Reproductive isolation accumulates gradually during allopatric divergence through drift and selection on different traits. Prezygotic barriers (mate choice, courtship incompatibility) often evolve first; postzygotic barriers (hybrid inviability, sterility) follow. The Dobzhansky-Muller model explains how independent mutations in different populations create reproductive incompatibilities.
You already know the categories of reproductive isolation — prezygotic barriers that prevent mating or fertilization, and postzygotic barriers that reduce hybrid fitness. And you know that speciation requires these barriers to form between populations. The question this topic addresses is: *how do these barriers actually accumulate during divergence?* The answer reveals that speciation is not a single event but a process, with different types of barriers arising at different stages and through different mechanisms.
Consider two populations of the same species separated by a geographic barrier — the classic allopatric scenario you have studied. In their separate environments, each population experiences different selection pressures and accumulates different mutations through drift. Over time, their courtship signals may diverge: one population's males evolve slightly different songs, colors, or pheromones in response to local conditions. If the populations later come into contact, females from one population may not recognize males from the other as suitable mates. This is a prezygotic barrier, and it tends to evolve relatively early because traits involved in mate recognition are often under strong sexual selection and can diverge rapidly. Temporal isolation (breeding at different times) and habitat isolation (preferring different microhabitats) can also arise early as populations adapt to different local environments.
Postzygotic barriers — hybrid inviability and hybrid sterility — typically take longer to accumulate because they require genetic incompatibilities between the diverging genomes. The Dobzhansky-Muller model explains how this happens without requiring any population to pass through a fitness valley. Imagine the ancestral population has genotype AABB at two interacting loci. Population 1 evolves to AAbb (mutation at the B locus), and Population 2 evolves to aaBB (mutation at the A locus). Each new allele works fine in its home genetic background. But a hybrid with genotype AaBb brings together the a and b alleles for the first time — a combination that was never tested by natural selection in either population. If these alleles interact badly (the protein products are incompatible, or they disrupt a shared developmental pathway), the hybrid is inviable or sterile. The key insight is that incompatibility arises not from deleterious mutations but from the novel combination of independently evolved alleles.
The accumulation of barriers follows a rough temporal sequence: behavioral and ecological prezygotic barriers first, then gametic isolation, then postzygotic inviability, and finally hybrid sterility. This ordering matters because it means that if populations make secondary contact early in divergence, prezygotic barriers may be weak and hybridization can reverse speciation. Reinforcement — natural selection strengthening prezygotic barriers when hybrids are unfit — can accelerate the completion of speciation. But reinforcement only works if postzygotic barriers are already partially in place, creating selection against hybridization. The full picture is one of accumulation and feedback: barriers build on each other, and the process accelerates as more barriers arise, until gene flow between the populations effectively ceases.