Photosynthesis, Respiration, and Thermal Energy Dissipation in Leaves of Two Phenotypes of Plantago media L. under Environmental Conditions

The ability to maintain the balance between the absorbed light energy and energy used in photosynthesis is a key factor of plant adaptation to variable environmental conditions. In this work, diurnal variations in photosynthesis, respiration, thermal energy dissipation, and the activity of the antioxidant system were studied in hoary plantain (Plantago media L.) growing on an open slope (sun plants) and under natural shading in the herbage (shade plants). The highest leaves net photosynthetic rate (Pn) was observed early in the morning and amounted to 2.6 and 9.2 mu mol CO2/m(2) s in shade and sun plants, respectively. In the daytime, the Pn values of sun plants decreased significantly (threefold) along with the decrease in stomatal conductance; changes of both parameters developed concurrently with the increase in insolation and air temperature. The Pn changes in leaves of shade plants were less pronounced and weakly dependent on stomatal conductance. The leaves of shade plants contained comparatively high amounts of soluble carbohydrates, whereas the sun plant leaves accumulated larger amounts of starch. In the daytime, nonphotochemical quenching (NPQ) of chlorophyll a fluorescence in photosystem II of sun plant leaves could be as large as 2.6 rel. units, which was four- to fivefold higher than NPQ in shade plants. In leaves of sun plants in the morning and evening hours, the ratio of cytochrome pathway (CP) and the alternative (AP) respiratory pathways was approximately 1.0, whereas this ratio decreased to 0.4 during the day, synchronously with an increase in NPQ. The CP/AP ratio in shade plant leaves remained constant throughout the diurnal cycle and equaled 1.4, indicating a comparatively high energy efficiency of respiration in shaded plants growing under the grass canopy. The leaves of sun plants featured an increased content of superoxide anion radical and hydrogen peroxide as well as the elevated activity of antioxidant enzymes (superoxide dismutase, guaiacol peroxidase) that control the accumulation of reactive oxygen species. The results emphasize the importance of coordinated changes in energy-dissipating processes and the activity of the antioxidant system for maintaining the energy and redox balance in phototrophic tissues during long-term adaptation of plants to environmental conditions, excessive insolation in particular.