BEL transcription factors in prominent Solanaceae crops: the missing pieces of the jigsaw in plant development.

MAIN CONCLUSION: Understanding BEL transcription factors roles in potato and tomato varies considerably with little overlap. The review suggests reciprocal use of gained results to proceed with the knowledge in both crops The proper development of organs that plants use for reproduction, like fruits or tubers, is crucial for the survival and competitiveness of the species and thus subject to strict regulations. Interestingly, the controls of potato (Solanum tuberosum) tuber and tomato (S. lycopersicum) fruit development use common mechanisms, including the action of the BEL transcription factors (TFs). Although more than ten BEL genes have been identified in either genome, only a few of them have been characterized. The review summarizes knowledge of BEL TFs' roles in these closely related Solanaceae species, focusing on those that are essential for tuberization in potato, namely StBEL5, StBEL11 and StBEL29, and for fruit development in tomato - SlBEL11, SlBL2 and SIBL4. Comprehension of the roles of individual BEL TFs, however, is not yet sufficient. Different levels of understanding of important characteristics are described, such as BEL transcript accumulation patterns, their mobility, BEL protein interaction with KNOX partners, subcellular localisation, and their target genes during initiation and development of the organs in question. A comparison of the knowledge on BEL TFs and their mechanisms of action in potato and tomato may provide inspiration for faster progress in the study of both models through the exchange of information and ideas. Both crops are extremely important for human nutrition. In addition, their production is likely to be threatened by the upcoming climate change, so there is a particular need for breeding using a deep knowledge of control mechanisms.

Light Drives and Temperature Modulates: Variation of Phenolic Compounds Profile in Relation to Photosynthesis in Spring Barley.

Accumulation and metabolic profile of phenolic compounds (PheCs; serving as UV-screening pigments and antioxidants) as well as carbon fixation rate (An) and plant growth are sensitive to irradiance and temperature. Since these factors are naturally co-acting in the environment, it is worthy to study the combined effects of these environmental factors to assess their possible physiological consequences. We investigated how low and high irradiance in combination with different temperatures modify the metabolic profile of PheCs and expression of genes involved in the antioxidative enzyme and PheCs biosynthesis, in relation to photosynthetic activity and availability of non-structural carbohydrates (NSC) in spring barley seedlings. High irradiance positively affected An, NSC, PheCs content, and antioxidant activity (AOX). High temperature led to decreased An, NSC, and increased dark respiration, whilst low temperature was accompanied by reduction of UV-A shielding but increase of PheCs content and AOX. Besides that, irradiance and temperature caused changes in the metabolic profile of PheCs, particularly alteration in homoorientin/isovitexin derivatives ratio, possibly related to demands on AOX-based protection. Moreover, we also observed changes in the ratio of sinapoyl-/feruloyl- acylated flavonoids, the function of which is not yet known. The data also strongly suggested that the NSC content may support the PheCs production.

Multi-Component Antioxidative System and Robust Carbohydrate Status, the Essence of Plant Arsenic Tolerance.

Arsenic (As) contaminates the food chain and decreases agricultural production through impairing plants, particularly due to oxidative stress. To better understand the As tolerance mechanisms, two contrasting tobacco genotypes: As-sensitive Nicotiana sylvestris and As-tolerant N.tabacum, cv. 'Wisconsin' were analyzed. The most meaningful differences were found in the carbohydrate status, neglected so far in the As context. In the tolerant genotype, contrary to the sensitive one, net photosynthesis rates and saccharide levels were unaffected by As exposure. Importantly, the total antioxidant capacity was far stronger in the As-tolerant genotype, based on higher antioxidants levels (e.g., phenolics, ascorbate, glutathione) and activities and/or appropriate localizations of antioxidative enzymes, manifested as reverse root/shoot activities in the selected genotypes. Accordingly, malondialdehyde levels, a lipid peroxidation marker, increased only in sensitive tobacco, indicating efficient membrane protection in As-tolerant species. We bring new evidence of the orchestrated action of a broad spectrum of both antioxidant enzymes and molecules essential for As stress coping. For the first time, we propose robust carbohydrate metabolism based on undisturbed photosynthesis to be crucial not only for subsidizing C and energy for defense but also for participating in direct reactive oxygen species (ROS) quenching. The collected data and suggestions can serve as a basis for the selection of plant As phytoremediators or for targeted breeding of tolerant crops.

