What are the ‘levels’ in levels of selection?

The levels of selection debate is generally taken to be a debate about how natural selection can occur at the various levels of biological organization. In this article, we argue that questions about levels of selection should be analysed separately from questions about levels of organization. In the deflationary proposal we defend, all that is necessary for multilevel selection is that there are cases in which particles are nested in collectives, and that both the collectives and the particles that compose them each separately undergo natural selection. We argue that adopting this deflationary account helps to disentangle the levels of selection and the levels of organization, and thereby contributes to advancing the levels of selection debate