Japanese written pseudowords can be conditioned to Japanese spoken words with positive, negative, and active emotions.

This study aimed to examine whether Japanese participants condition spoken words' meanings to written pseudowords. In Survey 1, we selected spoken words associated with negative (α = .91) and positive (α = .79) features for Experiment 1 and passive (α = .90) and active (α = .80) features for Experiment 2. In Experiment 1, participants evaluated four written pseudowords' emotional valence using a 7-point semantic differential scale (1: negative; 7: positive) before and after conditioning spoken words with negative, neutral, or positive features to each pseudoword. In the conditioning phase, participants read each pseudoword, listened to a spoken word, and verbally repeated each spoken word. The results showed that a pseudoword was conditioned to spoken words with positive and negative features. In Experiment 2, participants evaluated four pseudowords' activeness using a 7-point semantic differential scale (1: passive; 7: active) before and after conditioning spoken words of passive, neutral, and active features to each written pseudoword. In the conditioning phase, the participants read each written pseudoword, listened to a spoken word, and repeated the spoken word. The results showed that the activeness evaluations were more increased for pseudowords conditioned to spoken words of active and neutral features after conditioning than before conditioning but were unchanged for a pseudoword conditioned to those with passive features before and after conditioning. Additonally, Survey 2's results showed that although the positiveness and activeness responses of the words used in Experiments 1 and 2 were controlled well, the lack of significant differences among positiveness responses of words may influence the evaluative conditioning in Experiment 2. That is, when participants condition passive (low arousal) words' activeness (arousal) ratings to those of pseudowords, words' positiveness (valence) ratings would be important in the evaluative conditioning. Our findings suggest that participants can condition spoken word meanings of preference and activeness to those of written pseudowords. It also indicates that linguistically evaluative conditioning's effects are robust in a non-alphabetic language.

Intra- and inter-hemispheric network dynamics supporting object recognition and speech production.

We built normative brain atlases that animate millisecond-scale intra- and inter-hemispheric white matter-level connectivity dynamics supporting object recognition and speech production. We quantified electrocorticographic modulations during three naming tasks using event-related high-gamma activity from 1,114 nonepileptogenic intracranial electrodes (i.e., non-lesional areas unaffected by epileptiform discharges). Using this electrocorticography data, we visualized functional connectivity modulations defined as significant naming-related high-gamma modulations occurring simultaneously at two sites connected by direct white matter streamlines on diffusion-weighted imaging tractography. Immediately after stimulus onset, intra- and inter-hemispheric functional connectivity enhancements were confined mainly across modality-specific perceptual regions. During response preparation, left intra-hemispheric connectivity enhancements propagated in a posterior-to-anterior direction, involving the left precentral and prefrontal areas. After overt response onset, inter- and intra-hemispheric connectivity enhancements mainly encompassed precentral, postcentral, and superior-temporal (STG) gyri. We found task-specific connectivity enhancements during response preparation as follows. Picture naming enhanced activity along the left arcuate fasciculus between the inferior-temporal and precentral/posterior inferior-frontal (pIFG) gyri. Nonspeech environmental sound naming augmented functional connectivity via the left inferior longitudinal and fronto-occipital fasciculi between the medial-occipital and STG/pIFG. Auditory descriptive naming task enhanced usage of the left frontal U-fibers, involving the middle-frontal gyrus. Taken together, the commonly observed network enhancements include inter-hemispheric connectivity optimizing perceptual processing exerted in each hemisphere, left intra-hemispheric connectivity supporting semantic and lexical processing, and inter-hemispheric connectivity for symmetric oral movements during overt speech. Our atlases improve the currently available models of object recognition and speech production by adding neural dynamics via direct intra- and inter-hemispheric white matter tracts.

Bizarreness and Typicality Effects of Color on Object Recognition Memory.

