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Open Access Publications from the University of California

Department of Psychology

UC San Diego

Recent Work

Cover page of Prior episodic learning and the efficacy of retrieval practice.

Prior episodic learning and the efficacy of retrieval practice.


In three experiments we investigated how the level of study-based, episodic knowledge influences the efficacy of subsequent retrieval practice (testing) as a learning event. Possibilities are that the efficacy of a test, relative to a restudy control, decreases, increases, or is independent of the degree of prior study-based learning. The degree of study-based learning was manipulated by varying the number of item repetitions in the initial study phase between one and eight. Predictions of the dual-memory model of test-enhanced learning for the case of one study-phase repetition were used as a reference. Results support the hypothesis that the advantage of testing over restudy is independent of the degree of prior episodic learning, and they suggest that educators can apply cued-recall testing with the expectation that its efficacy is similar across varying levels of prior content learning. Implications for testing effect theory are discussed.

Cover page of Measuring retention within the adolescent brain cognitive development (ABCD)<sup>SM</sup> study.

Measuring retention within the adolescent brain cognitive development (ABCD)SM study.


The Adolescent Brain Cognitive Development (ABCD)SM study aims to retain a demographically diverse sample of youth and one parent across 21 sites throughout its 10-year protocol while minimizing selective (systematic) attrition. To evaluate the effectiveness of these efforts, the ABCD Retention Workgroup (RW) has employed a data-driven approach to examine, track, and intervene via three key metrics: (1) which youth completed visits late; (2) which youth missed visits; and (3) which youth withdrew from the study. The RW actively examines demographic (race, education level, family income) and site factors (visit satisfaction, distance from site, and enrollment in ancillary studies) to strategize efforts that will minimize disengagement and loss of participating youth and parents. Data showed that the most robust primary correlates of late visits were distance from study site, race, and parental education level. Race, lower parental education level, parental employment status, and lower family income were associated with higher odds of missed visits, while being enrolled in one of the ancillary studies was associated with lower odds of missed visits. Additionally, parents who were primary Spanish speakers withdrew at slightly higher rates. These findings provide insight into future targets for proactive retention efforts by the ABCD RW.

Cover page of Information normally considered task-irrelevant drives decision-making and affects premotor circuit recruitment.

Information normally considered task-irrelevant drives decision-making and affects premotor circuit recruitment.


Decision-making is a continuous and dynamic process with prior experience reflected in and used by the brain to guide adaptive behavior. However, most neurobiological studies constrain behavior and/or analyses to task-related variables, not accounting for the continuous internal and temporal space in which they occur. We show mice rely on information learned through recent and longer-term experience beyond just prior actions and reward - including checking behavior and the passage of time - to guide self-initiated, self-paced, and self-generated actions. These experiences are represented in secondary motor cortex (M2) activity and its projections into dorsal medial striatum (DMS). M2 integrates this information to bias strategy-level decision-making, and DMS projections reflect specific aspects of this recent experience to guide actions. This suggests diverse aspects of experience drive decision-making and its neural representation, and shows premotor corticostriatal circuits are crucial for using selective aspects of experiential information to guide adaptive behavior.

Cover page of Deletion of Six3 in post-proliferative neurons produces weakened SCN circadian output, improved metabolic function, and dwarfism in male mice.

Deletion of Six3 in post-proliferative neurons produces weakened SCN circadian output, improved metabolic function, and dwarfism in male mice.



The increasing prevalence of obesity makes it important to increase the understanding of the maturation and function of the neuronal integrators and regulators of metabolic function.


Behavioral, molecular, and physiological analyses of transgenic mice with Sine oculis 3 (Six3) deleted in mature neurons using the Synapsincreallele.


Conditional deletion of the homeodomain transcription factor Six3 in mature neurons causes dwarfism and weakens circadian wheel-running activity rhythms but increases general activity at night, and improves metabolic function, without impacting pubertal onset or fertility in males. The reduced growth in 6-week-old Six3fl/fl:Synapsincre (Six3syn) males correlates with increased somatostatin (SS) expression in the hypothalamus and reduced growth hormone (GH) in the pituitary. In contrast, 12-week-old Six3syn males have increased GH release, despite an increased number of the inhibitory SS neurons in the periventricular nucleus. GH is important in glucose metabolism, muscle function, and bone health. Interestingly, Six3syn males have improved glucose tolerance at 7, 12, and 18 weeks of age, which, in adulthood, is associated with increased % lean mass and increased metabolic rates. Further, 12-week-old Six3syn males have reduced bone mineralization and a lower bone mineral density, indicating that reduced GH levels during early life cause a long-term reduction in bone mineralization.


Our study points to the novel role of Six3 in post-proliferative neurons to regulate metabolic function through SS neuron control of GH release.

Minimizing Iridium Oxide Electrodes for High Visual Acuity Subretinal Stimulation.


Vision loss from diseases of the outer retina, such as age-related macular degeneration, is among the leading causes of irreversible blindness in the world today. The goal of retinal prosthetics is to replace the photo-sensing function of photoreceptors lost in these diseases with optoelectronic hardware to electrically stimulate patterns of retinal activity corresponding to vision. To enable high-resolution retinal prosthetics, the scale of stimulating electrodes must be significantly decreased from current designs; however, this reduces the amount of stimulating current that can be delivered. The efficacy of subretinal stimulation at electrode sizes suitable for high visual acuity retinal prosthesis are not well understood, particularly within the safe charge injection limits of electrode materials. Here, we measure retinal ganglion cell (RGC) responses in a mouse model of blindness to evaluate the stimulation efficacy of 10, 20, and 30 μm diameter iridium oxide electrodes within the electrode charge injection limits, focusing on measures of charge threshold and dynamic range. Stimulation thresholds were lower for smaller electrodes, but larger electrodes could elicit a greater dynamic range of spikes and recruited more ganglion cells within charge injection limits. These findings suggest a practical lower limit for planar electrode size and indicate strategies for maximizing stimulation thresholds and dynamic range.

