Understanding unique tolerance limits in Hydrocotyle verticillata: From submergence to water deficiency

Hydrocotyle verticillata can tolerate varying degrees of flooding, up to complete submergence, and is at the same time extremely sensitive to drought. Understanding the structural and biochemical principles of these unusual tolerance limits is of particular importance. We analyzed the effect of soil flooding, complete submergence (rooted plants and floating stems), and dehydration on root anatomy, alcohol dehydrogenase (ADH), heat shock proteins 70 (HSP70), hydrogen peroxide, and DNA integrity using light microscopy, biochemical and histological methods. It was shown that anatomical traits of adventitious roots with a triarch stele were similar in the plants growing under different conditions; the single-layered and thin-walled epidermis formed relatively short root hairs; essential air spaces were absent in the cortex parenchyma. Results on ADH clearly showed that anaerobic energetic metabolism in root apices and individual rhizoderma cells of the mature root zone was normal for this species in optimal and suboptimal conditions, while leaves changed metabolism to anaerobic in response to submergence. Alterations in the protein spectrum were accompanied by adequate up-regulation of HSP70 under different levels of flooding and dehydration/rehydration. These results appear to indicate a flooding adaptation strategy for H. verticillata based primarily on metabolic plasticity rather than morphoanatomical adaptations. Most notably, the resistance of this species to long-term submergence has been associated with strong ADH induction in leaves, transient activation of 70 kDa isoform of HSP70 and induction of 66 kDa isoform, as well as a significant delay in the accumulation of hydrogen peroxide and DNA degradation.