Root cultures of potato mutant lacking MSPI isoform, indispensable for photosynthetic light reactions, exhibit characteristics similar to intact plant roots.

Potato (Solanum tuberosum) mutant (ST) lacking one isoform of manganese-stabilizing protein (MSPI) of photosystem II exhibited besides spontaneous tuberization also growth changes with strongly impaired root system development. Previous studies revealed marked changes in carbohydrate levels and allocation within ST plant body. To verify causal relationship between changed carbohydrate balance and root growth restriction we engaged dark grown sucrose-supplied root organ-cultures of ST plants to exclude/confirm shoot effects. Unexpectedly, in ST root cultures we observed large alterations in growth and architecture as well as saccharide status similar to those found in the intact plant roots. The gene expression analysis, however, proved PsbO1 transcript (coding MSPI protein) neither in ST nor in WT root-organ cultures. Therefore, the results point to indirect effects of PsbO1 allele absence connected possibly with some epigenetic modulations.

Two facets of world arsenic problem solution: crop poisoning restriction and enforcement of phytoremediation.

MAIN CONCLUSION: This review provides insights into As toxicity in plants with focus on photosynthesis and sugar metabolism as important arsenic targets and simultaneously defence tools against accompanying oxidative stress. Heavy metal contamination is a great problem all over the world. Arsenic, a metalloid occurring naturally in the Earth's crust, also massively spreads out in the environment by human activities. Its accumulation in crops poses a severe health risk to humans and animals. Besides the restriction of human-caused contamination, there are two basic ways how to cope with the problem: first, to limit arsenic accumulation in harvestable parts of the crops; second, to make use of some arsenic hyperaccumulating plants for phytoremediation of contaminated soils and waters. Progress in the use of both strategies depends strongly on the level of our knowledge on the physiological and morphological processes resulting from arsenic exposure. Arsenic uptake is mediated preferentially by P and Si transporters and its accumulation substantially impairs plant metabolism at numerous levels including damages through oxidative stress. Rice is a predominantly studied crop where substantial progress has been made in understanding of the mechanisms of arsenic uptake, distribution, and detoxification, though many questions still remain. Full exploitation of plant potential for soil and water phytoremediations also requires deep understanding of the plant response to this toxic metalloid. The aim of this review is to summarize data regarding the effect of arsenic on plant physiology with a focus on mechanisms providing increased arsenic tolerance and/or hyperaccumulation. The emphasis is placed on the topic unjustifiably neglected in the previous reviews - i.e., carbohydrate metabolism, tightly connected to photosynthesis, and beside others involved in plant ability to cope with arsenic-induced oxidative and nitrosative stresses.

Carbohydrates and gibberellins relationship in potato tuberization.

Potato represents the third most important crop worldwide and therefore to understand regulations of tuber onset is crucial from both theoretical and practical points of view. Photosynthesis and related carbohydrate status along with phytohormone balance belong to the essential factors in regulation of plant development including storage organ formation. In our work we used potato (Solanum tuberosum) cv. Lada and its spontaneously tuberizing mutant (ST plants) grown in vitro under low carbohydrate availability (non-inductive conditions). Small plant phenotype and readiness to tuberization of ST plants was, however, not accompanied by lower gibberellins levels, as determined by UHPLC-MS/MS. Therefore, we focused on the other inducing factor, carbohydrate status. Using HPLC, we followed changes in carbohydrate distribution under mixotrophic (2.5% sucrose in medium) and photoautotrophic conditions (no sucrose addition and higher gas and light availability) and observed changes in soluble carbohydrate allocation and starch deposition, favouring basal stem part in mutants. In addition, the determination of tuber-inducing marker gene expressions revealed increased levels of StSP6A in ST leaves. Collectively these data point towards the possibility of two parallel cross-talking pathways (carbohydrate - and gibberellin- dependent ones) with the power of both to outcompete the other one when its signal is for some reason extraordinary strong.

