The Effects of Frequency, Variability, and Co-occurrence on Category Formation in Neural Systems.

Objects are grouped into categories through a complex combination of statistical and structural regularities. We sought to better understand the neural responses to the structural features of object categories that result from implicit learning. Adult participants were exposed to 32 object categories that contained three structural properties: frequency, variability, and co-occurrences, during an implicit learning task. After this exposure, participants completed a recognition task and were then presented with blocks of learned object categories during fMRI sessions. Analyses were performed by extracting data from ROIs placed throughout the fusiform gyri and lateral occipital cortex and comparing the effects of the different structural properties throughout the ROIs. Behaviorally, we found that symbol category recognition was supported by frequency, but not variability. Neurally, we found that sensitivity to object categories was greater in the right hemisphere and increased as ROIs were moved posteriorly. Frequency and variability altered the brain activation while processing object categories, although the presence of learned co-occurrences did not. Moreover, variability and co-occurrence interacted as a function of ROI, with the posterior fusiform gyrus being most sensitive to this relationship. This result suggests that variability may guide the learner to relevant co-occurrences and this is supported by the posterior ventral temporal cortex. Broadly, our results suggest that the internal features of the categories themselves are key factors in the category learning system.

Category structure guides the formation of neural representations.

Perceptual variability is often viewed as having multiple benefits in object learning and categorization. Despite the abundant results demonstrating benefits such as increased transfer of knowledge, the neural mechanisms underlying variability as well as the developmental trajectories of how variability precipitates changes to category boundaries are unknown. By manipulating an individual's exposure to variability of novel, metrically organized categories during an fMRI-adaptation paradigm, we were able to assess the functional differences between similarity and variability in category learning and generalization across two time-points in development: adulthood (n = 14) and late childhood (n = 13). During this study, participants were repeatedly exposed to category members from different distributions. After a period of adaptation, a deviant stimulus that differed from the expected distribution was then presented. This deviant differed in either an invariant dimension (a feature that remained consistent throughout presentation was altered) or a similarity dimension (a feature that changed throughout exposure was changed in a new dimension). Our results can be summarized in three main findings: (1) Variability during exposure recruited the right fusiform gyrus to a greater extent than tight exposure. (2) Deviant items were generalized based on the exemplar distributions during exposure, although children only generalized items if provided variable exposure. (3) Variability influenced release to a greater extent in children than adults. These results are discussed in relation to the variability and category learning literature more broadly.

Visual experiences during letter production contribute to the development of the neural systems supporting letter perception.

Letter production through handwriting creates visual experiences that may be important for the development of visual letter perception. We sought to better understand the neural responses to different visual percepts created during handwriting at different levels of experience. Three groups of participants, younger children, older children, and adults, ranging in age from 4.5 to 22 years old, were presented with dynamic and static presentations of their own handwritten letters, static presentations of an age-matched control's handwritten letters, and typeface letters during fMRI. First, data from each group were analyzed through a series of contrasts designed to highlight neural systems that were most sensitive to each visual experience in each age group. We found that younger children recruited ventral-temporal cortex during perception and this response was associated with the variability present in handwritten forms. Older children and adults also recruited ventral-temporal cortex; this response, however, was significant for typed letter forms but not variability. The adult response to typed letters was more distributed than in the children, including ventral-temporal, parietal, and frontal motor cortices. The adult response was also significant for one's own handwritten letters in left parietal cortex. Second, we compared responses among age groups. Compared to older children, younger children demonstrated a greater fusiform response associated with handwritten form variability. When compared to adults, younger children demonstrated a greater response to this variability in left parietal cortex. Our results suggest that the visual perception of the variability present in handwritten forms that occurs during handwriting may contribute to developmental changes in the neural systems that support letter perception.

An Analysis of the Brain Systems Involved with Producing Letters by Hand.