The bizarreness effect on object recognition is a strong phenomenon, but its influence has been inconsistent for bizarre object color. In this study, we manipulated three factors in separate experiments to determine whether a color bizarreness effect on object recognition memory would occur and, if not, why. Participants first saw (i.e., learned) object pictures that were either bizarrely or typically colored; they then completed a recognition memory test. In three experiments, we then manipulated (a) degree of color bizarreness (Experiment 1), (b) the orientation task (Experiment 2), and (c) additional demands for object identification (Experiment 3). In Experiment 1, we provided 49 undergraduate participants with object pictures whose colors were typical, moderately atypical, or bizarre and found no color bizarreness effect on recognition memory even for extremely bizarre colors. In Experiment 2, we manipulated the orientation task in that 28 young adult participants expressed their preferences for the pictures on a three-point scale while another 28 participants judged how natural the pictures were. Each orientation task group better recognized typically-colored rather than bizarrely-colored objects (typicality effect). In Experiment 3, we asked 27 young adults to identify the objects during the learning phase to ensure that they paid attention to the objects' bizarre colors; recognition memory was then unaffected by either color bizarreness or typicality. Thus, despite a general bizarreness effect in recognition memory, bizarre colors are less likely to influence object recognition memory.

Associative Learning of New Word Forms in a First Language (L1) and Haptic Referents in a Single-Day Experiment.

This study focused on the associative learning of new word forms in the first language and haptic stimuli. In this study, healthy Japanese participants performed three-step tasks. First, participants made nine subjective evaluations of haptic stimuli using five-point semantic differential scales (e.g., regarding stickiness, scored from 1 (not sticky) to 5 (sticky)). Second, the participants carried out learning and recognition tasks for associative pairs of new (meaningless) word forms in their first language (Japanese) and haptic stimulus (H condition), and performed learning and recognition tasks for new (meaningless) word forms only (W condition). The order of conditions was counterbalanced among participants. Third, participants performed free recall tasks. The results of the recognition tasks showed that the proportions and response times of the W condition were better and faster, respectively, than those of the H condition. Furthermore, preference of haptic features negatively correlated with free recall scores of the H condition; however, there was no significant difference between the free recall scores of the H and W conditions. Our results suggest that new word forms were learned better than associative pairs of new word forms and haptic stimuli in a single day of learning. Furthermore, the free recall performance of word forms associated with haptic features could also be affected by their subjective evaluation (preference).

Multisensory Connections of Novel Linguistic Stimuli in Japanese as a Native Language and Referential Tastes.

Previous findings have shown essential connections between linguistic and gustatory stimuli for people with autism or lexical gustatory synesthesia. We examined the associative learning of novel linguistic forms in Japanese as a native language and tastes (candies and chocolates) for healthy people. Healthy subjects performed four phases: (a) evaluation phase of gustatory features; (b) learning phases of novel linguistic form and gustatory stimulus pairs (G) or novel word forms (W); (c) recognition memory phases linked with G and W; and (d) free recall phase for G and W. In the recognition memory phases, the performance scores of W were higher than those of G, while there was no significant difference between response times of G and W. Additionally, no difference between recall performances in G and W was also shown. A subjective evaluation of gustatory features (sweetness) negatively correlated with the recall score for linguistic forms connected to the gustatory feature, whereas the accuracy rates of the recognition memory phase in G positively correlated with those of the free recall phase in G. Although learning of novel linguistic forms is more efficient than learning of the relationships between novel linguistic forms and tastes, gustatory features influence the free recall performances of linguistic forms linked with the tastes. These results may contribute to future applications to word learning not just for patients, but also healthy people.

Vowel Length Expands Perceptual and Emotional Evaluations in Written Japanese Sound-Symbolic Words.

In this study, we examined whether vowel length affected the perceptual and emotional evaluations of Japanese sound-symbolic words. The perceptual and emotional features of Japanese sound-symbolic words, which included short and long vowels, were evaluated by 209 native Japanese speakers. The results showed that subjective evaluations of familiarity, visual imageability, auditory imageability, tactile imageability, emotional valence, arousal, and length were significantly higher for sound-symbolic words with long vowels compared to those with short vowels. Additionally, a subjective evaluation of speed was significantly higher for written Japanese sound-symbolic words with short vowels than for those with long vowels. The current findings suggest that vowel length in written Japanese sound-symbolic words increases the perceptually and emotionally subjective evaluations of Japanese sound-symbolic words.

Your verbal questions beginning with 'what' will rapidly deactivate the left prefrontal cortex of listeners.

