{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,30]],"date-time":"2026-10-30T04:04:21Z","timestamp":1793333061833,"version":"4.5.0"},"reference-count":120,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T00:00:00Z","timestamp":1782864000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,5,8]],"date-time":"2026-05-08T00:00:00Z","timestamp":1778198400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["BioSystems"],"published-print":{"date-parts":[[2026,7]]},"DOI":"10.1016\/j.biosystems.2026.105812","type":"journal-article","created":{"date-parts":[[2026,5,9]],"date-time":"2026-05-09T00:00:34Z","timestamp":1778284834000},"page":"105812","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":1,"special_numbering":"C","title":["ARCH \u00d7 \u03a6: Sterols and the evolution of gated biological execution"],"prefix":"10.1016","volume":"265","author":[{"given":"Tahir","family":"Rahman","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Charles F.","family":"Zorumski","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"issue":"50","key":"10.1016\/j.biosystems.2026.105812_bib1","doi-asserted-by":"crossref","first-page":"11605","DOI":"10.1523\/JNEUROSCI.4173-05.2005","article-title":"Neurosteroid access to the GABAA receptor","volume":"25","author":"Akk","year":"2005","journal-title":"J. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib2","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1126\/science.1165939","article-title":"Serotonin mediates behavioral gregarization underlying swarm formation in desert locusts","volume":"323","author":"Anstey","year":"2009","journal-title":"Science"},{"key":"10.1016\/j.biosystems.2026.105812_bib3","first-page":"2636","article-title":"AP180-mediated inhibition of KCNQ2\/3 channels underlies neuronal hyperexcitability in niemann-pick type C1 disease neurons","volume":"27","author":"Bao","year":"2019","journal-title":"Cell Rep."},{"issue":"2","key":"10.1016\/j.biosystems.2026.105812_bib4","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1007\/s13752-013-0155-6","article-title":"From biosemiotics to code biology","volume":"9","author":"Barbieri","year":"2014","journal-title":"Biological Theory"},{"key":"10.1016\/j.biosystems.2026.105812_bib5","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.biosystems.2017.10.005","article-title":"What is code biology?","volume":"164","author":"Barbieri","year":"2018","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105812_bib6","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1016\/j.cels.2016.01.001","article-title":"A dynamical framework for the all-or-none G1\/S transition","volume":"2","author":"Barr","year":"2016","journal-title":"Cell Syst."},{"key":"10.1016\/j.biosystems.2026.105812_bib7","doi-asserted-by":"crossref","first-page":"963","DOI":"10.1016\/S0306-4530(98)00071-7","article-title":"Neurosteroids: a novel function of the brain","volume":"23","author":"Baulieu","year":"1998","journal-title":"Psychoneuroendocrinology"},{"key":"10.1016\/j.biosystems.2026.105812_bib8","doi-asserted-by":"crossref","first-page":"11893","DOI":"10.1073\/pnas.1513318112","article-title":"The genome of Aiptasia, a sea anemone model for coral symbiosis","volume":"112","author":"Baumgarten","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105812_bib9","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1002\/(SICI)1520-6327(199909)42:1<13::AID-ARCH3>3.0.CO;2-P","article-title":"Phytosterol metabolism and absorption in the generalist grasshopper, Schistocerca americana (Orthoptera: Acrididae)","volume":"42","author":"Behmer","year":"1999","journal-title":"Arch. Insect Biochem. Physiol."},{"key":"10.1016\/j.biosystems.2026.105812_bib10","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/S0022-1910(98)00131-0","article-title":"The nutritional significance of sterol metabolic constraints in the generalist grasshopper Schistocerca americana","volume":"45","author":"Behmer","year":"1999","journal-title":"J. Insect Physiol."},{"issue":"1","key":"10.1016\/j.biosystems.2026.105812_bib11","doi-asserted-by":"crossref","first-page":"1326","DOI":"10.1038\/s42003-024-06991-z","article-title":"Functional MRI of imprinting memory in awake newborn domestic chicks","volume":"7","author":"Behroozi","year":"2024","journal-title":"Commun. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib12","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1146\/annurev.biochem.71.110601.135425","article-title":"DNA replication in eukaryotic cells","volume":"71","author":"Bell","year":"2002","journal-title":"Annu. Rev. Biochem."