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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.  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
4.  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
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
6.  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
7.  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
8.  Left-lateralized N170 Effects of Visual Expertise in Reading: Evidence from Japanese Syllabic and Logographic Scripts 
Journal of cognitive neuroscience  2008;20(10):1878-1891.
The N170 component of the event-related potential (ERP) reflects experience-dependent neural changes in several forms of visual expertise, including expertise for visual words. Readers skilled in writing systems that link characters to phonemes (i.e., alphabetic writing) typically produce a left-lateralized N170 to visual word forms. This study examined the N170 in three Japanese scripts that link characters to larger phonological units. Participants were monolingual English speakers (EL1) and native Japanese speakers (JL1) who were also proficient in English. ERPs were collected using a 129-channel array, as participants performed a series of experiments viewing words or novel control stimuli in a repetition detection task. The N170 was strongly left-lateralized for all three Japanese scripts (including logographic Kanji characters) in JL1 participants, but bilateral in EL1 participants viewing these same stimuli. This demonstrates that left-lateralization of the N170 is dependent on specific reading expertise and is not limited to alphabetic scripts. Additional contrasts within the moraic Katakana script revealed equivalent N170 responses in JL1 speakers for familiar Katakana words and for Kanji words transcribed into novel Katakana words, suggesting that the N170 expertise effect is driven by script familiarity rather than familiarity with particular visual word forms. Finally, for English words and novel symbol string stimuli, both EL1 and JL1 subjects produced equivalent responses for the novel symbols, and more left-lateralized N170 responses for the English words, indicating that such effects are not limited to the first language. Taken together, these cross-linguistic results suggest that similar neural processes underlie visual expertise for print in very different writing systems.
PMCID: PMC4416222  PMID: 18370600
9.  Effect of Orthographic Processes on Letter Identity and Letter-Position Encoding in Dyslexic Children 
The ability to identify letters and encode their position is a crucial step of the word recognition process. However and despite their word identification problem, the ability of dyslexic children to encode letter identity and letter-position within strings was not systematically investigated. This study aimed at filling this gap and further explored how letter identity and letter-position encoding is modulated by letter context in developmental dyslexia. For this purpose, a letter-string comparison task was administered to French dyslexic children and two chronological age (CA) and reading age (RA)-matched control groups. Children had to judge whether two successively and briefly presented four-letter strings were identical or different. Letter-position and letter identity were manipulated through the transposition (e.g., RTGM vs. RMGT) or substitution of two letters (e.g., TSHF vs. TGHD). Non-words, pseudo-words, and words were used as stimuli to investigate sub-lexical and lexical effects on letter encoding. Dyslexic children showed both substitution and transposition detection problems relative to CA-controls. A substitution advantage over transpositions was only found for words in dyslexic children whereas it extended to pseudo-words in RA-controls and to all type of items in CA-controls. Letters were better identified in the dyslexic group when belonging to orthographically familiar strings. Letter-position encoding was very impaired in dyslexic children who did not show any word context effect in contrast to CA-controls. Overall, the current findings point to a strong letter identity and letter-position encoding disorder in developmental dyslexia.
PMCID: PMC3356879  PMID: 22661961
letter-string processing; letter-position encoding; letter-identity encoding; letter transposition; letter substitution; reading acquisition; dyslexic children
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.  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
13.  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
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.  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
16.  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
17.  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
18.  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
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.  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
21.  “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
22.  Parameters affecting the maintenance of negatively reinforced key pecking1 
Three negative reinforcement experiments employing a key-peck response are described. In Experiment I, pigeons shocked on the average of twice per minute (imposed condition) could produce, by pecking a key, an alternate condition with correlated stimuli. Delayed shocks were added, across sessions, to the alternate condition until pecking stopped. Two of three pigeons continued to peck despite a 100% increase in shock frequency. In Experiment II, pigeons were shocked in the imposed condition four times per minute. The postresponse delay to shock was held constant by delivering, in the alternate condition, the next shock, or the next two, three, or four shocks from the imposed-condition shock schedule. All three subjects continued to peck with no change in delay to the first two postresponse shocks but with a 75% reduction in shock frequency. In Experiment III, a response produced an immediate shock followed by a shock-free period. Three of four subjects continued to respond despite reduced delay to shock. Delay-to-shock or shock-frequency reduction was sufficient to maintain key pecking, but neither was necessary. The conditions that negatively reinforce the pigeon's key peck were similar to conditions that negatively reinforce the rat's bar press.
