If I have a sample of rats in which I want to measure protein levels in several different brain regions, the rats are of two genotypes and both sexes, and I want to take into account that the brain regions come from specific animals, which is the appropriate syntax for modeling this with lme4?
marker ~ genotype * sex * brain.region + (1|animal/brain.region)
marker ~ genotype * sex * brain.region + (animal|brain.region)