Pathogens, parasites, and their hosts are involved in evolutionary arms races. The evolution of resistance on the part of the host is often followed by counter-adaptation on the part of the pathogen. To date, the hypothesis that natural enemies can counter-adapt to the resistance encoded by secondary symbionts has not been tested, although it is likely that they do. First experimental avenues would be to examine natural enemy populations for genetic variation in the ability to remain unaffected by secondary symbiont mediated resistance, and to conduct experimental evolution studies where the capacity of natural enemies to evolve when faced with symbiont mediated resistance is examined.
The above experiments would determine whether counter-adaptation in the natural enemy is possible. If it is, it is not clear whether or not the trajectory of coevolution will differ in interactions between natural enemies and nuclear- and symbiont-encoded resistance factors. The difference in population biology of secondary symbiont and nuclear resistance alleles outlined above suggest they may do; symbiont-encoded resistance is more likely to be lost in the short term, leaving less scope for counter-adaptation.
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