},{"key":"10.1016\/j.biosystems.2026.105812_bib13","doi-asserted-by":"crossref","first-page":"309","DOI":"10.1016\/S0165-0173(98)00019-8","article-title":"What is the role of dopamine in reward: hedonic impact, reward learning, or incentive salience?","volume":"28","author":"Berridge","year":"1998","journal-title":"Brain Res. Rev."},{"key":"10.1016\/j.biosystems.2026.105812_bib14","doi-asserted-by":"crossref","first-page":"47","DOI":"10.3109\/10409238309102790","article-title":"Sterol structure and membrane function","volume":"14","author":"Bloch","year":"1983","journal-title":"CRC Crit. Rev. Biochem."},{"key":"10.1016\/j.biosystems.2026.105812_bib15","doi-asserted-by":"crossref","first-page":"1014","DOI":"10.1038\/ncb1101-1014","article-title":"Akt\/mTOR pathway is a crucial regulator of skeletal muscle hypertrophy and can prevent muscle atrophy in vivo","volume":"3","author":"Bodine","year":"2001","journal-title":"Nat. Cell Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib16","doi-asserted-by":"crossref","first-page":"3637","DOI":"10.1152\/jn.00686.2005","article-title":"Adaptive exponential integrate-and-fire model as an effective description of neuronal activity","volume":"94","author":"Brette","year":"2005","journal-title":"J. Neurophysiol."},{"key":"10.1016\/j.biosystems.2026.105812_bib17","doi-asserted-by":"crossref","first-page":"578","DOI":"10.1038\/nature23457","article-title":"The rise of algae in Cryogenian oceans and the emergence of animals","volume":"548","author":"Brocks","year":"2017","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib18","doi-asserted-by":"crossref","first-page":"598","DOI":"10.3390\/md18120598","article-title":"Sterol composition of sponges, cnidarians, arthropods, mollusks, and echinoderms from the deep northwest Atlantic: a comparison with shallow coastal Gulf of Mexico","volume":"18","author":"Carre\u00f3n-Palau","year":"2020","journal-title":"Mar. Drugs"},{"key":"10.1016\/j.biosystems.2026.105812_bib19","doi-asserted-by":"crossref","first-page":"1306","DOI":"10.1126\/science.aag1417","article-title":"Lysosomal cholesterol activates mTORC1 via an SLC38A9-Niemann-Pick C1 signaling complex","volume":"355","author":"Castellano","year":"2017","journal-title":"Science"},{"key":"10.1016\/j.biosystems.2026.105812_bib20","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1016\/S0005-2736(97)00075-8","article-title":"Effect of lipid structure on the dipole potential of phosphatidylcholine bilayers","volume":"1327","author":"Clarke","year":"1997","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.biosystems.2026.105812_bib21","doi-asserted-by":"crossref","first-page":"E10216","DOI":"10.1073\/pnas.1713897114","article-title":"Definition of the hypothalamic GnRH pulse generator in mice","volume":"114","author":"Clarkson","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105812_bib22","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.02882","article-title":"Three pools of plasma membrane cholesterol and their relation to cholesterol homeostasis","volume":"3","author":"Das","year":"2014","journal-title":"eLife"},{"key":"10.1016\/j.biosystems.2026.105812_bib23","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1093\/gbe\/evp036","article-title":"Phylogenomics of sterol synthesis: insights into the origin, evolution, and diversity of a key eukaryotic feature","volume":"1","author":"Desmond","year":"2009","journal-title":"Genome Biol. Evol."},{"key":"10.1016\/j.biosystems.2026.105812_bib24","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1007\/s00429-008-0189-x","article-title":"Brain structural and functional abnormalities in mood disorders: implications for neurocircuitry models of depression","volume":"213","author":"Drevets","year":"2008","journal-title":"Brain Struct. Funct."},{"key":"10.1016\/j.biosystems.2026.105812_bib25","doi-asserted-by":"crossref","first-page":"140","DOI":"10.1016\/S0955-0674(02)00314-9","article-title":"Self-perpetuating states in signal transduction: positive feedback, double-negative feedback and bistability","volume":"14","author":"Ferrell","year":"2002","journal-title":"Curr. Opin. Cell Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib26","doi-asserted-by":"crossref","first-page":"4849","DOI":"10.1523\/JNEUROSCI.17-12-04849.1997","article-title":"Dynamic changes in nucleus accumbens dopamine efflux during the Coolidge effect in male rats","volume":"17","author":"Fiorino","year":"1997","journal-title":"J. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib27","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1016\/S0006-3495(61)86902-6","article-title":"Impulses and physiological states in theoretical models of nerve membrane","volume":"1","author":"FitzHugh","year":"1961","journal-title":"Biophys. J."},{"key":"10.1016\/j.biosystems.2026.105812_bib28","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1038\/266830a0","article-title":"Monogamy and sex change by aggressive dominance in coral reef fish","volume":"266","author":"Fricke","year":"1977","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib29","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1038\/nrn2787","article-title":"The free-energy principle: a unified brain theory?","volume":"11","author":"Friston","year":"2010","journal-title":"Nat. Rev. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib30","first-page":"1693","article-title":"The Parkinson's puzzle","volume":"266","author":"Glickstein","year":"1999","journal-title":"Proc. Roy. Soc. Lond. B"},{"key":"10.1016\/j.biosystems.2026.105812_bib31","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1038\/nature21412","article-title":"Paleoproterozoic sterol biosynthesis and the rise of oxygen","volume":"543","author":"Gold","year":"2017","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib32","series-title":"The Chimpanzees of Gombe: Patterns of Behavior","author":"Goodall","year":"1986"},{"key":"10.1016\/j.biosystems.2026.105812_bib33","doi-asserted-by":"crossref","first-page":"1561","DOI":"10.1016\/j.bpj.2012.03.004","article-title":"Differential effect of cholesterol and its biosynthetic precursors on membrane dipole potential","volume":"102","author":"Haldar","year":"2012","journal-title":"Biophys. J."},{"key":"10.1016\/j.biosystems.2026.105812_bib34","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.43923","article-title":"Sterol transfer by atypical cholesterol-binding NPC2 proteins in coral-algal symbiosis","volume":"8","author":"Hambleton","year":"2019","journal-title":"eLife"},{"key":"10.1016\/j.biosystems.2026.105812_bib35","doi-asserted-by":"crossref","first-page":"338","DOI":"10.1038\/nrrheum.2014.15","article-title":"Diagnosis and management of neuropsychiatric SLE","volume":"10","author":"Hanly","year":"2014","journal-title":"Nat. Rev. Rheumatol."},{"key":"10.1016\/j.biosystems.2026.105812_bib36","doi-asserted-by":"crossref","first-page":"R29","DOI":"10.1016\/j.cub.2006.11.042","article-title":"Rapid conduction and the evolution of giant axons and myelinated fibers","volume":"17","author":"Hartline","year":"2007","journal-title":"Curr. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib37","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.88900.3","article-title":"Molecular tuning of sea anemone stinging","volume":"12","author":"He","year":"2023","journal-title":"eLife"},{"key":"10.1016\/j.biosystems.2026.105812_bib38","first-page":"118","article-title":"Accessible cholesterol is a key regulator of interferon-gamma signaling","volume":"38","author":"Heiser","year":"2023","journal-title":"Cell Metab."},{"key":"10.1016\/j.biosystems.2026.105812_bib39","doi-asserted-by":"crossref","first-page":"877","DOI":"10.1038\/nrn1787","article-title":"Critical period plasticity in local cortical circuits","volume":"6","author":"Hensch","year":"2005","journal-title":"Nat. Rev. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib40","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1038\/scientificamerican0358-81","article-title":"Imprinting in animals","volume":"198","author":"Hess","year":"1958","journal-title":"Sci. Am."},{"key":"10.1016\/j.biosystems.2026.105812_bib41","doi-asserted-by":"crossref","first-page":"500","DOI":"10.1113\/jphysiol.1952.sp004764","article-title":"A quantitative description of membrane current and its application to conduction and excitation in nerve","volume":"117","author":"Hodgkin","year":"1952","journal-title":"J. Physiol."},{"key":"10.1016\/j.biosystems.2026.105812_bib42","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1126\/science.6695186","article-title":"An ultrahigh-speed analysis of exocytosis: nematocyst discharge","volume":"223","author":"Holstein","year":"1984","journal-title":"Science"},{"key":"10.1016\/j.biosystems.2026.105812_bib43","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1038\/nrn1324","article-title":"Pathways of the past: the imprint of memory","volume":"5","author":"Horn","year":"2004","journal-title":"Nat. Rev. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib44","series-title":"Principles of Behavior","author":"Hull","year":"1943"},{"key":"10.1016\/j.biosystems.2026.105812_bib45","doi-asserted-by":"crossref","first-page":"185","DOI":"10.1254\/jjp.80.185","article-title":"Mechanism underlying the therapeutic effects of electroconvulsive therapy (ECT) on depression","volume":"80","author":"Ishihara","year":"1999","journal-title":"Jpn. J. Pharmacol."},{"key":"10.1016\/j.biosystems.2026.105812_bib46","doi-asserted-by":"crossref","first-page":"1569","DOI":"10.1109\/TNN.2003.820440","article-title":"Simple model of spiking neurons","volume":"14","author":"Izhikevich","year":"2003","journal-title":"IEEE Trans. Neural Network."},{"key":"10.1016\/j.biosystems.2026.105812_bib47","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1146\/annurev-ento-011019-025017","article-title":"Insect sterol nutrition: physiological mechanisms, ecology, and applications","volume":"65","author":"Jing","year":"2020","journal-title":"Annu. Rev. Entomol."},{"key":"10.1016\/j.biosystems.2026.105812_bib48","doi-asserted-by":"crossref","first-page":"2210","DOI":"10.1093\/brain\/awu133","article-title":"On the nature of seizure dynamics","volume":"137","author":"Jirsa","year":"2014","journal-title":"Brain"},{"key":"10.1016\/j.biosystems.2026.105812_bib49","doi-asserted-by":"crossref","first-page":"1298","DOI":"10.1021\/acschemneuro.3c00823","article-title":"Serotonin signaling through lipid membranes","volume":"15","author":"Kalinichenko","year":"2024","journal-title":"ACS Chem. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib50","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1242\/jeb.201.6.837","article-title":"Rattlesnake strike behavior: kinematics","volume":"201","author":"Kardong","year":"1998","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib51","series-title":"Cholesterol Systems in Insects and Animals","first-page":"1","article-title":"Sterols and insects","author":"Kircher","year":"1981"},{"key":"10.1016\/j.biosystems.2026.105812_bib52","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1016\/j.bbabio.2011.04.013","article-title":"The orange carotenoid protein in photoprotection of photosystem II in Cyanobacteria","volume":"1817","author":"Kirilovsky","year":"2012","journal-title":"Biochim. Biophys. Acta"},{"key":"10.1016\/j.biosystems.2026.105812_bib53","first-page":"827","article-title":"The significance of the Edinger-Westphal nucleus in non-visual function","volume":"32","author":"Kozicz","year":"2011","journal-title":"Peptides"},{"key":"10.1016\/j.biosystems.2026.105812_bib54","doi-asserted-by":"crossref","first-page":"857","DOI":"10.1126\/science.aai8878","article-title":"Improving photosynthesis and crop productivity by accelerating recovery from photoprotection","volume":"354","author":"Kromdijk","year":"2016","journal-title":"Science"},{"key":"10.1016\/j.biosystems.2026.105812_bib55","series-title":"Principles of Topological Psychology","author":"Lewin","year":"1936"},{"key":"10.1016\/j.biosystems.2026.105812_bib56","doi-asserted-by":"crossref","first-page":"391","DOI":"10.1038\/35000131","article-title":"A pigment-binding protein essential for regulation of photosynthetic light harvesting","volume":"403","author":"Li","year":"2000","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib57","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1038\/nature02373","article-title":"Crystal structure of spinach major light-harvesting complex at 2.72 \u00c5 resolution","volume":"428","author":"Liu","year":"2004","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib58","series-title":"The Pupil: Anatomy, Physiology, and Clinical Applications","author":"Loewenfeld","year":"1993"},{"key":"10.1016\/j.biosystems.2026.105812_bib59","series-title":"On Aggression","author":"Lorenz","year":"1963"},{"key":"10.1016\/j.biosystems.2026.105812_bib60","first-page":"961","article-title":"Association between early life adversity and children's brain structure and cognitive function","volume":"170","author":"Luby","year":"2016","journal-title":"JAMA Pediatr."