Complex visual-motor behaviors dominate human-environment interactions. Letter production, writing individual letters by hand, is an example of a complex visual-motor behavior composed of numerous behavioral components, including the required motor movements and the percepts that those motor movements create. By manipulating and isolating components of letter production, we provide experimental evidence that this complex visual-motor behavior is supported by a widespread neural system that is composed of smaller subsystems related to different sensorimotor components. Adult participants hand-printed letters with and without "ink" on an MR-safe digital writing tablet, perceived static and dynamic representations of their own handwritten letters, and perceived typeface letters during fMRI scanning. Our results can be summarized by three main findings: (1) Frontoparietal systems were associated with the motor component of letter production, whereas temporo-parietal systems were more associated with the visual component. (2) The more anterior regions of the left intraparietal sulcus were more associated with the motor component, whereas the more posterior regions were more associated with the visual component, with an area of visual-motor overlap in the posterior intraparietal sulcus. (3) The left posterior intraparietal sulcus and right fusiform gyrus responded similarly to both visual and motor components, and both regions also responded more during the perception of one's own handwritten letters compared with perceiving typed letters. These findings suggest that the neural systems recruited during complex visual-motor behaviors are composed of a set of interrelated sensorimotor subsystems that support the full behavior in different ways and, furthermore, that some of these subsystems can be rerecruited during passive perception in the absence of the full visual-motor behavior.

Dorsal stream function in the young child: an fMRI investigation of visually guided action.

Visually guided action is a ubiquitous component of human behavior, but the neural substrates that support the development of this behavior are unknown. Here we take an initial step in documenting visual-motor system development in the young (4- to 7-year-old) child. Through functional MRI and by using a new technique to measure the mechanisms underlying real-time visually guided action in the MRI environment, we demonstrate that children rely primarily on the IPS and cerebellum for this complex behavior. This pattern is consistent across three different visually guided actions, suggesting generalizability of these neural substrates across such tasks. However, minor differences in neural processing across tasks were also demonstrated. Overall, results are interpreted as demonstrating that the functions of the dorsal stream can be viewed as fairly mature in the young child. These results provide a benchmark for future studies that aim to understand the development of the neural circuitry for visually guided action.

Gesture for generalization: gesture facilitates flexible learning of words for actions on objects.

Verb learning is difficult for children (Gentner, ), partially because children have a bias to associate a novel verb not only with the action it represents, but also with the object on which it is learned (Kersten & Smith, ). Here we investigate how well 4- and 5-year-old children (N = 48) generalize novel verbs for actions on objects after doing or seeing the action (e.g., twisting a knob on an object) or after doing or seeing a gesture for the action (e.g., twisting in the air near an object). We find not only that children generalize more effectively through gesture experience, but also that this ability to generalize persists after a 24-hour delay.

A new statistical model for analyzing rating scale data pertaining to word meaning.

The concrete-abstract categorization scheme has guided several research programs. A popular way to classify words into one of these categories is to calculate a word's mean value in a Concreteness or Imageability rating scale. However, this procedure has several limitations. For instance, results can be highly distorted by outliers, ascribe differences among words when none may exist, and neglect rating trends in participants. We suggest using an alternative procedure to analyze rating scale data called median polish analysis (MPA). MPA is tolerant to outliers and accounts for information in multiple dimensions, including trends among participants. MPA performance can be readily evaluated using an effect size measure called analog R 2 and be integrated with bootstrap 95% confidence intervals, which can prevent assigning inexistent differences among words. To compare these analysis procedures, we asked 80 participants to rate a set of nouns and verbs using four different rating scales: Action, Concreteness, Imageability, and Multisensory. We analyzed the data using both two-way and three-way MPA models. We also calculated 95% CIs for the two-way models. Categorizing words with the Action scale revealed a continuum of word meaning for both nouns and verbs. The remaining scales produced dichotomous or stratified results for nouns, and continuous results for verbs. While the sample mean analysis generated continua irrespective of the rating scale, MPA differentiated among dichotomies and continua. We conclude that MPA allowed us to better classify words by discarding outliers, focusing on main trends, and considering the differences in rating criteria among participants.

Neural substrates of sensorimotor processes: letter writing and letter perception.

Writing and perceiving letters are thought to share similar neural substrates; however, what constitutes a neural representation for letters is currently debated. One hypothesis is that letter representation develops from sensorimotor experience resulting in an integrated set of modality-specific regions, whereas an alternative account suggests that letter representations may be abstract, independent of modality. Studies reviewed suggest that letter representation consists of a network of modality-responsive brain regions that may include an abstract component.

