Drastic environmental change and its effects on a planetary biosphere

Environmental conditions can change drastically and rapidly during the natural history of a planetary body. These changes affect the biosphere and can spur evolution via the mechanism of directional selection leading to the innovation of new processes and forms of life, or alternatively leading to the extinction of certain life forms. Based on the natural history of Earth, the effect on a planet's biosphere depends on three factors: (1) the nature and time scale of change, (2) the composition of the biosphere prior to change, and (3) the nature of the environment following the change. Though Earth has undergone various periods of drastic environmental change, life has shown an enormous resiliency and became more diverse and complex as a consequence of these events. Mars and Venus have undergone even larger environmental changes, both from habitable conditions under which the origin of life (or transfer of life from Earth) seem plausible, to a dry and cold planet punctuated by wetter conditions, and a hyperthermic greenhouse, respectively. Given its planetary history, life on Mars could have retreated to a psychrophilic lifestyle in the deep subsurface or to environmental near-surface niches, such as hydrothermal regions and caves. Further, strong directional selection could have pushed putative martian life to evolve alternating cycles between active and dormant forms, as well as the innovation of new traits adapted to challenging near-surface conditions. Life in the subsurface or on the surface of Venus seems impossible today, but microorganisms may have adapted to thrive in the lower cloud layer, possibly using a biochemical strategy analogous to Photosystem I and chemoautotrophic sulfur metabolism, and employing cycloocta sulfur for UV protection.