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The question of closure in biological systems is central to understanding the origins of the biological variation and complexity upon which various forms of selection act. Much of evolutionary theory, especially in the second half of the twentieth century, is concerned with the consequences of environmental selection acting on bio-diversity, but neglects questions of the origin of that diversity. This has permitted us to act as if an explanation of consequences was the ultimate explanation in biology. However, Darwin understood that evolution was both information driven and information constrained. The link between evolutionary constraints and closure can be profitably explored by starting with Darwin's notion of the primary of "the nature of the organism" over "the nature of the conditions" articulated in the sixth edition of Origin of Species. Contemporary ideas of self-organization, emergence, complexity, and inherent (developmental and phylogenetic) constraints can be seen as an elaboration and refinement of Darwin's views if we adopt the following perspective: (1) information is cheap, not costly, to produce, but may have costly consequences; and (2) information is produced by systems that are informationally closed but remain thermodynamically open.
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