The hierarchical structure of organisms: A scale and documentation of a trend in the maximum
The degree of hierarchical structure of organisms-the number of levels of nesting of lower-level entities within higher-level individuals-has apparently increased a number of times in the history of life, notably in the origin of the eukaryotic cell from an association of prokaryotic cells, of multicellular organisms from clones of eukaryotic cells, and of intergrated colonies from aggregates of multicellular individuals. Arranged in order of first occurrence, these three transitions suggest a trend, in particular a trend in the maximum, or an increase in the degree of hierarchical structure present in the hierarchically deepest organism on Earth. However, no rigorous documentation of such a trend-based on operational and consistent criteria for hierarchical levels-has been attempted. Also, the trajectory of increase has not been examined in any detail. One limitation is that no hierarchy scale has been developed with sufficient resolution to document more than these three major increases. Here, a higher-resolution scale is proposed in which hierarchical structure is decomposed into levels and sublevels, with levels reflecting number of layers of nestedness, and sublevels reflecting degree of individuation at the highest level. The scale is then used, together with the body-fossil record, to plot the trajectory of the maximum. Two alternative interpretations of the record are considered, and both reveal a long-term trend extending from the Archean through the early Phanerozoic. In one, the pattern of increase was incremental, with almost all sublevels arising precisely in order. The data also raise the possibility that waiting times for transitions between sublevels may have decreased with increasing hierarchical level (and with time). These last two findings-incremental increase in level and decreasing waiting times-are tentative, pending a study of possible biases in the fossil record.
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