A yeast mitotic activator sensitises the shoot apical meristem to become floral in day-neutral tobacco.

True day-neutral (DN) plants flower regardless of day-length and yet they flower at characteristic stages. DN Nicotiana tabacum cv. Samsun, makes about forty nodes before flowering. The question still persists whether flowering starts because leaves become physiologically able to export sufficient floral stimulus or the shoot apical meristem (SAM) acquires developmental competence to interpret its arrival. This question was addressed using tobacco expressing the Schizosaccharomyces pombe cell cycle gene, Spcdc25, as a tool. Spcdc25 expression induces early flowering and we tested a hypothesis that this phenotype arises because of premature floral competence of the SAM. Scions of vegetative Spcdc25 plants were grafted onto stocks of vegetative WT together with converse grafts and flowering onset followed (as the time since sowing and number of leaves formed till flowering). Spcdc25 plants flowered significantly earlier with fewer leaves, and, unlike WT, also formed flowers from axillary buds. Scions from vegetative Spcdc25 plants also flowered precociously when grafted to vegetative WT stocks. However, in a WT scion to Spcdc25 stock, the plants flowered at the same time as WT. SAMs from young vegetative Spcdc25 plants were elongated (increase in SAM convexity determined by tracing a circumference of SAM sections) with a pronounced meristem surface cell layers compared with WT. Presumably, Spcdc25 SAMs were competent for flowering earlier than WT and responded to florigenic signal produced even in young vegetative WT plants. Precocious reproductive competence in Spcdc25 SAMs comprised a pronounced mantle, a trait of prefloral SAMs. Hence, we propose that true DN plants export florigenic signal since early developmental stages but the SAM has to acquire competence to respond to the floral stimulus.

Regulatory dephosphorylation of CDK at G2/M in plants: yeast mitotic phosphatase cdc25 induces cytokinin-like effects in transgenic tobacco morphogenesis.

BACKGROUND: During the last three decades, the cell cycle and its control by cyclin-dependent kinases (CDKs) have been extensively studied in eukaryotes. This endeavour has produced an overall picture that basic mechanisms seem to be largely conserved among all eukaryotes. The intricate regulation of CDK activities includes, among others, CDK activation by CDC25 phosphatase at G2/M. In plants, however, studies of this regulation have lagged behind as a plant Cdc25 homologue or other unrelated phosphatase active at G2/M have not yet been identified. SCOPE: Failure to identify a plant mitotic CDK activatory phosphatase led to characterization of the effects of alien cdc25 gene expression in plants. Tobacco, expressing the Schizosaccharomyces pombe mitotic activator gene, Spcdc25, exhibited morphological, developmental and biochemical changes when compared with wild type (WT) and, importantly, increased CDK dephosphorylation at G2/M. Besides changes in leaf shape, internode length and root development, in day-neutral tobacco there was dramatically earlier onset of flowering with a disturbed acropetal floral capacity gradient typical of WT. In vitro, de novo organ formation revealed substantially earlier and more abundant formation of shoot primordia on Spcdc25 tobacco stem segments grown on shoot-inducing media when compared with WT. Moreover, in contrast to WT, stem segments from transgenic plants formed shoots even without application of exogenous growth regulator. Spcdc25-expressing BY-2 cells exhibited a reduced mitotic cell size due to a shortening of the G2 phase together with high activity of cyclin-dependent kinase, NtCDKB1, in early S-phase, S/G2 and early M-phase. Spcdc25-expressing tobacco ('Samsun') cell suspension cultures showed a clustered, more circular, cell phenotype compared with chains of elongated WT cells, and increased content of starch and soluble sugars. Taken together, Spcdc25 expression had cytokinin-like effects on the characteristics studied, although determination of endogenous cytokinin levels revealed a dramatic decrease in Spcdc25 transgenics. CONCLUSIONS: The data gained using the plants expressing yeast mitotic activator, Spcdc25, clearly argue for the existence and importance of activatory dephosphorylation at G2/M transition and its interaction with cytokinin signalling in plants. The observed cytokinin-like effects of Spcdc25 expression are consistent with the concept of interaction between cell cycle regulators and phytohormones during plant development. The G2/M control of the plant cell cycle, however, remains an elusive issue as doubts persist about the mode of activatory dephosphorylation, which in other eukaryotes is provided by Cdc25 phosphatase serving as a final all-or-nothing mitosis regulator.

