---
title: "Rem2 in CA1 Controls Sex-Dependent Spatial Learning via Glutamate Receptor Composition"
id: "biorxiv-16-the-small-gtpase-rem2-modulates-sex-dependent-spatial-learning-by-regulating"
canonical_url: "https://medichelpline.com/clinical-feed/biorxiv-16-the-small-gtpase-rem2-modulates-sex-dependent-spatial-learning-by-regulating"
content_type: "clinical_feed_article"
specialty: "Neurology"
source_name: "bioRxiv (Biomedical Preprints)"
source_url: "https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1?rss=1"
published_at: "2026-09-22T12:00:00.000Z"
evidence_level: "Verified Feed"
license: "CC-BY-NC-4.0 / Informational Use"
---
# Rem2 in CA1 Controls Sex-Dependent Spatial Learning via Glutamate Receptor Composition
## Provenance & Clinical Metadata
- **Canonical URL:** https://medichelpline.com/clinical-feed/biorxiv-16-the-small-gtpase-rem2-modulates-sex-dependent-spatial-learning-by-regulating
- **Specialty:** [Neurology](https://medichelpline.com/clinical-feed/neurology.md)
- **Primary Source:** bioRxiv (Biomedical Preprints)
- **Source URL:** [Original Journal Publication](https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1?rss=1)
- **Published At:** 2026-09-22T12:00:00.000Z
- **Evidence Rating:** Verified Feed
## Executive GIST (TL;DR)
- The small GTPase **Rem2** is a signaling molecule that influences synapse formation, dendritic and spine complexity, and CaMKII-dependent long-term potentiation; its role in learning and memory was previously unclear. - The authors deleted Rem2 selectively in dorsal **CA1** neurons in male and female mice to test effects on spatial learning using a spatial object recognition (SOR) task. - In baseline SOR behavior, male mice outperformed female mice; deleting Rem2 did not alter male performance but significantly improved performance in females so they matched males. - Protein profiling used an automated Western blot system to quantify all known **AMPAR** and **NMDAR** subunits in each mouse, enabling subunit-level comparisons across sex and genotype. - The sex-dependent behavioral change correlated with a Rem2-driven increase in **GluN2D** expression localized to interneurons in female mice only; this change was not observed in males. - Multivariate comparison using S-statistics showed that males and females normally exhibit different covariance structures across AMPAR and NMDAR composition; these sex differences in covariance were abolished when Rem2 was deleted from CA1 neurons. - The results indicate Rem2 maintains sex differences in glutamate receptor composition and associated spatial learning, making it the first reported signaling molecule to confer sexual dimorphism at excitatory synapses according to the authors. - The findings suggest Rem2 could be relevant to understanding sex-dependent features of neurodevelopmental and neurodegenerative disorders; the study is a preprint and has not been peer reviewed. - Funding was declared from the Charles E. Kaufman Foundation (KA2023-136491); authors declared no competing interests.
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Brew, Hassan E. Mohammed, Afrina Asad Meghla, Serendipity Coniglio, Eleanor Labriola, Regan Skudlarek, Kishore Kumar S. Narasimhan, Shashank M. Dravid, Gillian Queisser, Anna R. Moore, Victor M. Luna doi: https://doi.org/10.64898/2026.09.15.751881 This article is a preprint and has not been certified by peer review [[what does this mean?](https://www.biorxiv.org/about/FAQ#unrefereed)]. William E. Brew 1 Department of Biology, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=William%2BE.%2BBrew%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Brew%20WE&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AWilliam%2BE.%2BBrew%2B) Hassan E. Mohammed 2 Alzheimers Center at Temple, Department of Neural Sciences, Lewis Katz School of Medicine, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Hassan%2BE.%2BMohammed%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Mohammed%20HE&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AHassan%2BE.%2BMohammed%2B) Afrina Asad Meghla 3 Department of Mathematics, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Afrina%2BAsad%2BMeghla%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Meghla%20AA&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AAfrina%2BAsad%2BMeghla%2B) Serendipity Coniglio 2 Alzheimers Center at Temple, Department of Neural Sciences, Lewis Katz School of Medicine, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Serendipity%2BConiglio%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Coniglio%20S&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3ASerendipity%2BConiglio%2B) Eleanor Labriola 1 Department of Biology, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Eleanor%2BLabriola%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Labriola%20E&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AEleanor%2BLabriola%2B) Regan Skudlarek 1 Department of Biology, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Regan%2BSkudlarek%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Skudlarek%20R&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3ARegan%2BSkudlarek%2B) Kishore Kumar S. 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Moore 1 Department of Biology, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Anna%2BR.