Some views are better than others: evidence for a visual bias in object views self-generated by toddlers.

How objects are held determines how they are seen, and may thereby play an important developmental role in building visual object representations. Previous research suggests that toddlers, like adults, show themselves a disproportionate number of planar object views - that is, views in which the objects' axes of elongation are perpendicular or parallel to the line of sight. Here, three experiments address three explanations of this bias: (1) that the locations of interesting features of objects determine how they are held and thus how they are viewed; (2) that ease of holding determines object views; and (3) that there is a visual bias for planar views that exists independently of holding and of interesting surface properties. Children 18 to 24 months of age manually and visually explored novel objects (1) with interesting features centered in planar or ¾ views; (2) positioned inside Plexiglas boxes so that holding biased either planar or non-planar views; and (3) positioned inside Plexiglas spheres, so that no object properties directly influenced holding. Results indicate a visual bias for planar views that is influenced by interesting surface properties and ease of holding, but that continues to exist even when these factors push for alternative views.

Using the axis of elongation to align shapes: developmental changes between 18 and 24 months of age.

An object's axis of elongation serves as an important frame of reference for forming three-dimensional representations of object shape. By several recent accounts, the formation of these representations is also related to experiences of acting on objects. Four experiments examined 18- to 24-month-olds' (N=103) sensitivity to the elongated axis in action tasks that required extracting, comparing, and physically rotating an object so that its major axis was aligned with that of a visual standard. In Experiments 1 and 2, the older toddlers precisely rotated both simple and complexly shaped three-dimensional objects in insertion tasks where the visual standard was the rectangular contour defining the opening in a box. The younger toddlers performed poorly. Experiments 3 and 4 provide evidence on emerging abilities in extracting and using the most extended axis as a frame of reference for shape comparison. Experiment 3 showed that 18-month-olds could rotate an object to align its major axis with the direction of their own hand motion, and Experiment 4 showed that they could align the major axis of one object with that of another object of the exact same three-dimensional shape. The results are discussed in terms of theories of the development of three-dimensional shape representations, visual object recognition, and the role of action in these developments.

Neural correlates of gesture processing across human development.

Co-speech gesture facilitates learning to a greater degree in children than in adults, suggesting that the mechanisms underlying the processing of co-speech gesture differ as a function of development. We suggest that this may be partially due to children's lack of experience producing gesture, leading to differences in the recruitment of sensorimotor networks when comparing adults to children. Here, we investigated the neural substrates of gesture processing in a cross-sectional sample of 5-, 7.5-, and 10-year-old children and adults and focused on relative recruitment of a sensorimotor system that included the precentral gyrus (PCG) and the posterior middle temporal gyrus (pMTG). Children and adults were presented with videos in which communication occurred through different combinations of speech and gesture during a functional magnetic resonance imaging (fMRI) session. Results demonstrated that the PCG and pMTG were recruited to different extents in the two populations. We interpret these novel findings as supporting the idea that gesture perception (pMTG) is affected by a history of gesture production (PCG), revealing the importance of considering gesture processing as a sensorimotor process.

The Lateralizer: a tool for students to explore the divided brain.

Despite a profusion of popular misinformation about the left brain and right brain, there are functional differences between the left and right cerebral hemispheres in humans. Evidence from split-brain patients, individuals with unilateral brain damage, and neuroimaging studies suggest that each hemisphere may be specialized for certain cognitive processes. One way to easily explore these hemispheric asymmetries is with the divided visual field technique, where visual stimuli are presented on either the left or right side of the visual field and task performance is compared between these two conditions; any behavioral differences between the left and right visual fields may be interpreted as evidence for functional asymmetries between the left and right cerebral hemispheres. We developed a simple software package that implements the divided visual field technique, called the Lateralizer, and introduced this experimental approach as a problem-based learning module in a lower-division research methods course. Second-year undergraduate students used the Lateralizer to experimentally challenge and explore theories of the differences between the left and right cerebral hemispheres. Measured learning outcomes after active exploration with the Lateralizer, including new knowledge of brain anatomy and connectivity, were on par with those observed in an upper-division lecture course. Moreover, the project added to the students' research skill sets and seemed to foster an appreciation of the link between brain anatomy and function.