Cover page of The sex premium in religiously motivated moral judgment.

The sex premium in religiously motivated moral judgment.


Recent theorizing suggests that religious people's moral convictions are quite strategic (albeit unconsciously so), designed to make their worlds more amenable to their favored approaches to solving life's basic challenges. In a meta-analysis of 5 experiments and a preregistered replication, we find that religious identity places a sex premium on moral judgments, causing people to judge violations of conventional sexual morality as particularly objectionable. The sex premium is especially strong among highly religious people, and applies to both legal and illegal acts. Religion's influence on moral reasoning emphasizes conventional sexual norms, and may reflect the strategic projects to which religion has been applied throughout history. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Cover page of Experimental evidence that apologies promote forgiveness by communicating relationship value.

Experimental evidence that apologies promote forgiveness by communicating relationship value.


Robust evidence supports the importance of apologies for promoting forgiveness. Yet less is known about how apologies exert their effects. Here, we focus on their potential to promote forgiveness by way of increasing perceptions of relationship value. We used a method for directly testing these causal claims by manipulating both the independent variable and the proposed mediator. Namely, we use a 2 (Apology: yes vs. no) × 2 (Value: high vs. low) concurrent double-randomization design to test whether apologies cause forgiveness by affecting the same causal pathway as relationship value. In addition to supporting this causal claim, we also find that apologies had weaker effects on forgiveness when received from high-value transgressors, suggesting that the forgiveness-relevant information provided by apologies is redundant with relationship value. Taken together, these findings from a rigorous methodological paradigm help us parse out how apologies promote relationship repair.

Cover page of Target templates in low target-distractor discriminability visual search have higher resolution, but the advantage they provide is short-lived.

Target templates in low target-distractor discriminability visual search have higher resolution, but the advantage they provide is short-lived.


When you search repeatedly for a set of items among very similar distractors, does that make you more efficient in locating the targets? To address this, we had observers search for two categories of targets among the same set of distractors across trials. Visual and conceptual similarity of the stimuli were validated with a multidimensional scaling analysis, and separately using a deep neural network model. After a few blocks of visual search trials, the distractor set was replaced. In three experiments, we manipulated the level of discriminability between the targets and distractors before and after the distractors were replaced. Our results suggest that in the presence of repeated distractors, observers generally become more efficient. However, the difficulty of the search task does impact how efficient people are when the distractor set is replaced. Specifically, when the training is easy, people are more impaired in a difficult transfer test. We attribute this effect to the precision of the target template generated during training. In particular, a coarse target template is created when the target and distractors are easy to discriminate. These coarse target templates do not transfer well in a context with new distractors. This suggests that learning with more distinct targets and distractors can result in lower performance when context changes, but observers recover from this effect quickly (within a block of search trials).

Real-world objects are not stored in holistic representations in visual working memory.


When storing multiple objects in visual working memory, observers sometimes misattribute perceived features to incorrect locations or objects. These misattributions are called binding errors (or swaps) and have been previously demonstrated mostly in simple objects whose features are easy to encode independently and arbitrarily chosen, like colors and orientations. Here, we tested whether similar swaps can occur with real-world objects, where the connection between features is meaningful rather than arbitrary. In Experiments 1 and 2, observers were simultaneously shown four items from two object categories. Within a category, the two exemplars could be presented in either the same or different states (e.g., open/closed; full/empty). After a delay, both exemplars from one of the categories were probed, and participants had to recognize which exemplar went with which state. We found good memory for state information and exemplar information on their own, but a significant memory decrement for exemplar-state combinations, suggesting that binding was difficult for observers and swap errors occurred even for meaningful real-world objects. In Experiment 3, we used the same task, but in one-half of the trials, the locations of the exemplars were swapped at test. We found that there are more errors in general when the locations of exemplars were swapped. We concluded that the internal features of real-world objects are not perfectly bound in working memory, and location updates impair object and feature representations. Overall, we provide evidence that even real-world objects are not stored in an entirely unitized format in working memory.

Cover page of Different Effects of Alcohol Exposure on Action and Outcome-Related Orbitofrontal Cortex Activity.

Different Effects of Alcohol Exposure on Action and Outcome-Related Orbitofrontal Cortex Activity.


Alcohol dependence can result in long-lasting deficits to decision-making and action control. Neurobiological investigations have identified orbitofrontal cortex (OFC) as important for outcome-related contributions to goal-directed actions during decision-making. Prior work has shown that alcohol dependence induces long-lasting changes to OFC function that persist into protracted withdrawal and disrupts goal-directed control over actions. However, it is unclear whether these changes in function alter representation of action and outcome-related neural activity in OFC. Here, we used the well-validated chronic intermittent ethanol (CIE) exposure and withdrawal procedure to model alcohol dependence in mice and performed in vivo extracellular recordings during an instrumental task in which lever-press actions made for a food outcome. We found alcohol dependence disrupted goal-directed action control and increased OFC activity associated with lever-pressing but decreased OFC activity during outcome-related epochs. The ability to decode outcome-related information, but not action information, from OFC activity following CIE exposure was reduced. Hence, chronic alcohol exposure induced a long-lasting disruption to OFC function such that activity associated with actions was enhanced, but OFC activity contributions to outcome-related information was diminished. This has important implications for hypotheses regarding compulsive and habitual phenotypes observed in addiction.