},{"key":"10.1016\/j.biosystems.2026.105812_bib61","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1124\/mol.105.013573","article-title":"Capsaicin regulates voltage-dependent sodium channels by altering lipid bilayer elasticity","volume":"68","author":"Lundb\u00e6k","year":"2004","journal-title":"Mol. Pharmacol."},{"key":"10.1016\/j.biosystems.2026.105812_bib62","doi-asserted-by":"crossref","first-page":"38770","DOI":"10.1074\/jbc.M400831200","article-title":"Partitioning of the serotonin transporter into lipid microdomains modulates transport of serotonin","volume":"279","author":"Magnani","year":"2004","journal-title":"J. Biol. Chem."},{"key":"10.1016\/j.biosystems.2026.105812_bib63","doi-asserted-by":"crossref","DOI":"10.1016\/j.biosystems.2025.105516","article-title":"From code to archetype: toward a unified theory of biological, neural, and artificial artifacts","volume":"254","author":"Major","year":"2025","journal-title":"Biosystems"},{"key":"10.1016\/j.biosystems.2026.105812_bib64","doi-asserted-by":"crossref","first-page":"651","DOI":"10.1016\/j.neuron.2005.02.014","article-title":"Deep brain stimulation for treatment-resistant depression","volume":"45","author":"Mayberg","year":"2005","journal-title":"Neuron"},{"key":"10.1016\/j.biosystems.2026.105812_bib65","doi-asserted-by":"crossref","first-page":"1058","DOI":"10.1016\/S0140-6736(18)31551-4","article-title":"Brexanolone injection in post-partum depression: two multicentre, double-blind, randomised, placebo-controlled, phase 3 trials","volume":"392","author":"Meltzer-Brody","year":"2018","journal-title":"Lancet"},{"key":"10.1016\/j.biosystems.2026.105812_bib66","doi-asserted-by":"crossref","first-page":"971","DOI":"10.1126\/science.150.3699.971","article-title":"Pain mechanisms: a new theory","volume":"150","author":"Melzack","year":"1965","journal-title":"Science"},{"key":"10.1016\/j.biosystems.2026.105812_bib67","doi-asserted-by":"crossref","first-page":"2210","DOI":"10.1093\/gbe\/evu177","article-title":"The evolution of the four subunits of voltage-gated calcium channels: ancient roots, increasing complexity, and multiple losses","volume":"6","author":"Moran","year":"2014","journal-title":"Genome Biol. Evol."},{"key":"10.1016\/j.biosystems.2026.105812_bib68","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/S0006-3495(81)84782-0","article-title":"Voltage oscillations in the barnacle giant muscle fiber","volume":"35","author":"Morris","year":"1981","journal-title":"Biophys. J."},{"key":"10.1016\/j.biosystems.2026.105812_bib69","doi-asserted-by":"crossref","first-page":"R316","DOI":"10.1016\/j.cub.2006.03.089","article-title":"Nanosecond-scale kinetics of nematocyst discharge","volume":"16","author":"N\u00fcchter","year":"2006","journal-title":"Curr. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib70","doi-asserted-by":"crossref","first-page":"493","DOI":"10.1242\/jeb.61.2.493","article-title":"Control of moulting and metamorphosis in the tobacco hornworm, Manduca sexta (L.): cessation of juvenile hormone secretion as a trigger for pupation","volume":"61","author":"Nijhout","year":"1974","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib71","article-title":"Biosynthesis of ecdysteroids: Drosophila and other insect species","author":"Niwa","year":"2010","journal-title":"Ann. N.Y. Acad. Sci. Enzyme Res."},{"key":"10.1016\/j.biosystems.2026.105812_bib72","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1105\/tpc.10.7.1121","article-title":"Arabidopsis mutants define a central role for the xanthophyll cycle in the regulation of photosynthetic energy conversion","volume":"10","author":"Niyogi","year":"1998","journal-title":"Plant Cell"},{"key":"10.1016\/j.biosystems.2026.105812_bib73","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1016\/S0163-7827(01)00020-0","article-title":"Cholesterol interactions with phospholipids in membranes","volume":"41","author":"Ohvo-Rekil\u00e4","year":"2002","journal-title":"Prog. Lipid Res."},{"key":"10.1016\/j.biosystems.2026.105812_bib74","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1146\/annurev.mi.41.100187.001505","article-title":"Prokaryotic hopanoids and other polyterpenoid sterol surrogates","volume":"41","author":"Ourisson","year":"1987","journal-title":"Annu. Rev. Microbiol."},{"key":"10.1016\/j.biosystems.2026.105812_bib75","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/j.jad.2022.03.035","article-title":"Enhanced default mode network functional connectivity links with ECT response","volume":"306","author":"Pang","year":"2022","journal-title":"J. Affect. Disord."