The left prefrontal cortex is essential for verbal communication. It remains uncertain at what timing, to what extent, and what type of phrase initiates left-hemispheric dominant prefrontal activation during comprehension of spoken sentences. We clarified this issue by measuring event-related high-gamma activity during a task to respond to three-phrase questions configured in different orders. Questions beginning with a wh-interrogative deactivated the left posterior prefrontal cortex right after the 1st phrase offset and the anterior prefrontal cortex after the 2nd phrase offset. Left prefrontal high-gamma activity augmented subsequently and maximized around the 3rd phrase offset. Conversely, questions starting with a concrete phrase deactivated the right orbitofrontal region and then activated the left posterior prefrontal cortex after the 1st phrase offset. Regardless of sentence types, high-gamma activity emerged earlier, by one phrase, in the left posterior prefrontal than anterior prefrontal region. Sentences beginning with a wh-interrogative may initially deactivate the left prefrontal cortex to prioritize the bottom-up processing of upcoming auditory information. A concrete phrase may obliterate the inhibitory function of the right orbitofrontal region and facilitate top-down lexical prediction by the left prefrontal cortex. The left anterior prefrontal regions may be recruited for semantic integration of multiple concrete phrases.

The Relationships Between Initial Consonants in Japanese Sound Symbolic Words and Familiarity, Multi-Sensory Imageability, Emotional Valence, and Arousal.

Sound symbolic words consist of inevitable associations between sounds and meanings. We aimed to identify differences in familiarity, visual imageability, auditory imageability, tactile imageability, emotional valence, and arousal between Japanese sound symbolic words with voiced initial consonants (VCs; dakuon in Japanese; e.g., biribiri) and Japanese sound symbolic words with semi-voiced initial consonants (SVCs; handakuon in Japanese; e.g., piripiri), and between VCs (e.g., daradara) and Japanese sound symbolic words with voiceless initial consonants (VLCs; seion in Japanese; e.g., taratara). First, auditory imageability and arousal were significantly higher in VCs than SVCs, whereas familiarity, tactile imageability, and positive emotion (emotional valence) were significantly higher in SVCs than VCs. Second, visual imageability was higher in VCs than VLCs, while familiarity and positive emotion were higher in VLCs than VCs. Initial consonants in Japanese sound symbolic words could be associated with specific subjective evaluations such as familiarity, visual imageability, auditory imageability, tactile imageability, emotional valence, and arousal.

Semantics Based on the Physical Characteristics of Facial Expressions Used to Produce Japanese Vowels.

Previous studies have reported that verbal sounds are associated-non-arbitrarily-with specific meanings (e.g., sound symbolism and onomatopoeia), including visual forms of information such as facial expressions; however, it remains unclear how mouth shapes used to utter each vowel create our semantic impressions. We asked 81 Japanese participants to evaluate mouth shapes associated with five Japanese vowels by using 10 five-item semantic differential scales. The results reveal that the physical characteristics of the facial expressions (mouth shapes) induced specific evaluations. For example, the mouth shape made to voice the vowel "a" was the one with the biggest, widest, and highest facial components compared to other mouth shapes, and people perceived words containing that vowel sound as bigger. The mouth shapes used to pronounce the vowel "i" were perceived as more likable than the other four vowels. These findings indicate that the mouth shapes producing vowels imply specific meanings. Our study provides clues about the meaning of verbal sounds and what the facial expressions in communication represent to the perceiver.

Neural dynamics during the vocalization of 'uh' or 'um'.

People occasionally use filler phrases or pauses, such as "uh", "um", or "y'know," that interrupt the flow of a sentence and fill silent moments between ordinary (non-filler) phrases. It remains unknown which brain networks are engaged during the utterance of fillers. We addressed this question by quantifying event-related cortical high gamma activity at 70-110 Hz. During extraoperative electrocorticography recordings performed as part of the presurgical evaluation, patients with drug-resistant focal epilepsy were instructed to overtly explain, in a sentence, 'what is in the image (subject)', 'doing what (verb)', 'where (location)', and 'when (time)'. Time-frequency analysis revealed that the utterance of fillers, compared to that of ordinary words, was associated with a greater magnitude of high gamma augmentation in association and visual cortex of either hemisphere. Our preliminary results raise the hypothesis that filler utterance would often occur when large-scale networks across the association and visual cortex are engaged in cognitive processing, including lexical retrieval as well as verbal working memory and visual scene scanning.

Spatiotemporal dynamics of auditory and picture naming-related high-gamma modulations: A study of Japanese-speaking patients.

