Evolution Across Scales

Living systems are organized hierarchically, from molecules to ecosystems. How do the complex interactions between these scales influence evolution?

Here's an example of what I mean. Genetic elements (e.g. genes and non-coding regulatory sites) live on chromosomes. Chromosomes can be effectively shuffled through large mutations, altering the arrangement of genetic elements. Whether these large rearrangements fix in a population or not depends on which genetic elements are along for the ride. At the same time, seismic shifts in chromosomes alter the evolutionary landscape of the passengers. Factor in the three-dimensional shape of the genome, and things start to look very complicated - and very interesting.