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ENVIRONMENTAL STRESS AND ADAPTATION

Submergence-Induced Morphological, Anatomical, and Biochemical Responses in a Terrestrial Species Affect Gas Diffusion Resistance and Photosynthetic Performance

Department of Experimental Plant Ecology (L.M., E.J.W.V.) and Department of Experimental Botany (M.W.-A.), Radboud University Nijmegen, 6525 ED Nijmegen, The Netherlands; Department of Plant Ecophysiology, Utrecht University, 3584 CA Utrecht, The Netherlands (T.L.P.); and Laboratory of Plant Physiology, Department of Plant Biology, University of Groningen, 9750 AA Haren, The Netherlands (J.H.V.)

Gas exchange between the plant and the environment is severely hampered when plants are submerged, leading to oxygen and energy deficits. A straightforward way to reduce these shortages of oxygen and carbohydrates would be continued photosynthesis under water, but this possibility has received only little attention. Here, we combine several techniques to investigate the consequences of anatomical and biochemical responses of the terrestrial species Rumex palustris to submergence for different aspects of photosynthesis under water. The orientation of the chloroplasts in submergence-acclimated leaves was toward the epidermis instead of the intercellular spaces, indicating that underwater CO₂ diffuses through the cuticle and epidermis. Interestingly, both the cuticle thickness and the epidermal cell wall thickness were significantly reduced upon submergence, suggesting a considerable decrease in diffusion resistance. This decrease in diffusion resistance greatly facilitated underwater photosynthesis, as indicated by higher underwater photosynthesis rates in submergence-acclimated leaves at all CO₂ concentrations investigated. The increased availability of internal CO₂ in these "aquatic" leaves reduced photorespiration, and furthermore reduced excitation pressure of the electron transport system and, thus, the risk of photodamage. Acclimation to submergence also altered photosynthesis biochemistry as reduced Rubisco contents were observed in aquatic leaves, indicating a lower carboxylation capacity. Electron transport capacity was also reduced in these leaves but not as strongly as the reduction in Rubisco, indicating a substantial increase of the ratio between electron transport and carboxylation capacity upon submergence. This novel finding suggests that this ratio may be less conservative than previously thought.

^* Corresponding author; e-mail l.mommer{at}science.ru.nl; fax 31–24–3652409.

Received April 27, 2005; returned for revision May 20, 2005; accepted July 11, 2005.

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