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While prior work has focused on restricted\/hindered diffusion and membrane permeation, additional exchange mechanisms such as diffusion\u2010mediated exchange between dendritic shaft and spines in gray matter (GM) remain understudied. Here, we hypothesize that water diffusion within impermeable spiny dendrites can produce time\u2010dependent SDE signals indistinguishable from those arising from permeative exchange; and assess to what extent spine density impacts estimates of exchange time. Using Monte Carlo simulations and analytical solutions from the narrow escape problem, we quantify spine\u2013shaft and shaft\u2013spine exchange times, revealing characteristic times (1\u201350\u2009ms) comparable to permeative exchange estimates in the cortex. We show that a modified two\u2010compartment K\u00e4rger model accurately captures the time\u2010dependent SDE signal along spiny dendrites but yields exchange estimates that reflect total spine volume fraction rather than specific spine morphology. Simulations reveal that unaccounted diffusion\u2010mediated exchange from dendritic spines substantially biases Neurite Exchange Imaging (NEXI) estimates, inflating the apparent exchange rate and the inferred membrane permeability in proportion to spine volume fraction. Further, we propose an extended three\u2010compartment K\u00e4rger model (and its coarse\u2010grained version) incorporating both diffusion\u2010mediated exchange between dendritic shaft and spines and permeative exchange with extracellular space. Critically, while the three\u2010compartment K\u00e4rger model and its coarse\u2010grained version capture both exchange mechanisms, they cannot uniquely disentangle membrane permeability from spine volume fraction. Finally, we highlight the importance of accounting for dendritic spines when inferring membrane permeability from diffusion MRI exchange rates, as the additional membrane area contributed by spines increases the effective dendritic surface\u2010to\u2010volume ratio by approximately 24%\u201360% for realistic spine densities and, if neglected, leads to a corresponding overestimation of the true membrane permeability. These findings underscore the necessity of considering dendritic spine contributions when interpreting time\u2010dependent SDE data and caution against attributing exchange effects solely to membrane permeability. Our study further highlights the need for advanced acquisition and modeling approaches to differentiate permeative and diffusion\u2010mediated geometric exchange in GM.<\/jats:p>","DOI":"10.1002\/nbm.70382","type":"journal-article","created":{"date-parts":[[2026,9,12]],"date-time":"2026-09-12T00:24:46Z","timestamp":1789172686000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["The Role of Dendritic Spines in Water Exchange Measurements With Diffusion MRI: Time\u2010Dependent Single Diffusion Encoding MRI"],"prefix":"10.1002","volume":"39","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4044-7395","authenticated-orcid":false,"given":"Kadir","family":"\u015eim\u015fek","sequence":"first","affiliation":[{"name":"Cardiff University Brain Research Imaging Centre (CUBRIC) Cardiff University  Cardiff UK"},{"name":"School of Computer Science and Informatics Cardiff University  Cardiff 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