When researchers at Emory University in Atlanta trained mice to fear the smell of almonds (by pairing it with electric shocks), they found, to their consternation, that both the children and grandchildren of these mice were spontaneously afraid of the same smell. That is not supposed to happen. Generations of school children have been taught that the inheritance of acquired characteristics is impossible. A mouse should not be born with something its parents have learned during their lifetimes, any more than a mouse that loses its tail in an accident should give birth to tailless mice.

Modern evolutionary biology dates back to a synthesis that emerged around the $1940 \mathrm{~s}-60 \mathrm{~s}$, which married Charles Darwin's mechanism of natural selection with Gregor Mendel's discoveries of how genes are inherited. The traditional, and still dominant, view is that adaptations - from the human brain to the peacock's tail - are fully and satisfactorily explained by natural selection (and subsequent inheritance). Yet new evidence from genomics, epigenetics and developmental biology indicates that evolution is more complex than we once assumed.

In his book On Human Nature (1978), the evolutionary biologist Edward O Wilson claimed that human culture is held on a genetic leash. The metaphor needs revision. Imagine a dog-walker (the genes) struggling to retain control of a brawny mastiff (human culture). The pair's trajectory (the pathway of evolution) reflects the outcome of the struggle. Now imagine the same dog-walker struggling with multiple dogs, on leashes of varied lengths, with each dog tugging in different directions. All these tugs represent the influence of developmental factors, including epigenetics, antibodies and hormones passed on by parents, as well as the ecological legacies and culture they bequeath.

The received wisdom is that parental experiences can't affect the characters of their offspring. Except they do. The way that genes are expressed to produce an organism's phenotype - the actual characteristics it ends up with - is affected by chemicals that attach to them. Everything from diet to air pollution to parental behaviour can influence the addition or removal of these chemical marks, which switches genes on or off. Usually these so-called 'epigenetic' attachments are removed during the production of sperm and eggs cells, but it turns out that some escape the resetting process and are passed on to the next generation, along with the genes. This is known as 'epigenetic inheritance', and more and more studies are confirming that it really happens. Let's return to the almond-fearing mice. The inheritance of an epigenetic mark transmitted in the sperm is what led the mice's offspring to acquire an inherited fear.

Epigenetics is only part of the story. Through culture and society, humans and other animals inherit knowledge and skills acquired by their parents. All this complexity points to an evolutionary process in which genomes (over hundreds to thousands of generations), epigenetic modifications and inherited cultural factors (over several, perhaps tens or hundreds of generations), and parental effects (over single-generation timespans) collectively inform how organisms adapt. These extragenetic kinds of inheritance give organisms the flexibility to make rapid adjustments to environmental challenges, dragging genetic change in their wake - much like a rowdy pack of dogs.