Daniel J. Acheson

Increased alpha-band power during the retention of shapes and shape-location associations in visual short-term memory

Studies exploring the role of neural oscillations in cognition have revealed sustained increases in alpha-band (~8–14 Hz) power during the delay period of delayed-recognition short-term memory tasks. These increases have been proposed to reflect the inhibition, for example, of cortical areas representing task-irrelevant information, or of potentially interfering representations from previous trials. Another possibility, however, is that elevated delay-period alpha-band power (DPABP) reflects the selection and maintenance of information, rather than, or in addition to, the inhibition of task-irrelevant information. In the present study, we explored these possibilities using a delayed-recognition paradigm in which the presence and task relevance of shape information was systematically manipulated across trial blocks and electroencephalographic was used to measure alpha-band power. In the first trial block, participants remembered locations marked by identical black circles. The second block featured the same instructions, but locations were marked by unique shapes. The third block featured the same stimulus presentation as the second, but with pretrial instructions indicating, on a trial-by-trial basis, whether memory for shape or location was required, the other dimension being irrelevant. In the final block, participants remembered the unique pairing of shape and location for each stimulus. Results revealed minimal DPABP in each of the location-memory conditions, whether locations were marked with identical circles or with unique task-irrelevant shapes. In contrast, alpha-band power increases were observed in both the shape-memory condition, in which location was task irrelevant, and in the critical final condition, in which both shape and location were task relevant. These results provide support for the proposal that alpha-band oscillations reflect the retention of shape information and/or shape–location associations in short-term memory.

Stimulating the brain's language network: Syntactic ambiguity resolution after tms to the inferior frontal gyrus and middle temporal gyrus

The posterior middle temporal gyrus (MTG) and inferior frontal gyrus (IFG) are two critical nodes of the brains language network. Previous neuroimaging evidence has supported a dissociation in language comprehension in which parts of the MTG are involved in the retrieval of lexical syntactic information and the IFG in unification operations that maintain, select, and integrate multiple sources of information over time. In the present investigation, we tested for causal evidence of this dissociation by modulating activity in IFG and MTG using an offline TMS procedure: continuous theta-burst stimulation. Lexical–syntactic retrieval was manipulated by using sentences with and without a temporarily word-class (noun/verb) ambiguity (e.g., run). In one group of participants, TMS was applied to the IFG and MTG, and in a control group, no TMS was applied. Eye movements were recorded and quantified at two critical sentence regions: a temporarily ambiguous region and a disambiguating region. Results show that stimulation of the IFG led to a modulation of the ambiguity effect (ambiguous–unambiguous) at the disambiguating sentence region in three measures: first fixation durations, total reading times, and regressive eye movements into the region. Both IFG and MTG stimulation modulated the ambiguity effect for total reading times in the temporarily ambiguous sentence region relative to the control group. The current results demonstrate that an offline repetitive TMS protocol can have influences at a different point in time during online processing and provide causal evidence for IFG involvement in unification operations during sentence comprehension.

Attention for speaking: domain-general control from the anterior cingulate cortex in spoken word production

Accumulating evidence suggests that some degree of attentional control is required to regulate and monitor processes underlying speaking. Although progress has been made in delineating the neural substrates of the core language processes involved in speaking, substrates associated with regulatory and monitoring processes have remained relatively underspecified. We report the results of an fMRI study examining the neural substrates related to performance in three attention-demanding tasks varying in the amount of linguistic processing: vocal picture naming while ignoring distractors (picture-word interference, PWI); vocal color naming while ignoring distractors (Stroop); and manual object discrimination while ignoring spatial position (Simon task). All three tasks had congruent and incongruent stimuli, while PWI and Stroop also had neutral stimuli. Analyses focusing on common activation across tasks identified a portion of the dorsal anterior cingulate cortex (ACC) that was active in incongruent trials for all three tasks, suggesting that this region subserves a domain-general attentional control function. In the language tasks, this area showed increased activity for incongruent relative to congruent stimuli, consistent with the involvement of domain-general mechanisms of attentional control in word production. The two language tasks also showed activity in anterior-superior temporal gyrus (STG). Activity increased for neutral PWI stimuli (picture and word did not share the same semantic category) relative to incongruent (categorically related) and congruent stimuli. This finding is consistent with the involvement of language-specific areas in word production, possibly related to retrieval of lexical-semantic information from memory. The current results thus suggest that in addition to engaging language-specific areas for core linguistic processes, speaking also engages the ACC, a region that is likely implementing domain-general attentional control.

