Setting boundaries: Development of neural and behavioral event cognition in early childhood.

The ongoing stream of sensory experience is so complex and ever-changing that we tend to parse this experience at "event boundaries," which structures and strengthens memory. Memory processes undergo profound change across early childhood. Whether young children also divide their ongoing processing along event boundaries, and if those boundaries relate to memory, could provide important insight into the development of memory systems. In Study 1, 4-7-year-old children and adults segmented a cartoon, and we tested their memory. Children's event boundaries were more variable than adults' and differed in location and consistency of agreement. Older children's event segmentation was more adult-like than younger children's, and children who segmented events more like adults had better memory for those events. In Study 2, we asked whether these developmental differences in event segmentation had their roots in distinct neural representations. A separate group of 4-8-year-old children watched the same cartoon while undergoing an fMRI scan. In the right hippocampus, greater pattern dissimilarity across event boundaries compared to within events was evident for both child and adult behavioral boundaries, suggesting children and adults share similar event cognition. However, the boundaries identified by a data-driven Hidden Markov Model found that a different brain region-the left and right angular gyrus-aligned only with event boundaries defined by children. Overall, these data suggest that children's event cognition is reasonably well-developed by age 4 but continues to become more adult-like across early childhood. RESEARCH HIGHLIGHTS: Adults naturally break their experience into events, which structures and strengthens memory, but less is known about children's event perception and memory. Study 1 had adults and children segment and remember events from an animated show, and Study 2 compared those segmentations to other children's fMRI data. Children show better recognition and temporal order memory and more adult-like event segmentation with age, and children who segment more like adults have better memory. Children's and adults' behavioral boundaries mapped onto pattern similarity differences in hippocampus, and children's behavioral boundaries matched a data-driven model's boundaries in angular gyrus.

Understanding relational binding in early childhood: Interacting effects of overlap and delay.

Episodic memories typically share overlapping elements in distinctive combinations, and to be valuable for future behavior they need to withstand delays. There is relatively little work on whether children have special difficulty with overlap or withstanding delay. However, Yim, Dennis, and Sloutsky (Psychological Science, 2013, Vol. 24, pp. 2163-2172) suggested that extensive overlap is more problematic for younger children, and Darby and Sloutsky (Psychological Science, 2015, Vol. 26, pp. 1937-1946) reported that a 48-h delay period actually improves children's memory for overlapping pairs of items. In the current study, we asked how children's episodic memory is affected by stimulus overlap, delay, and age using visual stimuli containing either overlapping or unique item pairs. Children aged 4 and 6 years were tested both immediately and after a 24-h delay. As expected, older children performed better than younger children, and both age groups performed worse on overlapping pairs. Surprisingly, the 24-h delay had only a marginal effect on overall accuracy. Although there were no interactions, when errors were examined, there was evidence that delay buffered memory for overlapping pairs against cross-contextual confusion for younger children.

Children show adult-like hippocampal pattern similarity for familiar but not novel events.

The ability to detect differences among similar events in our lives is a crucial aspect of successful episodic memory performance, which develops across early childhood. The neural substrate of this ability is supported by operations in the medial temporal lobe (MTL). Here, we used representational similarity analysis (RSA) to measure neural pattern similarity in hippocampus, perirhinal cortex, and parahippocampal cortex for 4- to 10-year-old children and adults during naturalistic viewing of clips from the same compared to different movies. Further, we assessed the role of prior exposure to individual movie clips on pattern similarity in the MTL. In both age groups, neural pattern similarity in hippocampus was lower for clips drawn from the same movies compared to those drawn from different movies, suggesting that related content activates processes focused on keeping representations with shared content distinct. However, children showed this only for movies with which they had prior exposures, whereas adults showed the effect regardless of any prior exposures to the movies. These findings suggest that children require repeated exposure to stimuli to show adult-like MTL functioning in distinguishing among similar events.

Contingency of semantic generalization on episodic specificity varies across development.

Semantic memory-general knowledge of ideas and concepts-includes generalization processes that support inference. Episodic memory, on the other hand, preserves the specificity of individual events by binding together unique combinations of elements from an episode and relies on pattern separation to distinguish similar experiences. These two memory systems play complementary roles, supporting different mnemonic goals, but the nature and extent of their interdependence is unclear.1,2 Some models suggest that new information is encoded initially as hippocampus-dependent episodic memory and then, either through repetition or gist extraction, becomes semantic over time.3,4 These models also posit a neocortical route to semantic memory acquisition exists that can bypass the hippocampus.3 Both proposed routes are slow learning mechanisms, yet generalization can occur rapidly. Recent models suggest that fast generalization relies, in part, on the retrieval of individual but related episodes.5,6 Such episodic memory gating mechanisms render fast generalization contingent on the memory specificity of instances, a pattern that has been observed in adults.7,8 None of these models take into account the observation that generalization and episodic specificity have asynchronous developmental profiles, with generalization emerging years before episodic memory.9,10 We ask two questions about generalized and specific memory during early childhood: first, is rapid generalization contingent on remembering specific past memories? And second, does the strength or nature of this contingency differ across development? We found that the interdependence of generalization and episodic memory varies across development: generalization success in adults, but not in children, was contingent on context binding.

Dissecting the Fornix in Basic Memory Processes and Neuropsychiatric Disease: A Review.

Background: The fornix is the primary axonal tract of the hippocampus, connecting it to modulatory subcortical structures. This review reveals that fornix damage causes cognitive deficits that closely mirror those resulting from hippocampal lesions. Methods: We reviewed the literature on the fornix, spanning non-human animal lesion research, clinical case studies of human patients with fornix damage, as well as diffusion-weighted imaging (DWI) work that evaluates fornix microstructure in vivo. Results: The fornix is essential for memory formation because it serves as the conduit for theta rhythms and acetylcholine, as well as providing mnemonic representations to deep brain structures that guide motivated behavior, such as when and where to eat. In rodents and non-human primates, fornix lesions lead to deficits in conditioning, reversal learning, and navigation. In humans, damage to the fornix manifests as anterograde amnesia. DWI research reveals that the fornix plays a key role in mild cognitive impairment and Alzheimer's Disease, and can potentially predict conversion from the former to the latter. Emerging DWI findings link perturbations in this structure to schizophrenia, mood disorders, and eating disorders. Cutting-edge research has investigated how deep brain stimulation of the fornix can potentially attenuate memory loss, control epileptic seizures, and even improve mood. Conclusions: The fornix is essential to a fully functioning memory system and is implicated in nearly all neurological functions that rely on the hippocampus. Future research needs to use optimized DWI methods to study the fornix in vivo, which we discuss, given the difficult nature of fornix reconstruction. Impact Statement The fornix is a white matter tract that connects the hippocampus to several subcortical brain regions and is pivotal for episodic memory functioning. Functionally, the fornix transmits essential neurotransmitters, as well as theta rhythms, to the hippocampus. In addition, it is the conduit by which memories guide decisions. The fornix is biomedically important because lesions to this tract result in irreversible anterograde amnesia. Research using in vivo imaging methods has linked fornix pathology to cognitive aging, mild cognitive impairment, psychosis, epilepsy, and, importantly, Alzheimer's Disease.