This inheritance may take many forms and operate on many scales, with a multiplicity of systems of inheritance complementing the genes. From position and maternal effects on gene expression to epigenetic inheritance [6] to the active construction and intergenerational transmission of enduring niches,[2] development systems theory argues that not only inheritance but evolution as a whole can be understood only by taking into account a far wider range of ‘reproducers’ or ‘inheritance systems’ – genetic, epigenetic, behavioural and symbolic [7] – than neo-Darwinism’s ‘atomic’ genes and gene-like ‘replicators’.[8] DST regards every level of biological structure as susceptible to influence from all the structures by which they are surrounded, be it from above, below, or any other direction – a proposition that throws into question some of (popular and professional) biology’s most central and celebrated claims, not least the ‘central dogma’ of Mendelian genetics, any direct determination of phenotype by genotype, and the very notion that any aspect of biological (or psychological, or any other higher form) activity or experience is capable of direct or exhaustive genetic or evolutionary ‘explanation’.[9]