The gene can be de­scribed as the foun­da­tional con­cept of mod­ern bi­ol­ogy. As such, it has spilled over into daily dis­course, yet it is ac­knowl­edged among bi­ol­o­gists to be ill-de­fined. Here, fol­low­ing a short his­tory of the gene, I analyse crit­i­cally its role in in­her­i­tance, evo­lu­tion, de­vel­op­ment, and mor­pho­gen­e­sis. Wil­helm Jo­hannsen's geno­type-con­cep­tion, for­mu­lated in 1910, has been adopted as the foun­da­tion stone of ge­net­ics, giv­ing the gene a higher de­gree of promi­nence than is jus­ti­fied by the ev­i­dence. An analy­sis of the re­sults of the Long-Term Evo­lu­tion Ex­per­i­ment (LTEE) with E. coli bac­te­ria, grown over 60,000 gen­er­a­tions, does not sup­port spon­ta­neous gene mu­ta­tion as the source of vari­ance for nat­ural se­lec­tion. From this it fol­lows that the gene is not Mendel's unit of in­her­i­tance: that must be Jo­hannsen's trans­mis­sion-con­cep­tion at the ga­mete phe­no­type level, a form of in­her­i­tance that Jo­hannsen did not con­sider. Al­ter­na­tively, I con­tend that bi­ol­ogy viewed on the bases of ther­mo­dy­nam­ics, com­plex sys­tem dy­nam­ics and self-or­gan­i­sa­tion, pro­vides a new frame­work for the foun­da­tions of bi­ol­ogy. In this frame­work, the gene plays a pas­sive role as a vi­tal in­for­ma­tion store: it is the phe­no­type that plays the ac­tive role in in­her­i­tance, evo­lu­tion, de­vel­op­ment, and mor­pho­gen­e­sis.

The gene: An appraisal

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