Rupert Riedl's concept of burden forms a causal hypothesis on organismic integration and evolutionary constraints. Defined as the hierarchically nested interdependence of characters within the organism, burden was seen as (1) defining and conserving body plans and (2) constraining and directing evolutionary trajectories. A review of the components of the burden concept reveals important consistencies with the modern tenets of evo-devo. This concept differs from the current consensus of evolutionary theory in that it (1) grants evolution less options for changing tightly integrated, "locked-in" characters and (2) in deducing from this an ever decreasing freedom for evolution, with cyclism and typostrophism as resulting macroevolutionary phenomena. Despite these differences, I show that the burden concept was consistent with most major tenets of the Modern Synthesis, and Riedl attempted to explain patterns of large-scale evolutionary trends exclusively by microevolutionary (gradualistic) processes. The burden concept is fruitful and unique in its focus on hierarchically nested constraints and resembles the hierarchical architecture of gene regulatory networks. However, such networks are more high-dimensional and most of their components appear to be easier to evolve than Riedl's burden. Yet in combination with evolvability, a modified concept of burden might contribute substantially to the understanding of organismic integration and the long-term evolution of body plans.
(c) 2009 Wiley-Liss, Inc.