreasing response rate and reinforcement rate and increasing number of responses per reinforcement as a function of increasing schedule interval. High correlations were found between key pecking and treadle pressing for the measures of response rate, reinforcement rate, and responses per reinforcement. Regression equations allowed the prediction of treadle pressing from key pecking. More bursting occurred in responding to the key, and key pecking showed a more precise temporal discrimination than treadle pressing. A test for sequential dependencies between key and treadle responses showed significant dependencies not only under the concurrent procedure but also in data created artificially by merging key and treadle sequences from different pigeons under the concurrent procedure and from the same pigeon under the single-operant procedure. It seems likely that the sequential dependencies found were due to the independent action of the schedule on each operant and that behavioral dependencies did not occur with the concurrent training procedure. The key-peck operant does not appear to have any special qualities that preclude its use in discovering general laws of behavior, at least under the differential-reinforcement-of-low-rate schedule. The usefulness of the key peck in other situations requires direct experimental study.
PMCID: PMC1333511  PMID: 16811945
differential reinforcement of low rate; concurrent schedules; spaced responding; IRT distributions; treadle pressing; key pecking; pigeons
12.  A preliminary analysis of the dynamics of the pecking response in pigeons1 
The pecking response of pigeons is usually measured by a transducer that senses the presence or absence of a response. Typically, the response force as a function of time has not been accurately measured. Data were collected using a transducer specially designed to record the waveform of the pecking response in pigeons. Each response on the target surface of the transducer was reinforced and followed by a blackout. The response was stored on an oscilloscope screen and the peak force and duration of the response were recorded manually from the oscilloscope screen. The mean peak force of the response substantially exceeded the minimum criterion for reinforcement of 35 g (0.343 Newtons) of force. Photographs of the waveform of pecks on the transducer showed great variability in response force and demonstrated that the waveform produced by pecking the target surface was complex. The responses were frequently asymmetrical with the rise time shorter than the fall time, although no single verbal description could be applied to all of the waveforms. Bimodal peaks and double responses were observed and the first peck or response was usually larger than the second. A disadvantage of the transducer was that pecking produced oscillation of the transducer at its resonant frequency. In spite of this deficiency, the waveform of the peck was easily recognizable.
PMCID: PMC1333769  PMID: 16811441
13.  Visually guided catching and tracking skills in pigeons: A preliminary analysis 
Research on reaching, tracking, and catching in the pigeon has been hampered by limitations of technology. A new system was developed in which the target was a small rectangle presented on a video display terminal and the pecking response was detected with touch technology. The target moved up and down vertically with sinusoidal velocity. A coincidence between the location of the pigeon's beak and the cursor produced reinforcement. The pigeon pecked ahead and behind the target, but most pecks occurred behind the target so the dominant tracking strategy was lagging. The pigeon was adept at “catching” the target at many locations throughout the trajectory. Transfer of motor learning was tested on probe trials during which the trajectory changed from vertical to horizontal. On transfer trials the pigeons' dominant pattern of pecking immediately shifted from vertical to horizontal. The motor skill displayed by the pigeons was flexible and adaptive, suggesting that the pigeons had learned to track the cursor.
PMCID: PMC1339189  PMID: 16812602
tracking; catching; reaching; stimulus control by movement; transfer of training; motor behavior; peck; pigeons
14.  Effects of relative reinforcer frequency on complex color detection 
Pigeons were trained under a discrete-trials detection procedure in which one of a set of color stimuli was presented on the center key and a single response turned off the stimulus and illuminated two side keys. Single responses to one or the other side key produced occasional reinforcers depending on the value of the color stimulus. In Experiment 1, one color-stimulus set comprised 559, 564, 569, and 574 nm, and right-key pecks were occasionally reinforced following presentations of members of this set. The other stimulus set comprised 579, 584, 589, and 594 nm, and left-key pecks were occasionally reinforced following presentations of members of this set. Across seven experimental conditions, the left/(left + right) relative reinforcer frequency was varied from .1 to .9. In Experiment 2, one stimulus set contained only one member, 574 nm, and right-key responses were occasionally reinforced following its presentation. Over 12 experimental conditions, two manipulations were carried out. First, the number of stimuli comprising the other stimulus set was increased from one (579 nm) to two (579 and 584 nm) to three (579, 584, and 589 nm) and to four (579, 584, 589, and 594 nm), and left-key responses were reinforced occasionally following center-key presentations of members of this set. Second, for each stimulus combination, the left/(left + right) relative reinforcer frequency was varied from .1 to .5 to .9 across three experimental conditions. The principal finding of Experiments 1 and 2 was that reinforcers and stimuli interacted in their effects on behavior. In Experiment 3, pairs of adjacent stimuli (5 nm apart) in the range 559 to 594 nm were presented in each experimental condition, and the left/(left + right) relative reinforcer frequency was held constant at .5. The data from all three experiments were analyzed according to a detection model describing performance in multiple-stimulus two-response procedures. This model provided independent measures of stimulus discriminability, contingency discriminability, and bias. The analysis showed that (a) consistent with the color-naming function, pigeons were better able to discriminate between higher nanometer values than lower nanometer values; (b) their ability to discriminate between the stimuli was independent of the number of wavelengths comprising each stimulus set; (c) they allocated delivered reinforcers very accurately to the previously emitted response; and (d) no consistent biases emerged.
