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.
word reading; ERPs; visual cortex; visual orthography
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.
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.
Morphology; Letter Transposition; Hebrew; Masked-Priming
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.
behavioral contrast; response duration; differential reinforcement; elicited responding; response topography as an operant; additivity theory; multiple schedules; key peck; pigeons
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.
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.
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.
short-term memory; delayed stimulus control; relative recency; behavioral patterns; pigeons
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.
pigeon; autoshaping; conditional discrimination; open field; touchscreen; simultaneous discrimination
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.
variable-interval schedule; variable-ratio schedule; yoked schedules; time per reinforcement; responses per reinforcement; shaping; undergraduate laboratory course; key pecks; pigeons
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.
visual search; attention; priming; contrast; reaction time; reinforcement probability; reinforcement duration; peck; pigeon
The duration of the pigeon's key peck was differentially reinforced in either a trials or a free-operant procedure. Mean emitted peck duration was a power function of the duration required for food delivery to occur. The exponents of the power function differed considerably from those observed in earlier research involving longer duration responses in pigeons and other species. The coefficients of variation also did not correspond with those of the earlier research on other responses, nor did consideration of the durations actually reinforced resolve the differences. Duration was neither a function of response rate nor of intermittency of reinforcement. Key-peck duration was changed in an orderly way by differential reinforcement. However, it appeared to be more strongly determined by its duration in the absence of differential reinforcement than were longer duration responses.
temporal psychophysics; key-peck duration; differential reinforcement; base duration; key peck; pigeons
In a two-key pigeon chamber, variable-interval reinforcement was scheduled for a specified number of pecks, emitted either on a single key or in a particular sequence on the two keys. Although the distribution of pecks between the two keys was affected by whether pecks were required on one or on both keys, the total pecks emitted was not; the change from a one-key to a two-key requirement simply moved some pecks from one key to the other. Thus, each peck preceding the one that produced the reinforcer contributed independently to the subsequent rate of responding; the contribution of a particular peck in the sequence was determined by the time between its emission and the delivery of the reinforcer (delay of reinforcement), and was identified by the proportion of pecks moved from one key to the other when the response requirement at that point in the sequence was moved from one key to the other.
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.
tracking; catching; reaching; stimulus control by movement; transfer of training; motor behavior; peck; pigeons
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.
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.
Variations of the symbolic delayed-matching-to-sample procedure were used to study a pigeon's memory for a small number of pecks. In the first experiment a choice of a left or right sidekey after a delay or retention interval was reinforced if a bird had not pecked at all or had pecked exactly once, before the delay, respectively. In the second experiment a choice of a red or green sidekey, regardless of its position, was reinforced if a bird had not pecked at all or had pecked exactly twice, respectively. In the first experiment a bird could orient toward the correct choice during the delay, whereas it could not in the second experiment. In a third experiment a feature-probing method was used to study a pigeon's memory for a number of pecks in the context of certain other pecks. The results showed that a pigeon can remember a small number of pecks for one-half to one minute or more and that the percent correct is a decreasing function of the log retention interval. When a second number of pecks is different from the first number, memory for the first number lasts only a few seconds. When a second number is the same, memory lasts considerably longer. The more recent number of pecks is remembered better. The results are interpreted in terms of a theory which holds that a reinforcer, in general, may act on a subjects' memory for recent behavior to generate patterns of behavior.
short-term memory; symbolic delayed-matching-to-sample; feature probing; bias; memory strength; behavioral patterns; key pecks; pigeons
Pigeons' responses to a uniformly illuminated response key were either reinforced on a variable-interval one-minute schedule of reinforcement or extinguished for one-minute periods. When 1.5 second signals were presented at the beginning of each component, so as to differentially predict reinforcement, the pigeons pecked at the signals, at rates higher than rates during the remainder of the component. When the brief signals were not differentially predictive of reinforcement, pecking in their presence decreased to near zero levels. Similar results were obtained with signals based upon colors and upon line orientations. Changes in rates of (unreinforced) pecking occurred during the signal whether pigeons responded differentially during the remainder of the component or not. Experiment II demonstrated that the presence of the signal correlated with extinction was not necessary for pecking to develop at the signal which preceded the component in which responding was intermittently reinforced. The experiments demonstrated a clear dissociation of respondent control from operant control of a response. In addition, operant behavior was shown to be relatively insensitive to differing rates of reinforcement, as compared to the sensitivity of respondent behavior to differing rates of reinforcement produced by the very same operant behavior.