Vision for action in toddlers: the posting task.

Three experiments examine 18- to 24-month-old (N = 78) toddlers' ability to spatially orient objects by their major axes for insertion into a slot. This is a simplified version of the posting task that is commonly used to measure dorsal stream functioning. The experiments identify marked developmental changes in children's ability to preorient objects for insertion, with 18-month-olds failing completely and 24-month-olds succeeding easily. In marked contrast, 18-month-olds preorient their empty hands for insertion into the same slots. This developmental dissociation between aligning hands and aligning objects to slots suggests that the key developmental change is in action with the goal of object-to-object alignment versus action on an object.

Protracted Neural Development of Dorsal Motor Systems During Handwriting and the Relation to Early Literacy Skills.

Handwriting is a complex visual-motor skill that affects early reading development. A large body of work has demonstrated that handwriting is supported by a widespread neural system comprising ventral-temporal, parietal, and frontal motor regions in adults. Recent work has demonstrated that this neural system is largely established by 8 years of age, suggesting that the development of this system occurs in young children who are still learning to read and write. We made use of a novel MRI-compatible writing tablet that allowed us to measure brain activation in 5-8-year-old children during handwriting. We compared activation during handwriting in children and adults to provide information concerning the developmental trajectory of the neural system that supports handwriting. We found that parietal and frontal motor involvement during handwriting in children is different from adults, suggesting that the neural system that supports handwriting changes over the course of development. Furthermore, we found that parietal and frontal motor activation correlated with a literacy composite score in our child sample, suggesting that the individual differences in the dorsal response during handwriting are related to individual differences in emerging literacy skills. Our results suggest that components of the widespread neural system supporting handwriting develop at different rates and provide insight into the mechanisms underlying the contributions of handwriting to early literacy development.

"Neural differences in expert guitarists during over-learned non-standard visuomotor mapping of abstract versus concrete information".

Using visual information to perform actions is a fundamental aspect of human behavior. Musicians commonly translate visual information into action using both concrete and abstract visual information. We exposed expert guitarists to four types of familiar visual depictions of action instruction including musical notation (very abstract), tablature (abstract), chord diagrams (more concrete), and actual pictures of guitars chords being formed (very concrete). These were shown during fMRI scanning as the guitarists formed the appropriate chords (as visually depicted) on a magnet safe guitar fret board with strings, or where they simply viewed the visual stimuli without an action. Whole brain contrasts revealed that the right precuneus was more active for abstract instruction while an occipito-insular circuit was more active for concrete instruction. The current findings highlight that the degree of over-learned visual abstraction is an important factor modulating visual-motor processing.

The neural correlates of attempting to suppress negative versus neutral memories.

We performed an event-related fMRI study comparing attempts at suppressing recall of negative versus neutral memories. The hippocampus is crucial for successful explicit recall. Hippocampal activation has been shown to decrease during the suppression of previously learned neutral words. However, different effects may occur in the case of emotional memories. Participants first learned 40 word pairs consisting of a cue and either a neutral or a negative target. During fMRI scanning, the participants were shown the cues and were instructed to recall the targets or to suppress the targets, using attentional distraction. Similar right-lateralized frontoparietal regions were activated more during suppression than during recall, regardless of emotion. However, we show for the first time that lowered hippocampal activation occurs during the suppression of neutral, but not negative, words. Coinciding with this sustained hippocampal activation, the amygdala, insula, anterior cingulate, and fusiform gyrus showed greater activation during the suppression of negative memories than during suppression of neutral memories. Thus, during attempts to suppress negative memories, regions involved in the emotional and sensory aspects of memory reactivate, along with regions indexing conscious recall. Revealing the neural correlates and mechanisms of the suppression of negative memories has relevance for disorders such as posttraumatic stress disorder, in which traumatic memories often intrude and are associated with avoidance. Supplemental materials for this article may be downloaded from http://cabn.psychonomic-journals.org/content/supplemental.