OBJECTIVE: To characterize the spatiotemporal dynamics of auditory and picture naming-related cortical activation in Japanese-speaking patients. METHODS: Ten patients were assigned auditory naming and picture naming tasks during extraoperative intracranial EEG recording in a tertiary epilepsy center. Time-frequency analysis determined at what electrode sites and at what time windows during each task the amplitude of high-gamma activity (65-95 Hz) was modulated. RESULTS: The superior-temporal gyrus on each hemisphere showed high-gamma augmentation during sentence listening, whereas the left middle-temporal and inferior-frontal gyri showed high-gamma augmentation peaking around stimulus offset. Auditory naming-specific high-gamma augmentation was noted in the bilateral superior-temporal gyri as well as left frontal-parietal-temporal perisylvian network regions, whereas picture naming-specific augmentation was noted in the occipital-fusiform regions, bilaterally. The inferior pre- and postcentral gyri on each hemisphere showed modality-common high-gamma augmentation time-locked to overt responses. CONCLUSIONS: The spatiotemporal dynamics of auditory and picture naming-related high-gamma augmentation in Japanese-speaking patients were qualitatively similar to those previously reported in studies of English-speaking patients. SIGNIFICANCE: The cortical dynamics for auditory sentence recognition are at least partly shared by cohorts speaking two distinct languages. Multicenter studies regarding the clinical utility of high-gamma language mapping across Eastern and Western hemispheres may be feasible.

Neural dynamics of verbal working memory in auditory description naming.

Auditory naming is suggested to require verbal working memory (WM) operations in addition to speech sound perception during the sentence listening period and semantic/syntactic processing during the subsequent judgement period. We attempted to dissect cortical activations attributable to verbal WM from those otherwise involved in answering auditory sentence questions. We studied 19 patients who underwent electrocorticography recordings and measured high-gamma activity during auditory naming and WM tasks. In the auditory naming task, inferior-precentral high-gamma activity was augmented during sentence listening, and the magnitude of augmentation was independently correlated to that during the WM task maintenance period as well as patient age. High-gamma augmentation during the WM task scanning period accounted for high-gamma variance during the naming task judgement period in some of the left frontal association neocortex regions (most significantly in the middle-frontal, less in the inferior-frontal, and least in the orbitofrontal gyrus). Inferior-frontal high-gamma augmentation was left-hemispheric dominant during naming task judgement but rather symmetric during WM scanning. Left orbitofrontal high-gamma augmentation was evident only during the naming task judgement period but minimal during the WM task scanning period. The inferior-precentral regions may exert WM maintenance during sentence listening, and such maintenance function may be gradually strengthened as the brain matures. The left frontal association neocortex may have a dorsal-to-ventral gradient in functional roles during naming task judgement. Namely, left middle-frontal activation may be well-attributable to WM scanning function, whereas left orbitofrontal activation may be attributable less to WM scanning but more largely to syntactic/semantic processing.

Presurgical language mapping using event-related high-gamma activity: The Detroit procedure.

A number of investigators have reported that event-related augmentation of high-gamma activity at 70-110 Hz on electrocorticography (ECoG) can localize functionally-important brain regions in children and adults who undergo epilepsy surgery. The advantages of ECoG-based language mapping over the gold-standard stimulation include: (i) lack of stimulation-induced seizures, (ii) better sensitivity of localization of language areas in young children, and (iii) shorter patient participant time. Despite its potential utility, ECoG-based language mapping is far less commonly practiced than stimulation mapping. Here, we have provided video presentations to explain, point-by-point, our own hardware setting and time-frequency analysis procedures. We also have provided standardized auditory stimuli, in multiple languages, ready to be used for ECoG-based language mapping. Finally, we discussed the technical aspects of ECoG-based mapping, including its pitfalls, to facilitate appropriate interpretation of the data.

Spatio-temporal dynamics of working memory maintenance and scanning of verbal information.

OBJECTIVE: During verbal communication, humans briefly maintain mental representations of speech sounds conveying verbal information, and constantly scan these representations for comparison to incoming information. We determined the spatio-temporal dynamics of such short-term maintenance and subsequent scanning of verbal information, by intracranially measuring high-gamma activity at 70-110Hz during a working memory task. METHODS: Patients listened to a stimulus set of two or four spoken letters and were instructed to remember those letters over a two-second interval, following which they were asked to determine if a subsequent target letter had been presented earlier in that trial's stimulus set. RESULTS: Auditory presentation of letter stimuli sequentially elicited high-gamma augmentation bilaterally in the superior-temporal and pre-central gyri. During the two-second maintenance period, high-gamma activity was augmented in the left pre-central gyrus, and this effect was larger during the maintenance of stimulus sets consisting of four compared to two letters. During the scanning period following target presentation, high-gamma augmentation involved the left inferior-frontal and supra-marginal gyri. CONCLUSIONS: Short-term maintenance of verbal information is, at least in part, supported by the left pre-central gyrus, whereas scanning by the left inferior-frontal and supra-marginal gyri. SIGNIFICANCE: The cortical structures involved in short-term maintenance and scanning of speech stimuli were segregated with an excellent temporal resolution.