PMCID: PMC1333624  PMID: 16812019
aversive control; avoidance; delayed shock; shock frequency; key pecking; pigeons
23.  Effects of search cost on foraging and feeding: a three-component chain analysis. 
An experiment determined whether pigeons minimize number of key pecks per food delivery and maintain their baseline intake of food while key pecking on a three-component chain schedule. Pigeons at either 80% or 100% body weight obtained all their food during baseline and contingency sessions. During baseline sessions, pecks on the left and center keys had no consequences; each peck on the right key activated the feeder. During contingency sessions, pigeons key pecked on a three-component chain schedule simulating components of a foraging chain. In the search component either 3, 9 or 15 key pecks (varied parametrically across blocks of sessions) on the left key produced a stimulus on the middle key, indicating an encounter with either the low-cost prey (3 key pecks) or an equally probable high-cost prey (21 key pecks). In the procurement component the pigeon pecked either: (a) the left key once, thus returning to the search component, or (b) the middle key either 3 or 21 times, which activated the right response key. In the handling component one peck on the right key operated the feeder. The pigeons always procured the low-cost prey and minimized the number of key pecks per hopper by procuring the high-cost prey when the search-cost ratio was high (15 key pecks) but not when it was low (3 key pecks). All pigeons maintained their baselines of eating during contingency sessions by key pecking more frequently and eating more efficiently. The 80% body-weight birds produced higher overall rates of key pecking and eating. These results have implications for ecological theories of optimal foraging and for psychological theories of learned performance.
PMCID: PMC1348078  PMID: 6502067
24.  Decision rules and signal detectability in a reinforcement-density discrimination 
Two probabilistic schedules of reinforcement, one richer in reinforcement, the other leaner, were overlapping stimuli to be discriminated in a choice situation. One of two schedules was in effect for 12 seconds. Then, during a 6-second choice period, the first left-key peck was reinforced if the richer schedule had been in effect, and the first right-key peck was reinforced if the leaner schedule had been in effect. The two schedule stimuli may be viewed as two binomial distributions of the number of reinforcement opportunities. Each schedule yielded different frequencies of 16 substimuli. Each substimulus had a particular type of outcome pattern for the 12 seconds during which a schedule was in effect, and consisted of four consecutive light-cued 3-second T-cycles, each having 0 or 1 reinforced center-key pecks. Substimuli therefore contained 0 to 4 reinforcers. On any 3-second cycle, the first center-key peck darkened that key and was reinforced with probability .75 or .25 in the richer or leaner schedules, respectively. In terms of the theory of signal detection, detectability neared the maximum possible d′ for all four pigeons. Left-key peck probability increased when number of reinforcers in a substimulus increased, when these occurred closer to choice, or when pellets were larger for correct left-key pecks than for correct right-key pecks. Averaged over different temporal patterns of reinforcement in a substimulus, substimuli with the same number of reinforcers produced choice probabilities that matched relative expected payoff rather than maximized one alternative.
PMCID: PMC1332883  PMID: 16812141
discrimination; decision rules; reinforcement; memory; matching; maximizing; signal detection; T-schedule; key peck; pigeon
25.  Yoked variable-ratio and variable-interval responding in pigeons1 
Pigeons' key pecks were maintained by variable-ratio or variable-interval schedules of food reinforcement. For pairs of pigeons in one group, variable-ratio reinforcement was arranged for one pigeon's pecks; for the second pigeon, reinforcement was arranged according to a variable-interval schedule yoked to the interreinforcement times produced by the first pigeon. For pairs of pigeons in another group, variable-interval reinforcement was arranged for one pigeon's pecks; for the second pigeon, reinforcement was arranged according to a variable-ratio schedule yoked to the interreinforcement responses produced by the first pigeon. For each pair, the yoking procedure was maintained for four or five consecutive sessions of 50 reinforcements each. In more than three-quarters of the pairs, variable-ratio response rates were higher than variable-interval rates within two sessions; in all cases, the rate difference developed within four sessions.
PMCID: PMC1333627  PMID: 16812022
variable-interval schedule; variable-ratio schedule; yoked schedules; time per reinforcement; responses per reinforcement; shaping; undergraduate laboratory course; key pecks; pigeons

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