elicited and operant responding; stimulus control; trace stimuli; mixed schedules; key peck; pigeons
Pigeons were trained to peck at red or green keys presented simultaneously in discrete trials. In one experiment, reinforcements were arranged by concurrent variable-interval schedules. The proportion of responses to green approximately matched the proportion of reinforcements produced by pecking green. Detailed analysis of responding revealed a systematic decrease in the probability of switching from green to red within sequences of trials after reinforcement. This trend corresponded to sequential changes in the relative frequency of reinforcement, and not to sequential changes in probability of reinforcement. In a second experiment, reinforcements were scheduled probabilistically every seventh trial. Even though there were no contingencies on pecking during the first six post-reinforcement trials, choices of green on the first response after reinforcement matched the proportion of reinforcements for pecking green. These results extend the generality of overall matching under concurrent reinforcement.
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.
Pigeons were trained on a modified three-key matching-to-sample procedure, in which only one comparison key (rather than two) was lighted after an observing response to the center-key standard. Pecks on keys of matching comparison hues were reinforced. When non-matching hues appeared as the initially lighted comparisons, the nonmatching hue terminated and the matching hue appeared on the other side key only if the pigeon did not peck the nonmatching comparison for 4.8 sec. Pecks to the nonmatching hue reset the 4.8-sec delay interval. Three hues were used during acquisition. During transfer tests, two novel hues were substituted individually or together for one or two of the training hues. Latencies to the novel side-key hue were shortest when a novel matching hue appeared as the standard on the center key, and were essentially identical to baseline matching latencies. In contrast, when a novel hue appeared as either a standard or comparison in a nonmatching combination, latencies increased with increasing separation between the noevel hue and the nonmatching hue. These transfer data demonstrate the concept of hue matching.
Pigeons emitted almost exclusively short-duration key pecks (shorter than 20 msec) when on negative automaintenance procedures, in which pecks prevented reinforcement. Peck durations under fixed-interval and fixed-ratio reinforcement schedules were generally two to five times longer than pecks under a negative automaintenance schedule. However, initial key pecks were of short duration, independent of procedure. The frequency of short-duration pecks was insensitive to differential reinforcement, while the frequency of long-duration pecks was sensitive to differential reinforcement. It is proposed that short-duration pecks arise from the pigeon's normal feeding pattern and are directly enhanced by food presentation, while long-duration pecks are controlled by the contingent effects of food presentation. The implications of the existence of two classes of pecks for the functional definition of operants and the separation of phylogenetic and ontogenetic sources of control of key pecking are discussed.
In two discrete-trial delayed-detection experiments, six pigeons were trained on dependent concurrent variable-interval schedules. Pecking a red side key was reinforced when the brighter of two white lights (S1) had been presented on the center key, and pecking a green side key was reinforced when the duller of two white lights (S2) had been presented on the center key. Incorrect responses were red side-key pecks following S2 presentations and green side-key pecks following S1 presentations; these resulted in three-second blackouts. In Experiment 1, the time between presentation of S1 or S2 on the center key and the onset of the red and green side keys was varied nonsystematically from 0.06 seconds to 19.69 seconds across experimental conditions. Stimulus discriminability decreased as the stimulus-choice delay increased. A rectangular-hyperbolic function better described this decrease in discriminability over time than did a negative-exponential function. In Experiment 2, at each of three stimulus-choice delays (0.06, 3.85, and 10.36 seconds), relative reinforcer frequency for correct responses to the red and green side keys was varied by changing the values of the dependent concurrent variable-interval schedules. The sensitivity of choice to relative reinforcer frequency was independent of the decrease in stimulus discriminability with increasing stimulus-choice delay.
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.
aversive control; avoidance; delayed shock; shock frequency; key pecking; pigeons
The results of a number of recent studies suggest that acquisitions of autoshaped key pecking in pigeons is affected by the similarity of the grain-hopper stimulus and response-key stimulus. In Experiment 1 this hypothesis was tested by training independent groups of pigeons to key peck under six different hopper-stimulus and key-stimulus similarity conditions, and three procedures containing either immediate reinforcement, variable delay of reinforcement, or omission of reinforcement for key pecking. Number of trials to acquisition was found to be related to the similarity variable. Maintained responding was affected by the response-reinforcer contingency. This effect was found both within and between subjects. Under two of the contingencies (automaintenance and omission), maintained responding continued to be affected by the similarity of the hopper stimulus and key stimulus. In Experiment 2 pigeons were given omission training with a hopper light on or off. Both acquisition and maintenance of key pecking were facilitated by the presence of the hopper light. The present findings suggest that much of the responding reported in various automatic shaping and training procedures may reflect the effects of key stimulus/food stimulus similarity.
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.