{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:03:22Z","timestamp":1792785802669,"version":"4.5.0"},"reference-count":62,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"am","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/100009633","name":"Eunice Kennedy Shriver National Institute of Child Health and Human Development","doi-asserted-by":"publisher","award":["R01HD110170"],"award-info":[{"award-number":["R01HD110170"]}],"id":[{"id":"10.13039\/100009633","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Thomas Harriot College of Arts and Sciences at East Carolina University","id":[{"id":"https:\/\/ror.org\/01vx35703","id-type":"ROR","asserted-by":"crossref"}]},{"name":"East Carolina University Research and Economic Development Office","id":[{"id":"https:\/\/ror.org\/01vx35703","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Mammalian sperm must undergo capacitation, a process marked by substantial phenotypic heterogeneity among sperm. How heterogeneity relates to fertilizing potential remains unresolved, in part because sperm subpopulations are typically analyzed independently despite being intrinsically interdependent. We combine flow cytometry with statistical modeling to quantify how sperm subpopulation compositions respond to a range of capacitation signal intensity in vitro. Using cauda epididymal mouse sperm, we systematically varied extracellular bicarbonate and free calcium\u2014key regulators of capacitation\u2014and classified millions of individual cells into four irreversible physiological states. We show that bicarbonate and calcium interact nonlinearly to redistribute sperm across these subpopulations, revealing responses that would be otherwise obscured in measurements lacking single-cell resolution. Elevated intracellular calcium was associated with increased cell death, while the highest proportions of live, acrosome-reacted sperm occurred under relatively low extracellular calcium conditions. We then applied a hierarchical Dirichlet\u2013multinomial regression model that accounts for multinomial sampling noise and between-male variability, yielding posterior probability surfaces that describe how sperm subpopulations reallocate across functional states as signaling conditions change. We then tested the model predictions using IVF and found that the proportion of live, acrosome reacted sperm was not predictive of early embryonic developmental success. Together, these results suggest that capacitation, though tightly regulated at the single cell scale, is a stochastic process shaped by structured phenotypic heterogeneity at the cell population scale. This framework provides a quantitative foundation for linking sperm subpopulation composition to measures of fertility competence.<\/jats:p>","DOI":"10.1093\/biolre\/ioag207","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T12:01:00Z","timestamp":1790164860000},"source":"Crossref","is-referenced-by-count":0,"title":["Phenotypic heterogeneity during sperm capacitation: a compositional analysis"],"prefix":"10.1093","author":[{"given":"Benjamin M","family":"Brisard","sequence":"first","affiliation":[{"name":"Department of Biology, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aishwarya P","family":"Halder","sequence":"additional","affiliation":[{"name":"Department of Biology, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aidan","family":"Charles","sequence":"additional","affiliation":[{"name":"Department of Biology, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amy L","family":"Ward","sequence":"additional","affiliation":[{"name":"Department of Biology, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maryam","family":"Asadi","sequence":"additional","affiliation":[{"name":"Department of Physics, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David","family":"Hart","sequence":"additional","affiliation":[{"name":"Department of Computer Science, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Debajit","family":"Bhowmick","sequence":"additional","affiliation":[{"name":"Flow Cytometry and Cell Sorting Shared Resource (FCCSSR), Saint Jude Children's Hospital , Memphis, TN ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul","family":"Vos","sequence":"additional","affiliation":[{"name":"Department of Public Health, Brody School of Medicine, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6628-6329","authenticated-orcid":false,"given":"Cameron A","family":"Schmidt","sequence":"additional","affiliation":[{"name":"Department of Biology, East Carolina University , Greenville, NC ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"2026100514071489200_ref1","doi-asserted-by":"publisher","first-page":"127","DOI":"10.1093\/biolre\/ioy045","article-title":"Sperm acrosome reaction: its site and role in fertilization","volume":"99","author":"Hirohashi","year":"2018","journal-title":"Biol Reprod"},{"key":"2026100514071489200_ref2","doi-asserted-by":"publisher","first-page":"860","DOI":"10.1002\/mrd.22663","article-title":"Chang\u2019s meaning of capacitation: a molecular perspective","volume":"83","author":"Gervasi","year":"2016","journal-title":"Mol Reprod Dev"},{"key":"2026100514071489200_ref3","doi-asserted-by":"publisher","first-page":"1041","DOI":"10.1016\/j.febslet.2010.02.005","article-title":"The SLO3 sperm-specific potassium channel plays a vital role in male fertility","volume":"584","author":"Santi","year":"2010","journal-title":"FEBS Lett"},{"key":"2026100514071489200_ref4","doi-asserted-by":"publisher","first-page":"1149","DOI":"10.1016\/j.bbrc.2014.05.100","article-title":"Intracellular pH in sperm physiology","volume":"450","author":"Nishigaki","year":"2014","journal-title":"Biochem Biophys Res Commun"},{"key":"2026100514071489200_ref5","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/j.devcel.2005.06.007","article-title":"The \u2018soluble\u2019 adenylyl cyclase in sperm mediates multiple signaling events required for fertilization","volume":"9","author":"Hess","year":"2005","journal-title":"Dev Cell"},{"key":"2026100514071489200_ref6","doi-asserted-by":"publisher","first-page":"1129","DOI":"10.1242\/dev.121.4.1129","article-title":"Capacitation of mouse spermatozoa I. 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