Electrophysiological evidence that inhibition supports lexical selection in picture naming.

We investigated the neural basis of inhibitory control during lexical selection. Participants overtly named pictures while response times (RTs) and event-related brain potentials (ERPs) were recorded. The difficulty of lexical selection was manipulated by using object and action pictures with high name agreement (few response candidates) versus low name agreement (many response candidates). To assess the involvement of inhibition, we conducted delta plot analyses of naming RTs and examined the N2 component of the ERP. We found longer mean naming RTs and a larger N2 amplitude in the low relative to the high name agreement condition. For action naming we found a negative correlation between the slopes of the slowest delta segment and the difference in N2 amplitude between the low and high name agreement conditions. The converging behavioral and electrophysiological evidence suggests that selective inhibition is engaged to reduce competition during lexical selection in picture naming.

Conflict monitoring in speech production: Physiological evidence from bilingual picture naming

Self-monitoring in production is critical to correct performance, and recent accounts suggest that such monitoring may occur via the detection of response conflict. The error-related negativity (ERN) is a response-locked event-related potential (ERP) that is sensitive to response conflict. The present study examines whether response conflict is detected in production by exploring a situation where multiple outputs are activated: the bilingual naming of form-related equivalents (i.e. cognates). ERPs were recorded while German-Dutch bilinguals named pictures in their first and second languages. Although cognates were named faster than non-cognates, response conflict was evident in the form of a larger ERN-like response for cognates and adaptation effects on naming, as the magnitude of cognate facilitation was smaller following the naming of cognates. Given that signals of response conflict are present during correct naming, the present results suggest that such conflict may serve as a reliable signal for monitoring in speech production.

Twisting tongues to test for conflict-monitoring in speech production.

A number of recent studies have hypothesized that monitoring in speech production may occur via domain-general mechanisms responsible for the detection of response conflict. Outside of language, two ERP components have consistently been elicited in conflict-inducing tasks (e.g., the flanker task): the stimulus-locked N2 on correct trials, and the response-locked error-related negativity (ERN). The present investigation used these electrophysiological markers to test whether a common response conflict monitor is responsible for monitoring in speech and non-speech tasks. Electroencephalography (EEG) was recorded while participants performed a tongue twister (TT) task and a manual version of the flanker task. In the TT task, people rapidly read sequences of four nonwords arranged in TT and non-TT patterns three times. In the flanker task, people responded with a left/right button press to a center-facing arrow, and conflict was manipulated by the congruency of the flanking arrows. Behavioral results showed typical effects of both tasks, with increased error rates and slower speech onset times for TT relative to non-TT trials and for incongruent relative to congruent flanker trials. In the flanker task, stimulus-locked EEG analyses replicated previous results, with a larger N2 for incongruent relative to congruent trials, and a response-locked ERN. In the TT task, stimulus-locked analyses revealed broad, frontally-distributed differences beginning around 50 ms and lasting until just before speech initiation, with TT trials more negative than non-TT trials; response-locked analyses revealed an ERN. Correlation across these measures showed some correlations within a task, but little evidence of systematic cross-task correlation. Although the present results do not speak against conflict signals from the production system serving as cues to self-monitoring, they are not consistent with signatures of response conflict being mediated by a single, domain-general conflict monitor.

