Vegetation-derived abscisic acid and four terpenes enforce dormancy in seeds of the post-fire annual,Nicotiana attenuata

The native tobacco,Nicotiana attenuata, synchronizes its germination with the immediate post-fire environment with a combination of germination stimulants found in wood smoke and inhibitors from the unburned litter of the dominant vegetation. The inhibitors override the stimulants and prevent seeds from germinating maladaptively in unburned habitats adjacent to burns. To understand the physiological basis of this environmental control of germination, we tested several previously isolated signals, phytohormones and their respective biosynthesis inhibitors. The germination inhibitors methyl jasmonate (MeJA, a constituent of sagebrush litter), bornane-2,5-dione (BD, a constituent of juniper litter extract, JLE) and JLE did not alter abscisic acid (ABA) content of imbibed seeds. Treatment with the ABA biosynthesis inhibitor, fluridone, inhibited the dormancy-inducing effects of BD, JLE and MeJA, but surprisingly did not affect endogenous ABA levels in treated seeds. However, ABA leached from litter of the species, which dominate the plant community before fires, plays an important role in germination control. We conclude thatN. attenuataseeds, which can lie dormant in the soil for 150 years between fires, time their germination with the post-fire environment by responding to smoke, ABA and four terpenes (BD, 1,8-cineole, β-thujaplicin and camphor) leaching from the litter of the dominant vegetation.