Early biases and developmental changes in self-generated object views.

Object recognition depends on the seen views of objects. These views depend in part on the perceivers' own actions as they select and show object views to themselves. The self-selection of object views from manual exploration of objects during infancy and childhood may be particularly informative about the human object recognition system and its development. Here, we report for the first time on the structure of object views generated by 12 to 36 month old children (N = 54) and for comparison adults (N = 17) during manual and visual exploration of objects. Object views were recorded via a tiny video camera placed low on the participant's forehead. The findings indicate two viewing biases that grow rapidly in the first three years: a bias for planar views and for views of objects in an upright position. These biases also strongly characterize adult viewing. We discuss the implications of these findings for a developmentally complete theory of object recognition.

Ecological validity of experimental set-up affects parietal involvement during letter production.

Studies of symbol production using fMRI often use techniques that introduce an artificial pairing between motor production and visual perception. These techniques allow participants to see their own output by recording their pen trajectories using a touchscreen-only tablet and displaying these productions on a mirror placed above their head. We recently developed an MR-safe writing tablet with video display that allows participants to see their own hand and their own productions while producing symbols in real time on the surface where they are producing them-allowing for more ecologically valid fMRI studies of production. We conducted a study to determine whether the participation of posterior parietal cortex during symbol production was affected by the pairing of motor production and visual feedback associated with the two types of tablets. We performed ROI analyses in intraparietal sulcus while adult participants produced letters to dictation using either a touchscreen-only tablet (no visual guidance of the hand) (n = 14) or using a touchscreen-and-video-display tablet (visual guidance of the hand) (n = 14). We found that left posterior intraparietal sulcus was more active during production with the touchscreen-only tablet than during production with the touchscreen-and-video-display tablet. These results suggest that posterior parietal involvement during production tasks is associated with the somewhat artificial visual-motor pairing that is introduced by the techniques used in some studies of symbol production.

Constraining Stroke Order During Manual Symbol Learning Hinders Subsequent Recognition in Children Under 4 1/2 Years.

In the age of technology, writing by hand has become less common than texting and keyboarding. Learning letters by hand, however, has been shown to have profound developmental importance. One aspect of writing by hand that has been understudied is the effect of learning symbols stroke-by-stroke, a dynamic action that does not occur with keyboarding. We trained children to draw novel symbols in either an instructed stroke order or in a self-directed stroke order and tested: (1) whether learning novel symbols in a self-directed stroke order benefits subsequent recognition more than learning in a specified stroke order, (2) whether seeing novel symbols unfold in the stroke order that was taught would aid in recognition, and (3) whether any effects are age-dependent. Our results demonstrate that producing a symbol with a self-directed stroke order provides more benefit to symbol recognition than instructed stroke orders in 4.0-4.5-year-old children but not in 4.5-5.0-year-old children. We found, further, that the observed recognition benefits were not affected by seeing the symbol unfold in the same stroke order it was learned during testing, suggesting that the learning was not reliant upon the exact stroke order experienced during learning. These results stress the importance of allowing children to produce symbols in a self-directed manner and, by extension, that constraining how a child learns to write can adversely affect subsequent recognition.

When writing impairs reading: letter perception's susceptibility to motor interference.

The effect of writing on the concurrent visual perception of letters was investigated in a series of studies using an interference paradigm. Participants drew shapes and letters while simultaneously visually identifying letters and shapes embedded in noise. Experiments 1-3 demonstrated that letter perception, but not the perception of shapes, was affected by motor interference. This suggests a strong link between the perception of letters and the neural substrates engaged during writing. The overlap both in category (letter vs. shape) and in the perceptual similarity of the features (straight vs. curvy) of the seen and drawn items determined the amount of interference. Experiment 4 demonstrated that intentional production of letters is not necessary for the interference to occur, because passive movement of the hand in the shape of letters also interfered with letter perception. When passive movements were used, however, only the category of the drawn items (letters vs. shapes), but not the perceptual similarity, had an influence, suggesting that motor representations for letters may selectively influence visual perception of letters through proprioceptive feedback, with an additional influence of perceptual similarity that depends on motor programs.