Adaptations for an amphibious life: changes in leaf morphology, growth rate, carbon and nitrogen investment, and reproduction during adjustment to emersion by the freshwater macrophyte Littorella uniflora

Littorella uniflora (L.) Ascherson is a small, perennial, amphibious rhizophyte of rosette life-form which is common along the margins of lakes, tarns and reservoirs where water-level fluctuations are often rapid and unpredictable. The majority of plants are continuously submersed and reproduce vegetatively, but a small proportion become completely emersed for variable lengths of time, when flowering and seed set occur. To find out how L. uniflora adjusts to sudden emersion we studied the plants at a reservoir where water level falls each spring and remains low throughout the summer; L. uniflora adjusted very quickly showing a degree of phenotypic plasticity not expected in a 'stress tolerator', including the production of a new set of terrestrial leaves with reduced lacunal volume and increased stomatal density, a rapid increase in leaf growth rate, and flowering within 3–4 wk. Comparison of terrestrial L. uniflora with aquatic plants growing permanently submersed in lake and tarn habitats showed that three to fourfold more carbon (C) and nitrogen (N) was incorporated into above-ground biomass by emersed plants. However, ramet production in the aquatic environment appeared to be more costly, in terms of C and N invested, than terrestrial flower and seed production. The combination of continuous, submersed vegetative spread with the capacity for a high degree of phenotypic plasticity allowing some flower and seed production to occur during brief periods of emersion seems to account for the success of this plant in the amphibious niche.