The riddle of mitochondrial alkaline/neutral invertases: A novel Arabidopsis isoform mainly present in reproductive tissues and involved in root ROS production.

Alkaline/neutral invertases (A/N-Inv), glucosidases that irreversibly hydrolyze sucrose into glucose and fructose, play significant roles in plant growth, development, and stress adaptation. They occur as multiple isoforms located in the cytosol or organelles. In Arabidopsis thaliana, two mitochondrial A/N-Inv genes (A/N-InvA and A/N-InvC) have already been investigated. In this study, we functionally characterized A/N-InvH, a third Arabidopsis gene coding for a mitochondrial-targeted protein. The phenotypic analysis of knockout mutant plants (invh) showed a severely reduced shoot growth, while root development was not affected. The emergence of the first floral bud and the opening of the first flower were the most affected stages, presenting a significant delay. A/N-InvH transcription is markedly active in reproductive tissues. It is also expressed in the elongation and apical meristem root zones. Our results show that A/N-InvH expression is not evident in photosynthetic tissues, despite being of relevance in developmental processes and mitochondrial functional status. NaCl and mannitol treatments increased A/N-InvH expression twofold in the columella root cap. Moreover, the absence of A/N-InvH prevented ROS formation, not only in invh roots of salt- and ABA-treated seedlings but also in invh control roots. We hypothesize that this isoform may take part in the ROS/sugar (sucrose or its hydrolysis products) signaling pathway network, involved in reproductive tissue development, cell elongation, and abiotic stress responses.

A mitochondrial alkaline/neutral invertase isoform (A/N-InvC) functions in developmental energy-demanding processes in Arabidopsis.

Recent findings demonstrate that alkaline/neutral invertases (A/N-Invs), enzymes that catalyze the breakdown of sucrose into glucose and fructose, are essential proteins in plant life. The fact that different isoforms are present in multiple locations makes them candidates for the coordination of metabolic processes. In the present study, we functionally characterized the encoding gene of a novel A/N-Inv (named A/N-InvC) from Arabidopsis, which localizes in mitochondria. A/N-InvC is expressed in roots, in aerial parts (shoots and leaves) and flowers. A detailed phenotypic analysis of knockout mutant plants (invc) reveals an impaired growth phenotype. Shoot growth was severely reduced, but root development was not affected as reported for A/N-InvA mutant (inva) plants. Remarkably, germination and flowering, two energy demanding processes, were the most affected stages. The effect of exogenous growth regulators led us to suggest that A/N-InvC may be modulating hormone balance in relation to the radicle emergence. We also show that oxygen consumption is reduced in inva and invc in comparison with wild-type plants, indicating that both organelle isoenzymes may play a fundamental role in mitochondrion functionality. Taken together, our results emphasize the involvement of mitochondrial A/N-Invs in developmental processes and uncover the possibility of playing different roles for the two isoforms located in the organelle.

Genetic variability for leaf growth rate and duration under water deficit in sunflower: analysis of responses at cell, organ, and plant level.

Plants under water deficit reduce leaf growth, thereby reducing transpiration rate at the expense of reduced photosynthesis. The objective of this work was to analyse the response of leaf growth to water deficit in several sunflower genotypes in order to identify and quantitatively describe sources of genetic variability for this trait that could be used to develop crop varieties adapted to specific scenarios. The genetic variability of the response of leaf growth to water deficit was assessed among 18 sunflower (Helianthus annuus L.) inbred lines representing a broad range of genetic diversity. Plants were subjected to long-term, constant-level, water-deficit treatments, and the response to water deficit quantified by means of growth models at cell-, leaf-, and plant-scale. Significant variation among lines was found for the response of leaf expansion rate and of leaf growth duration, with an equal contribution of these responses to the variability in the reduction of leaf area. Increased leaf growth duration under water deficit is usually suggested to be caused by changes in the activity of cell-wall enzymes, but the present results suggest that the duration of epidermal cell division plays a key role in this response. Intrinsic genotypic responses of rate and duration at a cellular scale were linked to genotypic differences in whole-plant leaf area profile to water deficit. The results suggest that rate and duration responses are the result of different physiological mechanisms, and therefore capable of being combined to increase the variability in leaf area response to water deficit.

Rewatering plants after a long water-deficit treatment reveals that leaf epidermal cells retain their ability to expand after the leaf has apparently reached its final size.

BACKGROUND AND AIMS: Leaves expand during a given period of time until they reach their final size and form, which is called determinate growth. Duration of leaf expansion is stable when expressed in thermal-time and in the absence of stress, and consequently it is often proposed that it is controlled by a robust programme at the plant scale. The usual hypothesis is that growth cessation occurs when cell expansion becomes limited by an irreversible tightening of cell wall, and that leaf size is fixed once cell expansion ceases. The objective of this paper was to test whether leaf expansion could be restored by rewatering plants after a long soil water-deficit period. METHODS: Four experiments were performed on two different species (Arabidopsis thaliana and Helianthus annuus) in which the area of leaves that had apparently reached their final size was measured upon reversal of water stresses of different intensities and durations. KEY RESULTS: Re-growth of leaves that had apparently reached their final size occurred in both species, and its magnitude depended only on the time elapsed from growth cessation to rewatering. Leaf area increased up to 186% in A. thaliana and up to 88% in H. annuus after rewatering, with respect to the leaves of plants that remained under water deficit. Re-growth was accounted for by cell expansion. Increase in leaf area represented actual growth and not only a reversible change due to increased turgor. CONCLUSIONS: After the leaf has ceased to grow, leaf cells retain their ability to expand for several days before leaf size becomes fixed. A response window was identified in both species, during which the extent of leaf area recovery decreased with time after the 'initial' leaf growth cessation. These results suggest that re-growth after rewatering of leaves having apparently attained their final size could be a generalized phenomenon, at least in dicotyledonous plants.