},{"key":"10.1016\/j.biosystems.2026.105812_bib76","doi-asserted-by":"crossref","first-page":"294","DOI":"10.1242\/jeb.19.3.294","article-title":"The excitation of nematocysts","volume":"19","author":"Pantin","year":"1942","journal-title":"J. Exp. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib77","doi-asserted-by":"crossref","first-page":"17290","DOI":"10.1523\/JNEUROSCI.2619-13.2013","article-title":"The major brain cholesterol metabolite 24(S)-hydroxycholesterol is a potent allosteric modulator of N-methyl-D-aspartate receptors","volume":"33","author":"Paul","year":"2013","journal-title":"J. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib78","doi-asserted-by":"crossref","first-page":"5464","DOI":"10.1073\/pnas.1117206109","article-title":"Electroconvulsive therapy reduces frontal cortical connectivity in severe depressive disorder","volume":"109","author":"Perrin","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105812_bib79","series-title":"Drive: Neurobiological and Molecular Mechanisms of Sexual Motivation","author":"Pfaff","year":"1999"},{"key":"10.1016\/j.biosystems.2026.105812_bib80","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1016\/j.plipres.2011.06.002","article-title":"Cholesterol metabolism in neurons and astrocytes","volume":"50","author":"Pfrieger","year":"2011","journal-title":"Prog. Lipid Res."},{"key":"10.1016\/j.biosystems.2026.105812_bib81","first-page":"455","article-title":"Lipids of gelatinous Antarctic zooplankton: cnidaria and Ctenophora","volume":"35","author":"Phleger","year":"2000","journal-title":"Lipids"},{"key":"10.1016\/j.biosystems.2026.105812_bib82","doi-asserted-by":"crossref","first-page":"346","DOI":"10.1038\/ncb954","article-title":"Building a cell cycle oscillator: hysteresis and bistability in the activation of Cdc2","volume":"5","author":"Pomerening","year":"2003","journal-title":"Nat. Cell Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib83","doi-asserted-by":"crossref","first-page":"1219","DOI":"10.1073\/pnas.0610286104","article-title":"All four CatSper ion channel proteins are required for male fertility and sperm cell hyperactivation","volume":"104","author":"Qi","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105812_bib84","doi-asserted-by":"crossref","first-page":"12527","DOI":"10.1073\/pnas.221454998","article-title":"A voltage-gated ion channel expressed specifically in spermatozoa","volume":"98","author":"Quill","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105812_bib85","article-title":"A multiplicative behavioral model of DNA replication initiation in cells","volume":"20","author":"Rahman","year":"2025","journal-title":"Open Life Sci."},{"key":"10.1016\/j.biosystems.2026.105812_bib86","doi-asserted-by":"crossref","DOI":"10.3389\/fpsyt.2025.1669530","article-title":"The ARCH model: a neuroevolutionary framework for behavioral execution","volume":"16","author":"Rahman","year":"2025","journal-title":"Front. Psychiatr."},{"key":"10.1016\/j.biosystems.2026.105812_bib87","doi-asserted-by":"crossref","DOI":"10.1016\/j.plantsci.2026.113010","article-title":"Threshold decision-making in the Venus flytrap","volume":"365","author":"Rahman","year":"2026","journal-title":"Plant Sci."},{"key":"10.1016\/j.biosystems.2026.105812_bib88","doi-asserted-by":"crossref","DOI":"10.1037\/tam0000257","article-title":"Archetype killers","author":"Rahman","year":"2025","journal-title":"J. Threat Assess. Manag"},{"key":"10.1016\/j.biosystems.2026.105812_bib89","article-title":"Sex change in clownfish as an ARCH-governed biological decision. Horm","volume":"180","author":"Rahman","year":"2026","journal-title":"Beyond Behav."},{"key":"10.1016\/j.biosystems.2026.105812_bib90","doi-asserted-by":"crossref","DOI":"10.1111\/jne.13028","article-title":"Neurosteroid replacement therapy for catamenial epilepsy, postpartum depression and neuroendocrine disorders in women","volume":"34","author":"Reddy","year":"2022","journal-title":"J. Neuroendocrinol."