BackgroundTool use is defined as the manipulation of an inanimate object to change the position or form of a separate object. The expansion of cognitive niches and tool-use capabilities probably stimulated each other in hominid evolution. To understand the causes of cognitive expansion in humans, we need to know the behavioral and neural basis of tool use. Although a wide range of animals exhibit tool use in nature, most studies have focused on primates and birds on behavioral or psychological levels and did not directly address questions of which neural modifications contributed to the emergence of tool use. To investigate such questions, an animal model suitable for cellular and molecular manipulations is needed.Methodology/Principal FindingsWe demonstrated for the first time that rodents can be trained to use tools. Through a step-by-step training procedure, we trained degus (Octodon degus) to use a rake-like tool with their forelimbs to retrieve otherwise out-of-reach rewards. Eventually, they mastered effective use of the tool, moving it in an elegant trajectory. After the degus were well trained, probe tests that examined whether they showed functional understanding of the tool were performed. Degus did not hesitate to use tools of different size, colors, and shapes, but were reluctant to use the tool with a raised nonfunctional blade. Thus, degus understood the functional and physical properties of the tool after extensive training.Conclusions/SignificanceOur findings suggest that tool use is not a specific faculty resulting from higher intelligence, but is a specific combination of more general cognitive faculties. Studying the brains and behaviors of trained rodents can provide insights into how higher cognitive functions might be broken down into more general faculties, and also what cellular and molecular mechanisms are involved in the emergence of such cognitive functions.

The ability to hierarchically combine multiple objects to construct a self-embedded structure is assumed to be unique to primates and parrots. However, we observed that Degus, a species of rodents, spontaneously constructed this structure while they were being trained for an unrelated vocal operant task. With a large dust-bath bowl, a medium sized food cup, and a small toy ball, they built a triplet "Chinese boxes." The strategy by Degus in constructing this structure was similar with that by primates: first they mainly constructed pairs of the objects and then proceeded to triplets. The relation between the evolutionary emergence of hierarchical construction and language acquisition is discussed.

This study examines the neural substrate of perspective-taking by analyzing the electroencephalographic (EEG) activity elicited by the auditory comprehension of sentences for which the comprehender had to adopt the perspective of the person described in them. Recent studies suggest that the ability of perspective-taking can be an integrative function of temporal and spatial information processing. We thus examined the independence and possible interaction of human perspective shifts and temporal perspective-taking by utilizing Japanese subsidiary verbs for giving, namely -ageru and -kureru. We manipulated human perspective shifts and temporal perspective-taking independently in experimental sentences by syntactically changing the subject and the object between the speaker and a third person, while we manipulated the tense to be past or non-past tense via sentence-final particles ru/ta (non-past/past). The EEG analyses via electrodes indicated the suppression of the β band for human perspective shifts in sentences in non-past tense and the absence of such suppression in sentences in past tense. The analyses for the clusters of independent components indicated β suppression for past tense against non-past tense in sentences without a human perspective shift. This response pattern suggests a close relationship between human perspective shifting and temporal perspective-taking. The β suppression for the human perspective shift in our experiment can be understood as a replication of the previous EEG findings observed for perspective-taking in the presentation of visual images. The preceding findings and our result suggest that the ability or the function of perspective-taking is not specific to the modality. Furthermore, the generator of the β suppression for past tense against non-past tense without human perspective shifting was localized in the precuneus, which is consistent with recent findings indicating that the precuneus is deeply involved in time perception.

This study attempts to detect the differences in event-related potentials (ERPs) associated with two syntactic processes: the syntactic integration of discontinuous dependency and the detection of a violation of the syntactic island constraint. We recorded the electroencephalogram elicited by complex sentences in Japanese that included a dependency between a quantifier and its head noun, in which we changed the word order of the two words to manipulate the presence and absence of a syntactic integration and a syntactic island violation while keeping the lexical items and construction unchanged. We found significant negative and positive deflections for the syntactic integration only when a quantifier preceded its head noun. We also observed significant negative and positive deflections for the syntactic island violation, for which the negativity was more salient when a quantifier preceded its head noun. This study is the first to report a late positive ERP for a violation of the syntactic island constraints in Japanese, and the results showed that the ERP elicited by syntactic integration and that by syntactic island violation were different in terms of their latency, topography, and duration. More importantly, the ERPs elicited by the two syntactic processes were biphasic, and the amplitudes of the negative ERP and of the positive ERP were positively correlated. This positive correlation could be a characteristic of syntactic processing because it contrasted with the negative correlation reported for the ERP elicited by semantic anomalies in English. Furthermore, the amplitude of the ERP for syntactic integration was negatively correlated with the individual capacity of working memory (WM). That is, a reader with greater WM capacity showed smaller negativity and positivity for the syntactic integration, whereas the amplitude for the syntactic island violation showed no significant correlation with the individual capacity of WM. Our results suggested that linguistic ERPs functionally interacted with each other and that the ERP involving the retention and the retrieval of a distant word could be constrained by the individual differences in WM capacity. We discuss the possible reasons for the contrast between English and Japanese on the basis of the cross-linguistic differences in the two languages.

This study examines the neural substrate of the understanding of human relationships in verbal communication with Japanese honorific sentences as experimental materials. We manipulated two types of Japanese verbs specifically used to represent respect for others, i.e., exalted and humble verbs, which represent respect for the person in the subject and the person in the object, respectively. We visually presented appropriate and anomalous sentences containing the two types of verbs and analyzed the electroencephalogram elicited by the verbs. We observed significant parietal negativity at a latency of approximately 400 ms for anomalous verbs compared with appropriate verbs. This parietal negativity could be a manifestation of the pragmatic process used to integrate the linguistic forms with the human relationships represented in the sentences. The topographies of these event-related potentials (ERPs) corresponded well with those of ERPs for two second-person pronouns in Chinese (plain ni and respectful nin). This correspondence suggests that the pragmatic integration process in honorific expressions is cross-linguistically common in part. Furthermore, we assessed the source localization by means of independent component (IC) analysis and dipole fitting and observed a significant difference in ERP between the honorific and control sentences in the IC cluster centered in the precentral gyrus and in the cluster centered in the medial part of the occipital lobe, which corresponded well with the functional magnetic resonance imaging findings for Japanese honorification. We also found several significant differences in the time-frequency analyses for the medial occipital cluster. These significant differences in the medial occipital cluster suggested that the circuit of the theory of mind was involved in the processing of Japanese honorification. Our results suggest that pragmatic and syntactic processing are performed in parallel because the person to be respected must fulfill the grammatical function appropriate for the honorific verb.