Hierarchy is Detrimental for Human Cooperation

Studies of animal behavior consistently demonstrate that the social environment impacts cooperation, yet the effect of social dynamics has been largely excluded from studies of human cooperation. Here, we introduce a novel approach inspired by nonhuman primate research to address how social hierarchies impact human cooperation. Participants competed to earn hierarchy positions and then could cooperate with another individual in the hierarchy by investing in a common effort. Cooperation was achieved if the combined investments exceeded a threshold, and the higher ranked individual distributed the spoils unless control was contested by the partner. Compared to a condition lacking hierarchy, cooperation declined in the presence of a hierarchy due to a decrease in investment by lower ranked individuals. Furthermore, hierarchy was detrimental to cooperation regardless of whether it was earned or arbitrary. These findings mirror results from nonhuman primates and demonstrate that hierarchies are detrimental to cooperation. However, these results deviate from nonhuman primate findings by demonstrating that human behavior is responsive to changing hierarchical structures and suggests partnership dynamics that may improve cooperation. This work introduces a controlled way to investigate the social influences on human behavior, and demonstrates the evolutionary continuity of human behavior with other primate species.

Effects of semantic integration on subject–verb agreement: evidence from Dutch

The generation of subject–verb agreement is a central component of grammatical encoding. It is sensitive to conceptual and grammatical influences, but the interplay between these factors is still not fully understood. We investigate how semantic integration of the subject noun phrase (‘the secretary of/with the governor’) and the Local Noun Number (‘the secretary with the governor/governors’) affect the ease of selecting the verb form. Two hypotheses are assessed: according to the notional hypothesis, integration encourages the assignment of the singular notional number to the noun phrase and facilitates the choice of the singular verb form. According to the lexical interference hypothesis, integration strengthens the competition between nouns within the subject phrase, making it harder to select the verb form when the nouns mismatch in number. In two experiments, adult speakers of Dutch completed spoken preambles (Experiment 1) or selected appropriate verb forms (Experiment 2). Results showed facilitatory effects of semantic integration (fewer errors and faster responses with increasing integration). These effects did not interact with the effects of the Local Noun Number (slower response times and higher error rates for mismatching than for matching noun numbers). The findings thus support the notional hypothesis and a model of agreement where conceptual and lexical factors independently contribute to the determination of the number of the subject noun phrase and, ultimately, the verb.

Keeping it simple: studying grammatical encoding with lexically reduced item sets

Compared to the large body of work on lexical access, little research has been done on grammatical encoding in language production. An exception is the generation of subject-verb agreement. Here, two key findings have been reported: (1) speakers make more agreement errors when the head and local noun of a phrase mismatch in number than when they match [e.g., the key to the cabinet(s)]; and (2) this attraction effect is asymmetric, with stronger attraction for singular than for plural head nouns. Although these findings are robust, the cognitive processes leading to agreement errors and their significance for the generation of correct agreement are not fully understood. We propose that future studies of agreement, and grammatical encoding in general, may benefit from using paradigms that tightly control the variability of the lexical content of the material. We report two experiments illustrating this approach. In both of them, the experimental items featured combinations of four nouns, four color adjectives, and two prepositions. In Experiment 1, native speakers of Dutch described pictures in sentences such as the circle next to the stars is blue. In Experiment 2, they carried out a forced-choice task, where they read subject noun phrases (e.g., the circle next to the stars) and selected the correct verb-phrase (is blue or are blue) with a button press. Both experiments showed an attraction effect, with more errors after subject phrases with mismatching, compared to matching head and local nouns. This effect was stronger for singular than plural heads, replicating the attraction asymmetry. In contrast, the response times recorded in Experiment 2 showed similar attraction effects for singular and plural head nouns. These results demonstrate that critical agreement phenomena can be elicited reliably in lexically reduced contexts. We discuss the theoretical implications of the findings and the potential and limitations of studies using lexically simple materials.