},{"key":"10.1016\/j.biosystems.2026.105812_bib91","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1016\/j.nurt.2009.01.006","article-title":"Neurosteroid replacement therapy for catamenial epilepsy","volume":"6","author":"Reddy","year":"2009","journal-title":"Neurotherapeutics"},{"key":"10.1016\/j.biosystems.2026.105812_bib92","first-page":"899","article-title":"Hormones and Drosophila development","volume":"vol. 2","author":"Riddiford","year":"1993"},{"key":"10.1016\/j.biosystems.2026.105812_bib93","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1038\/nature06262","article-title":"Identification of a mechanism of photoprotective energy dissipation in higher plants","volume":"450","author":"Ruban","year":"2007","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib94","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1177\/1073858410373835","article-title":"Cholesterol: a novel regulatory role in myelin formation","volume":"17","author":"Saher","year":"2011","journal-title":"Neuroscientist"},{"key":"10.1016\/j.biosystems.2026.105812_bib95","series-title":"Behave: the Biology of Humans at our Best and Worst","author":"Sapolsky","year":"2017"},{"key":"10.1016\/j.biosystems.2026.105812_bib96","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1136\/jnnp.37.2.152","article-title":"Impulse conduction in multiple sclerosis","volume":"37","author":"Schauf","year":"1974","journal-title":"J. Neurol. Neurosurg. Psychiatry"},{"key":"10.1016\/j.biosystems.2026.105812_bib97","doi-asserted-by":"crossref","first-page":"481","DOI":"10.3389\/fphys.2016.00481","article-title":"Physiology and evolution of voltage-gated calcium channels in early diverging animal phyla: cnidaria, Placozoa, Porifera and Ctenophora","volume":"7","author":"Senatore","year":"2016","journal-title":"Front. Physiol."},{"key":"10.1016\/j.biosystems.2026.105812_bib98","doi-asserted-by":"crossref","first-page":"6667","DOI":"10.1523\/JNEUROSCI.1399-04.2004","article-title":"Slow actions of neuroactive steroids at GABAA receptors","volume":"24","author":"Shu","year":"2004","journal-title":"J. Neurosci."},{"key":"10.1016\/j.biosystems.2026.105812_bib99","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1038\/35036052","article-title":"Lipid rafts and signal transduction","volume":"1","author":"Simons","year":"2000","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib100","doi-asserted-by":"crossref","first-page":"3895","DOI":"10.1073\/pnas.071527998","article-title":"Gregarious behavior in desert locusts is evoked by touching their back legs","volume":"98","author":"Simpson","year":"2001","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105812_bib101","doi-asserted-by":"crossref","first-page":"899","DOI":"10.1111\/jnc.15228","article-title":"24S-hydroxycholesterol: cellular effects and variations in brain diseases","volume":"157","author":"Sodero","year":"2021","journal-title":"J. Neurochem."},{"key":"10.1016\/j.biosystems.2026.105812_bib102","doi-asserted-by":"crossref","first-page":"267","DOI":"10.1038\/270267a0","article-title":"Modulation of electrical activity in Aplysia neurones by cholesterol","volume":"270","author":"Stephens","year":"1977","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib103","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1098\/rstb.2006.1837","article-title":"Steroids, triterpenoids and molecular oxygen","volume":"361","author":"Summons","year":"2006","journal-title":"Phil. Trans. Roy. Soc. Lond. B"},{"key":"10.1016\/j.biosystems.2026.105812_bib104","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1086\/708947","article-title":"Activated L-type calcium channels inhibit chemosensitized nematocyst discharge from sea anemone tentacles","volume":"238","author":"Thorington","year":"2020","journal-title":"Biol. Bull."},{"key":"10.1016\/j.biosystems.2026.105812_bib105","series-title":"The study of instinct","author":"Tinbergen","year":"1951"},{"key":"10.1016\/j.biosystems.2026.105812_bib106","series-title":"The Structural Biology of Palms","author":"Tomlinson","year":"1990"},{"key":"10.1016\/j.biosystems.2026.105812_bib107","first-page":"R1104","article-title":"Programmed cell death, paraptosis and the origins of metamorphosis","volume":"29","author":"Truman","year":"2019","journal-title":"Curr. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib108","first-page":"121","article-title":"Neurosteroid biosynthesis in vertebrate brains","volume":"124","author":"Tsutsui","year":"1999","journal-title":"Comp. Biochem. Physiol., C"},{"key":"10.1016\/j.biosystems.2026.105812_bib109","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1006\/jtbi.2001.2293","article-title":"Regulation of the eukaryotic cell cycle: molecular antagonism, hysteresis, and irreversible transitions","volume":"210","author":"Tyson","year":"2001","journal-title":"J. Theor. Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib110","doi-asserted-by":"crossref","first-page":"139","DOI":"10.4161\/psb.2.3.4217","article-title":"Closing of Venus flytrap by electrical stimulation of motor cells","volume":"2","author":"Volkov","year":"2007","journal-title":"Plant Signal. Behav."},{"key":"10.1016\/j.biosystems.2026.105812_bib111","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1152\/japplphysiol.00685.2018","article-title":"Stimuli and sensors that initiate skeletal muscle hypertrophy following resistance exercise","volume":"126","author":"Wackerhage","year":"2019","journal-title":"J. Appl. Physiol."},{"key":"10.1016\/j.biosystems.2026.105812_bib112","doi-asserted-by":"crossref","first-page":"1589","DOI":"10.1126\/science.2564698","article-title":"Cnidocyte mechanoreceptors are tuned to the movements of swimming prey by chemoreceptors","volume":"243","author":"Watson","year":"1989","journal-title":"Science"},{"key":"10.1016\/j.biosystems.2026.105812_bib113","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.57578","article-title":"A molecular filter for the cnidarian stinging response","volume":"9","author":"Weir","year":"2020","journal-title":"eLife"},{"key":"10.1016\/j.biosystems.2026.105812_bib114","doi-asserted-by":"crossref","first-page":"12905","DOI":"10.1073\/pnas.1208255109","article-title":"Discovery, taxonomic distribution, and phenotypic characterization of a gene required for 3-methylhopanoid production","volume":"109","author":"Welander","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/j.biosystems.2026.105812_bib115","doi-asserted-by":"crossref","first-page":"1371","DOI":"10.1016\/j.biocel.2010.05.012","article-title":"Human exercise-induced skeletal muscle hypertrophy is an intrinsic process","volume":"42","author":"West","year":"2010","journal-title":"Int. J. Biochem. Cell Biol."},{"key":"10.1016\/j.biosystems.2026.105812_bib116","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1038\/133725b0","article-title":"Factors controlling moulting and \u2018Metamorphosis\u2019 in an insect","volume":"133","author":"Wigglesworth","year":"1934","journal-title":"Nature"},{"key":"10.1016\/j.biosystems.2026.105812_bib117","doi-asserted-by":"crossref","first-page":"1081","DOI":"10.1038\/ncomms2088","article-title":"Thyroid hormone determines the start of the sensitive period of imprinting and primes later learning","volume":"3","author":"Yamaguchi","year":"2012","journal-title":"Nat. Commun."},{"key":"10.1016\/j.biosystems.2026.105812_bib118","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1146\/annurev-ento-120811-153608","article-title":"Ecdysone control of developmental transitions: lessons from Drosophila research","volume":"58","author":"Yamanaka","year":"2013","journal-title":"Annu. Rev. Entomol."},{"key":"10.1016\/j.biosystems.2026.105812_bib119","doi-asserted-by":"crossref","DOI":"10.1016\/j.neubiorev.2025.106119","article-title":"New directions in neurosteroid therapeutics in neuropsychiatry","author":"Zorumski","year":"2025","journal-title":"Neurosci. Biobehav. Rev."},{"key":"10.1016\/j.biosystems.2026.105812_bib120","doi-asserted-by":"crossref","DOI":"10.1016\/j.ynstr.2019.100196","article-title":"Neurosteroids as novel antidepressants and anxiolytics: GABAA receptors and beyond","volume":"11","author":"Zorumski","year":"2019","journal-title":"Neurobiol. Stress"}],"container-title":["BioSystems"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S030326472600122X?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S030326472600122X?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T11:37:53Z","timestamp":1781609873000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S030326472600122X"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7]]},"references-count":120,"alternative-id":["S030326472600122X"],"URL":"https:\/\/doi.org\/10.1016\/j.biosystems.2026.105812","relation":{},"ISSN":["0303-2647"],"issn-type":[{"value":"0303-2647","type":"print"}],"subject":[],"published":{"date-parts":[[2026,7]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"ARCH \u00d7 \u03a6: Sterols and the evolution of gated biological execution","name":"articletitle","label":"Article Title"},{"value":"BioSystems","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.biosystems.2026.105812","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 The Authors. Published by Elsevier B.V.","name":"copyright","label":"Copyright"}],"article-number":"105812"}}