PMCID: PMC1338923  PMID: 16812580
signal detection; multiple-stimulus control; stimulus discriminability; contingency discriminability; bias; key peck; pigeons
15.  Frequency of reinforcement as a determinant of extinction-induced aggression during errorless discrimination learning. 
Seven pigeons were trained to discriminate without errors between a green keylight and a dark key. The key-pecking response was reinforced in the presence of green, and extinction was in effect in the presence of the dark key. The opportunity to attack a restrained target pigeon was present only during extinction. Both variable-interval 30-sec and fixed-ratio 1 schedules of reinforcement during the positive stimulus induced a higher rate of attack during extinction than a variable-interval 5-min schedule. The highest rate of attack during extinction occurred during the first 20 sec after the positive stimulus terminated. Hence, the withdrawal of the positive condition, rather than the consequences of the pecking response during extinction, appears to be one of the primary factors responsible for attack between pigeons during extinction. Behavioral contrast, defined as a decrease in the rate of responding when the positive stimulus was presented alone, was obtained from the four birds that displayed the lowest overall rates of attack while the three birds with the highest attack rates did not display behavioral contrast. For the birds without contrast, components of the attack response during the positive stimulus presumably competed with and reduced the rate of pecking the key, thereby recluding behavioral contrast.
PMCID: PMC1333327  PMID: 1167899
16.  Key-peck durations under behavioral contrast and differential reinforcement 
Pigeons were maintained on a multiple schedule in which both components were variable-interval one-minute schedules. When they were switched to a condition in which one component was extinction, behavioral contrast was observed. The median durations of the key pecks in the unchanged component did not decrease in size. The results are incompatible with a theory of behavioral contrast which considers the added pecks to be short-duration responses. In a second experiment, pigeons were required to emit short-duration key pecks in one component of a multiple schedule, and long-duration pecks in the other. Two of three pigeons learned to emit responses appropriate to the requirements of the component in effect, suggesting that the duration of the key-peck response is sensitive to differential reinforcement.
PMCID: PMC1332994  PMID: 16812185
behavioral contrast; response duration; differential reinforcement; elicited responding; response topography as an operant; additivity theory; multiple schedules; key peck; pigeons
17.  A yoked-chamber comparison of concurrent and multiple schedules1 
Pigeons were exposed to alternative pairs of variable-interval schedules correlated with red and green lights on one key (the food key). In one experimental chamber, responses on a white key (the changeover key) changed the color of the food key and initiated a 2-sec changeover delay. Pigeons in a second chamber obtained food by pecking on a colored key whenever the pigeons in the first (concurrent) chamber had obtained food for a peck on that key color. There was no changeover key in the second (multiple) chamber: changeover responses in the first chamber alternated the schedules and colors in both chambers. The pigeons in both chambers emitted the same proportion of responses on each of the variable-interval schedules, and mastered discrimination reversals at the same rate. The pigeons differed only in their absolute response rates, which were greater under the concurrent schedules. In a second experiment, changes in key color occurred automatically, with different proportions of time allocated to the two variable-interval schedules. Matching of relative response frequency to relative reinforcement frequency was affected by the relative amounts of time in each component, by rate of changeovers, and by manipulations of the variable-interval scheduling.