Tobacco cells transformed with the fission yeast Spcdc25 mitotic inducer display growth and morphological characteristics as well as starch and sugar status evocable by cytokinin application.

In plants, the G2/M control of cell cycle remains an elusive issue as doubts persist about activatory dephosphorylation--in other eukaryotes provided by CDC25 phosphatase and serving as a final all-or-nothing mitosis regulator. We report on the effects of tobacco (Nicotiana tabacum L., cv. Samsun) transformation with fission yeast (Schizosaccharomyces pombe) cdc25 (Spcdc25) on cell characteristics. Transformed cell suspension cultures showed higher dry mass accumulation during the exponential phase and clustered more circular cell phenotypes compared to chains of elongated WT cells. Similar cell parameters, as in the transformants, can be induced in WT by cytokinins. Spcdc25 cells, after cytokinin treatment, showed giant cell clusters and growth inhibition. In addition, Spcdc25 expression led to altered carbohydrate status: increased starch and soluble sugars with higher sucrose:hexoses ratio, inducible in WT by cytokinin treatment. Taken together, the Spcdc25 transformation had a cytokinin-like effect on studied characteristics. However, endogenous cytokinin determination revealed markedly lower cytokinin levels in Spcdc25 transformants. This indicates that the cells sense Spcdc25 expression as an increased cytokinin availability, manifested by changed cell morphology, and in consequence decrease endogenous cytokinin levels. Clearly, the results on cell growth and morphology are consistent with the model of G2/M control including cytokinin-regulated activatory dephosphorylation. Nevertheless, no clear link is obvious between Spcdc25 transformation and carbohydrate status and thus the observed cytokinin-like effect on carbohydrate levels poses a problem. Hence, we propose that Spcdc25-induced higher CDK(s) activity at G2/M generates a signal-modifying carbohydrate metabolism to meet high energy and C demands of forthcoming cell division.

The fission yeast mitotic activator cdc25 and sucrose induce early flowering synergistically in the day-neutral Nicotiana tabacum cv. Samsun.

Here, the tobacco (Nicotiana tabacum) day-neutral (DN) cv. Samsun transformed with the Schizosaccharomyces pombe mitotic activator gene Spcdc25 was used to study the onset of flowering. Wild type (WT) and cdc25 plants were grown from seeds in vitro until they were 20 cm high. Apical and basal nodes were then subcultured repeatedly and the regenerated plants were used to document time to flowering and the number of leaves formed before flowering. Three sucrose treatments (3, 5 or 7% (weight/volume)) were used and measurements of leaf endogenous soluble carbohydrates were performed. In the 3% treatment, cdc25 plants flowered but WT plants did not. The higher sucrose treatments enabled WT flowering; two-thirds of the plants flowered at 5%, while all plants flowered at 7% sucrose. However, in all treatments, cdc25 plants exhibited significantly earlier flowering and fewer leaves compared with wild type. Remarkably, a typical acropetal flowering gradient in WT plants did not occur in cdc25 plants. In cdc25 leaves, there were significantly higher amounts of endogenous sugars with a higher proportion of sucrose compared with WT. Our data demonstrate that Spcdc25 expression and sucrose act synergistically to induce precocious flowering.