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However, how amino acid protonation alters protein hydration and solubility is an open question since Kuntz and Kauzmann proposed p\n                    <jats:italic toggle=\"yes\">H<\/jats:italic>\n                    -driven protein desolvation in 1974. Here, in a series of high-resolution cryoelectron microscopy structures of a protein complex at different p\n                    <jats:italic toggle=\"yes\">H<\/jats:italic>\n                    values (from p\n                    <jats:italic toggle=\"yes\">H<\/jats:italic>\n                    9.0 to 3.5), we examined thousands of observable hydration sites. Cryoelectron microscopy data, in agreement with constant-p\n                    <jats:italic toggle=\"yes\">H<\/jats:italic>\n                    molecular dynamics simulations, show that nearly half of protein-bound waters exchanged with the bulk solvent upon acidification, with ~100 waters lost per p\n                    <jats:italic toggle=\"yes\">H<\/jats:italic>\n                    unit per molecule. The loss of waters was most significant around the side chains of glutamate and aspartate residues while specific polar residues, mostly asparagine, anchored persistent waters. A positionally conserved hydration layer was observed across all p\n                    <jats:italic toggle=\"yes\">H<\/jats:italic>\n                    conditions, accounting for 40% of resolved waters. Those waters displayed denser packing than less persistent waters, forming a p\n                    <jats:italic toggle=\"yes\">H<\/jats:italic>\n                    -independent solvation shell. Acid-induced water exchange also displaced bound iron, providing a mechanistic link between solvation and metal release. Our findings demonstrate the core principles of acid-driven protein desolvation, resolving a 50-y-old biochemical hypothesis.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2525949123","type":"journal-article","created":{"date-parts":[[2026,3,5]],"date-time":"2026-03-05T18:58:39Z","timestamp":1772737119000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":6,"title":["Direct evidence of acid-driven protein desolvation"],"prefix":"10.1073","volume":"123","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2155-5000","authenticated-orcid":false,"given":"Farzad","family":"Hamdi","sequence":"first","affiliation":[{"name":"Department of Integrative Structural Biochemistry, Institute of Biochemistry and Biotechnology, Martin Luther University Halle-Wittenberg","place":["Halle\/Saale, Germany"]},{"name":"Interdisciplinary Research Center HALOmem, Charles Tanford Protein Center, Martin Luther University 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