PMCID: PMC1333980  PMID: 16811609
18.  Constraints on Letter-in-String Identification in Peripheral Vision: Effects of Number of Flankers and Deployment of Attention 
Effects of non-adjacent flanking elements on crowding of letter stimuli were examined in experiments manipulating the number of flanking elements and the deployment of spatial attention. To this end, identification accuracy of single letters was compared with identification of letter targets surrounded by two, four, or six flanking elements placed symmetrically left and right of the target. Target stimuli were presented left or right of a central fixation, and appeared either unilaterally or with an equivalent number of characters in the contralateral visual field (bilateral presentation). Experiment 1A tested letter targets with random letter flankers, and Experiments 1B and 2 tested letter targets with Xs as flanking stimuli. The results revealed a number of flankers effect that extended beyond standard two-flanker crowding. Flanker interference was stronger with random letter flankers compared with homogeneous Xs, and performance was systematically better under unilateral presentation conditions compared with bilateral presentation. Furthermore, the difference between the zero-flanker and two-flanker conditions was significantly greater under bilateral presentation, whereas the difference between two-flankers and four-flankers did not differ across unilateral and bilateral presentation. The complete pattern of results can be captured by the independent contributions of excessive feature integration and deployment of spatial attention to letter-in-string visibility.
PMCID: PMC3595509  PMID: 23494610
letter perception; crowding; non-adjacent flankers; number of flankers; spatial attention
19.  Independence of stimulus discriminability from absolute rate of reinforcement in a signal-detection procedure. 
Three experiments are reported in which two pigeons were trained to detect differences in stimulus duration under varying levels of absolute rate of reinforcement. Two red stimuli, differing in duration, were arranged probabilistically on the center key of a three-key chamber. On completion of the center-key duration, the center keylight was extinguished and the two side keys were illuminated white. Correct responses were left-key pecks following the shorter duration and right-key pecks following the longer duration. In Experiment 1, relative rate of reinforcement for correct responses was held constant and absolute rate of reinforcement was varied in seven conditions from continuous reinforcement to a variable-interval 90-second schedule. In Experiment 2, relative rate of reinforcement was manipulated across three different absolute rates of reinforcement (continuous reinforcement, variable-interval 15-second, and variable-interval 45-second). Stimulus discriminability was unaffected by changes in absolute or relative rates of reinforcement. Experiment 3 showed that discriminability was also unaffected by arranging the same consequences (three-second blackout) for unreinforced correct responses and errors.
PMCID: PMC1333153  PMID: 7097151
20.  Concurrent performances: reinforcement interaction and response independence1 
When a pigeon's pecks on two keys were reinforced concurrently by two independent variable-interval (VI) schedules, one for each key, the response rate on either key was given by the equation: R1=Kr1/(r1+r2)5/6, where R is response rate, r is reinforcement rate, and the subscripts 1 and 2 indicate keys 1 and 2. When the constant, K, was determined for a given pigeon in one schedule sequence, the equation predicted that pigeon's response rates in a second schedule sequence. The equation derived from two characteristics of the performance: the total response rate on the two keys was proportional to the one-sixth power of the total reinforcement rate provided by the two VI schedules; and, the pigeon matched the relative response rate on a key to the relative reinforcement rate for that key. The equation states that response rate on one key depends in part on reinforcement rate for the other key, but implies that it does not depend on response rate on the other key. This independence of response rates on the two keys was demonstrated by presenting a stimulus to the pigeon whenever one key's schedule programmed reinforcement. This maintained the reinforcement rate for that key, but reduced the response rate almost to zero. The response rate on the other key, nevertheless, continued to vary with reinforcement rates according to the equation.
PMCID: PMC1404302  PMID: 14019312
21.  Choice performance in several concurrent key-peck treadle-press reinforcement schedules1 
Five pigeons were exposed to several concurrent variable-interval food reinforcement schedules. For three subjects, one component of the schedule required a key-pecking response, the other a treadle-pressing response. For the other two subjects, both schedule components required treadle-pressing responses. The relative probability of reinforcement associated with the manipulanda was varied from 0 to 1.0 in 13 experimental conditions for the Key-Treadle subjects and nine conditions for the Treadle-Treadle subjects. The results indicated that the logarithms of relative time spent responding, and the logarithms of relative number of responses emitted on a manipulandum, approximated direct linear functions of logarithms of the relative frequencies of reinforcement associated with that manipulandum. No systematic bias in favor of time spent key pecking over time spent treadle pressing was apparent for the Key-Treadle subjects. All subjects exhibited undermatching, in that the ratios of time and response allocation at the alternatives systematically differed from the ratios of reinforcers obtained from the alternatives in the direction of indifference. Key pecking appeared to have no special link to food beyond treadle pressing or what would be expected on the basis of the reinforcement dependencies alone.