%2BMoore%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Moore%20AR&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AAnna%2BR.%2BMoore%2B) Victor M. Luna 2 Alzheimers Center at Temple, Department of Neural Sciences, Lewis Katz School of Medicine, Temple University; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Victor%2BM.%2BLuna%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Luna%20VM&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AVictor%2BM.%2BLuna%2B) * For correspondence: victor.luna@temple.edu * [Abstract](https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1)[](https://www.biorxiv.org/panels_ajax_tab/biorxiv_tab_art/node:5801075/1) * [Info/History](https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1.article-info)[](https://www.biorxiv.org/panels_ajax_tab/biorxiv_tab_info/node:5801075/1) * [Metrics](https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1.article-metrics)[](https://www.biorxiv.org/panels_ajax_tab/article_tab_metrics/node:5801075/1) * [ Preview PDF](https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1.full.pdf+html)[](https://www.biorxiv.org/panels_ajax_tab/biorxiv_tab_pdf/node:5801075/1) ![Loading](https://www.biorxiv.org/sites/all/modules/contrib/panels_ajax_tab/images/loading.gif) ## Abstract The small GTPase Rem2 is a key signaling molecule involved in synaptic formation, dendritic complexity, spine formation, and regulation of CaMKII-dependent long-term potentiation (LTP). However, it remains unclear how Rem2 explicitly impacts learning and memory. To address this issue, we deleted Rem2 specifically in dorsal CA1 neurons of male and female mice and assessed spatial learning using a spatial object recognition (SOR) task. We found that males outperform females in this behavioral assay and that deleting Rem2 did not impact spatial learning in males. In contrast, Rem2 deletion significantly improved spatial learning in females enabling them to perform as well as males in the SOR task. Using an automated Western blot system to assess all known AMPA- and NMDA-mediated glutamate receptor (AMPAR and NMDAR) subunits in each mouse, we found that the sex-dependent change in SOR we observed was likely due to Rem2 increasing GluN2D expression in the interneurons of female mice only. Furthermore, using S-statistics to compare the overall AMPAR and NMDAR composition among groups, we found that males and females normally have divergent covariance structures but this divergence is eliminated when Rem2 is deleted from CA1 neurons. These results reveal an unexpected role of Rem2 in maintaining sex differences in glutamate receptor composition and spatial learning abilities. To our knowledge, this is the first demonstration of a signaling molecule that confers sexual dimorphism to excitatory synapses. As such, Rem2 may play a critical role in understanding how sex-dependent symptoms of neurodevelopmental and neurodegenerative disorders arise. ### Competing Interest Statement The authors have declared no competing interest. ## Funder Information Declared Charles E. Kaufman Foundation, KA2023-136491 Copyright The copyright holder for this preprint is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. All rights reserved. No reuse allowed without permission. bioRxiv and medRxiv thank the following for their generous financial support: > The Chan Zuckerberg Initiative, Cold Spring Harbor Laboratory, the Sergey Brin Family Foundation, California Institute of Technology, Centre National de la Recherche Scientifique, Fred Hutchinson Cancer Center, Imperial College London, Massachusetts Institute of Technology, Stanford University, The University of Edinburgh, University of Washington, and Vrije Universiteit Amsterdam. [Donate to openRxiv ](https://www.zeffy.com/en-US/donation-form/donate-to-make-a-difference-10981) [ Back to top](https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1?rss=1#page) [ Previous](https://www.biorxiv.org/content/10.64898/2026.09.13.751220v2 "Categorical Color Structure Persists in Color Vision Deficiency")[Next ](https://www.biorxiv.org/content/10.64898/2026.09.20.752975v1 "INO80 rapidly shuttles nucleosomes between chromatin barriers") Posted September 22, 2026. [ Download PDF](https://www.biorxiv.org/content/10.64898/2026.09.15.751881v1.full.pdf) Print/Save Options [Download PDF](https://www.biorxiv.org/content/biorxiv/early/2026/09/22/2026.09.15.751881.full.pdf)Full Text & In-line FiguresXML [More Info](https://www.biorxiv.org/about/FAQ#PrintOptions "More Information on Print/Save Options") [ Email](https://www.biorxiv.org/ "Email this Article") [ Share](https://www.biorxiv.org/) The small GTPase Rem2 modulates sex-dependent spatial learning by regulating CA1 glutamate receptor composition William E. Brew, Hassan E. Mohammed, Afrina Asad Meghla, Serendipity Coniglio, Eleanor Labriola, Regan Skudlarek, Kishore Kumar S. Narasimhan, Shashank M. Dravid, Gillian Queisser, Anna R. Moore, Victor M. Luna bioRxiv 2026.09.15.751881; doi: https://doi.org/10.64898/2026.09.15.751881 This article is a preprint and has not been certified by peer review [[what does this mean?](https://www.biorxiv.org/about/FAQ#unrefereed)]. 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