Neural mechanisms underlying the reward-related enhancement of motivation when remembering episodic memories with high difficulty.

The motivation to receive rewards enhances episodic memories, and the motivation is modulated by task difficulty. In episodic retrieval, however, functional neuroimaging evidence regarding the motivation that mediates interactions between reward and task difficulty is scarce. The present fMRI study investigated this issue. During encoding performed without fMRI, participants encoded Japanese words using either deep or shallow strategies, which led to variation in difficulty level during subsequent retrieval. During retrieval with fMRI, participants recognized the target words in either high or low monetary reward conditions. In the behavioral results, a reward-related enhancement of memory was found only when the memory retrieval was difficult, and the rewarding effect on subjective motivation was greater in the retrieval of memories with high difficulty than those with low difficulty. The fMRI data showed that reward-related increases in the activation of the substantia nigra/ventral tegmental area (SN/VTA), medial temporal lobe (MTL), dorsomedial prefrontal cortex (dmPFC), and dorsolateral prefrontal cortex (dlPFC) were greater during the retrieval of memories with high difficulty than those with low difficulty. Furthermore, reward-related enhancement of functional connectivity between the SN/VTA and MTL and between the SN/VTA and dmPFC during the retrieval of memories with high difficulty was significantly correlated with reward-related increases of retrieval accuracy and subjective motivation. The reward-related enhancement of episodic retrieval and retrieval-related motivation could be most effective when the level of retrieval difficulty is optimized. Such reward-related enhancement of memory and motivation could be modulated by a network including the reward-related SN/VTA, motivation-related dmPFC, and memory-related MTL. Hum Brain Mapp 38:3428-3443, 2017. © 2017 Wiley Periodicals, Inc.

Three- and four-dimensional mapping of speech and language in patients with epilepsy.

We have provided 3-D and 4D mapping of speech and language function based upon the results of direct cortical stimulation and event-related modulation of electrocorticography signals. Patients estimated to have right-hemispheric language dominance were excluded. Thus, 100 patients who underwent two-stage epilepsy surgery with chronic electrocorticography recording were studied. An older group consisted of 84 patients at least 10 years of age (7367 artefact-free non-epileptic electrodes), whereas a younger group included 16 children younger than age 10 (1438 electrodes). The probability of symptoms transiently induced by electrical stimulation was delineated on a 3D average surface image. The electrocorticography amplitude changes of high-gamma (70-110 Hz) and beta (15-30 Hz) activities during an auditory-naming task were animated on the average surface image in a 4D manner. Thereby, high-gamma augmentation and beta attenuation were treated as summary measures of cortical activation. Stimulation data indicated the causal relationship between (i) superior-temporal gyrus of either hemisphere and auditory hallucination; (ii) left superior-/middle-temporal gyri and receptive aphasia; (iii) widespread temporal/frontal lobe regions of the left hemisphere and expressive aphasia; and (iv) bilateral precentral/left posterior superior-frontal regions and speech arrest. On electrocorticography analysis, high-gamma augmentation involved the bilateral superior-temporal and precentral gyri immediately following question onset; at the same time, high-gamma activity was attenuated in the left orbitofrontal gyrus. High-gamma activity was augmented in the left temporal/frontal lobe regions, as well as left inferior-parietal and cingulate regions, maximally around question offset, with high-gamma augmentation in the left pars orbitalis inferior-frontal, middle-frontal, and inferior-parietal regions preceded by high-gamma attenuation in the contralateral homotopic regions. Immediately before verbal response, high-gamma augmentation involved the posterior superior-frontal and pre/postcentral regions, bilaterally. Beta-attenuation was spatially and temporally correlated with high-gamma augmentation in general but with exceptions. The younger and older groups shared similar spatial-temporal profiles of high-gamma and beta modulation; except, the younger group failed to show left-dominant activation in the rostral middle-frontal and pars orbitalis inferior-frontal regions around stimulus offset. The human brain may rapidly and alternately activate and deactivate cortical areas advantageous or obtrusive to function directed toward speech and language at a given moment. Increased left-dominant activation in the anterior frontal structures in the older age group may reflect developmental consolidation of the language system. The results of our functional mapping may be useful in predicting, across not only space but also time and patient age, sites specific to language function for presurgical evaluation of focal epilepsy.