PMCID: PMC1332746  PMID: 16812046
concurrent schedules; matching contrast; treadle press; key peck; pigeons
22.  Absence of shock-elicited aggression in pigeons1 
Two pigeons that attacked a taxidermically prepared target pigeon during a schedule of positive reinforcement for key pecking, and two that did not, were shocked through implanted electrodes in the presence of the target. Shock intensities of 2 and 4 mA, durations of 0.1 and 1.3 sec, and frequencies of 2, 6, 20, and 35 per minute were delivered across 16 sessions with 180 shocks per session. No pigeon attacked the target; one pecked the shockplug on its back. The two pigeons that had not attacked during the positive reinforcement schedules were conditioned to peck the target for food reinforcement before another 16 sessions of shock. No attack was observed in these shock sessions. During subsequent positive reinforcement of key pecking, the target was attacked by the two pigeons that had originally attacked and by one that had not. Absence of shock-elicited attack in these pigeons may be related to the parameters of the experiment or may be yet another instance of the absence of shock-elicited attack in the class Aves. At least under the present conditions, it was not possible to predict the level of attack during electric shock from the level of attack during schedules of positive reinforcement for key pecking.
PMCID: PMC1333194  PMID: 4856078
23.  Contrast as seen in visual search reaction times. 
Three pigeons searched arrays of alphabetic letters displayed on computer monitors. On each trial, either an A or an E appeared, and the reaction time and accuracy with which the bird pecked at this target were measured. In each block of trials, each target (A or E) was displayed alone, or together with a number of distractor letters (2 or 18) that varied in their similarity to the target. During a baseline series of sessions, responses to the A and to the E each yielded food reinforcement on 10% of the trials. In the next series of sessions, reinforcement continued at 10% for A, but rose to 30% for E. In a final series, these reinforcement conditions were reversed. As expected, reaction times increased with target-distractor similarity and (for similar distractors) with the number of distractors. Increased reinforcement of E had no effect on reaction times to E, but produced a very consistent increase in reaction times to A; the average increase was constant across the various display conditions. Reversal of the differential reinforcement conditions reversed this contrast effect. Analysis of the reaction time distributions indicated that increased reinforcement to E decreased the momentary probability of response to A by a constant amount, regardless of display conditions. These results are discussed in relation to theories of contrast, memory, and of the search image.
PMCID: PMC1339175  PMID: 2584913
24.  Studies of operant and reflexive key pecks in the pigeon1 
The duration of pigeons' key pecks was studied in three experiments. Experiment I revealed that key pecks early in exposure to continuous reinforcement were of short duration, as were key pecks observed on an omission procedure in which pecks prevented food delivery. Key pecks later in exposure to continuous reinforcement, and those that occurred on positive automaintenance procedures, were of long duration. In Experiment II, pigeons were exposed to fixed-interval and fixed-ratio reinforcement schedules, and durations were recorded separately for each quarter of each interval or ratio. On fixed interval, durations were shorter in the first quarter of each interval than in subsequent quarters; on fixed ratio, durations were longer in the first quarter of the ratio than in subsequent quarters. These data parallel observations of concurrent operant responding and salivation in dogs. In Experiment III, pigeons were exposed to a discrete trial, differential-reinforcement-of-low-rate 6-sec schedule. Durations of responses in the first 2 sec of the trial were substantially shorter than those of responses that occurred later. The data from all three experiments support the view that the pigeon's “key peck” actually consists of two subclasses of peck, one reflexive and one operant.
PMCID: PMC1333594  PMID: 16811992
peck duration; continuous reinforcement; omission; fixed interval; fixed ratio; differential reinforcement of low rate; pigeon
25.  “Automaintenance”: the role of reinforcement1 
A key was illuminated on the average of every 30 sec for a duration of 6 sec and this was followed by food presentations. When key pecks in the presence of the light produced immediate access to grain (autoshaping procedure) pigeons were likely to peck. When pecks terminated the keylight but prevented access to grain (automaintenance procedure) pigeons were much less likely to peck. Seven of 12 pigeons failed to develop responding during the automaintenance procedure. Four of the five pigeons that responded during the automaintenance procedure were exposed to a procedure in which responses could not immediately terminate the light. Three of the four ceased to respond during optimal automaintenance conditions, suggesting that the response-dependent offset of the keylight had been reinforcing their pecking. Responding during the automaintenance procedure was eliminated for a fifth pigeon by eliminating the contiguity of light-offset and food-onset on those trials in which the pigeon did not peck. These results suggest that: (1) automaintenance (unlike autoshaping) is not an effective procedure for reliably generating responding; (2) responding that does occur during the automaintenance procedure is reinforced by the response-dependent offset of the keylight.
PMCID: PMC1333175  PMID: 16811725

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