Leaf senescence is a genetically controlled dismantling programme that enables plants to efficiently remobilise nutrients to new growing sinks. It involves substantial metabolic reprogramming whose timing is affected by developmental and environmental signals. Plant hormones have long been known to affect the timing of leaf senescence, but they also affect plant development and stress responses. It has therefore been difficult to tease apart how the different hormones regulate the onset and progression of leaf senescence, i.e., whether they directly affect leaf senescence or affect it indirectly by altering the developmental programme or by altering plants' response to stress. Here we review research on hormonal regulation of leaf senescence and propose that hormones affect senescence through differential responses to developmental and environmental signals. We suggest that leaf senescence strictly depends on developmental changes, after which senescence can be induced, depending on the type of hormonal and environmental cues.

Hormonal regulation of leaf senescence through integration of developmental and stress signals.

https://repositorio.iniap.gob.ec/bitstream/41000/4838/1/iniapscR2016v250p188.pdf

Saponin determination, expression analysis and functional characterization of saponin biosynthetic genes in Chenopodium quinoa leaves.

Leaf senescence is an important developmental programmed degeneration process that dramatically affects crop quality and yield. The regulation of senescence is highly complex. Although senescence regulatory genes have been well characterized in model species such as Arabidopsis and rice, there is little information on the control of this process in cotton. Here, the senescence process in cotton (Gossypium hirsutum L.) leaves was investigated over a time course including young leaf, mature leaf and leaf samples from different senescence stages using RNA-Seq. Of 24,846 genes detected by mapping the tags to Gossypium genomes, 3,624 genes were identified as differentially expressed during leaf senescence. There was some overlap between the genes identified here and senescence-associated genes previously identified in other species. Most of the genes related to photosynthesis, chlorophyll metabolism and carbon fixation were downregulated; whereas those for plant hormone signal transduction were upregulated. Quantitative real-time PCR was used to evaluate the results of RNA-Seq for gene expression profiles. Furthermore, 519 differentially expressed transcription factors were identified, notably WRKY, bHLH and C3H. In addition, 960 genes involved in the metabolism and regulation of eight hormones were identified, of which many genes involved in the abscisic acid, brassinosteroid, jasmonic acid, salicylic acid and ethylene pathways were upregulated, indicating that these hormone-related genes might play crucial roles in cotton leaf development and senescence. However, most auxin, cytokinin and gibberellin pathway-related genes were downregulated, suggesting that these three hormones may act as negative regulators of senescence. This is the first high-resolution, multiple time-course, genome-wide comprehensive analysis of gene expression in cotton. These data are the most comprehensive dataset currently available for cotton leaf senescence, and will serve as a useful resource for unraveling the functions of many specific genes involved in cotton leaf development and senescence.

https://link.springer.com/content/pdf/10.1186%2Fs12870-015-0433-5.pdf

Global analysis of the Gossypium hirsutum L. Transcriptome during leaf senescence by RNA-Seq.

Many plant pathogens show interactions with host development. Pathogens may modify plant development according to their nutritional demands. Conversely, plant development influences pathogen growth. Biotrophic pathogens often delay senescence to keep host cells alive, and resistance is achieved by senescence-like processes in the host. Necrotrophic pathogens promote senescence in the host, and preventing early senescence is a resistance strategy of plants. For hemibiotrophic pathogens both patterns may apply. Most signaling pathways are involved in both developmental and defense reactions. Increasing knowledge about the molecular components allows to distinguish signaling branches, cross-talk and regulatory nodes that may influence the outcome of an infection. In this review, recent reports on major molecular players and their role in senescence and in pathogen response are reviewed. Examples of pathosystems with strong developmental implications illustrate the molecular basis of selected control strategies. A study of gene expression in the interaction between the hemibiotrophic vascular pathogen Verticillium longisporum and its cruciferous hosts shows processes that are fine-tuned to counteract early senescence and to achieve resistance. The complexity of the processes involved reflects the complex genetic control of quantitative disease resistance, and understanding the relationship between disease, development and resistance will support resistance breeding.

https://www.mdpi.com/2223-7747/4/3/449/pdf

Keeping Control: The Role of Senescence and Development in Plant Pathogenesis and Defense

Leaf senescence is a degenerative process that is genetically controlled and involves nutrient remobilization prior to the death of leaf tissues. