Odor-mediated push-pull pollination in cycads.

The reproductive organs of some plants self-heat, release scent, and attract pollinators. The relations among these processes are not well understood, especially in the more ancient, nonflowering gymnosperm lineages. We describe the influence of plant volatiles in an obligate pollination mutualism between an Australian Macrozamia cycad (a gymnosperm with male and female individuals) and its specialist thrips pollinator, Cycadothrips chadwicki. Pollen-laden thrips leave male cycad cones en masse during the daily thermogenic phase, when cone temperatures and volatile emissions increase dramatically and thrips are repelled. As thermogenesis declines, total volatile emissions diminish and cones attract thrips, resulting in pollination of female cones. Behavioral and electrophysiological tests on thrips reveal that variations in b-myrcene and ocimene emissions by male and female cones are sufficient to explain the observed sequential thrips' repellence (push) and attraction (pull). These dynamic interactions represent complex adaptations that enhance the likelihood of pollination and may reflect an intermediate state in the evolution of biotic pollination.