High-gamma power changes after cognitive intervention: preliminary results from twenty-one senior adult subjects.

INTRODUCTION: Brain-imaging techniques have begun to be popular in evaluating the effectiveness of cognitive intervention training. Although gamma activities are rarely used as an index of training effects, they have several characteristics that suggest their potential suitability for this purpose. This pilot study examined whether cognitive training in elderly people affected the high-gamma activity associated with attentional processing and whether high-gamma power changes were related to changes in behavioral performance. METHODS: We analyzed (MEG) magnetoencephalography data obtained from 35 healthy elderly subjects (60-75 years old) who had participated in our previous intervention study in which the subjects were randomly assigned to one of the three types of intervention groups: Group V trained in a vehicle with a newly developed onboard cognitive training program, Group P trained with a similar program but on a personal computer, and Group C was trained to solve a crossword puzzle as an active control group. High-gamma (52-100 Hz) activity during a three-stimulus visual oddball task was measured before and after training. As a result of exclusion in the MEG data analysis stage, the final sample consisted of five subjects in Group V, nine subjects in Group P, and seven subjects in Group C. RESULTS: Results showed that high-gamma activities were differently altered between groups after cognitive intervention. In particular, members of Group V, who showed significant improvements in cognitive function after training, exhibited increased high-gamma power in the left middle frontal gyrus during top-down anticipatory target processing. High-gamma power changes in this region were also associated with changes in behavioral performance. CONCLUSIONS: Our preliminary results suggest the usefulness of high-gamma activities as an index of the effectiveness of cognitive training in elderly subjects.

High-gamma activity in an attention network predicts individual differences in elderly adults' behavioral performance.

The current study used a magnetoencephalogram to investigate the relationship between high-gamma (52-100 Hz) activity within an attention network and individual differences in behavioral performance among healthy elderly adults. We analyzed brain activity in 41 elderly subjects performing a 3-stimulus visual oddball task. In addition to the average amplitude of event-related fields in the left intraparietal sulcus (IPS), high-gamma power in the left middle frontal gyrus (MFG), the strength of high-gamma imaginary coherence between the right MFG and the left MFG, and those between the right MFG and the left thalamus predicted individual differences in reaction time. In addition, high-gamma power in the left MFG was correlated with task accuracy, whereas high-gamma power in the left thalamus and left IPS was correlated with individual processing speed. The direction of correlations indicated that higher high-gamma power or coherence in an attention network was associated with better task performance and, presumably, higher cognitive function. Thus, high-gamma activity in different regions of this attention network differentially contributed to attentional processing, and such activity could be a fundamental process associated with individual differences in cognitive aging.

Remembering with gains and losses: effects of monetary reward and punishment on successful encoding activation of source memories.

The motivation of getting rewards or avoiding punishments reinforces learning behaviors. Although the neural mechanisms underlying the effect of rewards on episodic memory have been demonstrated, there is little evidence of the effect of punishments on this memory. Our functional magnetic resonance imaging (fMRI) study investigated the effects of monetary rewards and punishments on activation during the encoding of source memories. During encoding, participants memorized words (item) and locations of presented words (source) under 3 conditions (Reward, Punishment, and Control). During retrieval, participants retrieved item and source memories of the words and were rewarded or penalized according to their performance. Source memories encoded with rewards or punishments were remembered better than those without such encoding. fMRI data demonstrated that the ventral tegmental area and substantia nigra and nucleus accumbens activations reflected both the processes of reward and punishment, whereas insular activation increased as a linear function of punishment. Activation in the hippocampus and parahippocampal cortex predicted subsequent retrieval success of source memories. Additionally, correlations between these reward/punishment-related regions and the hippocampus were significant. The successful encoding of source memories could be enhanced by punishments and rewards, and interactions between reward/punishment-related regions and memory-related regions could contribute to memory enhancement by reward and/or punishment.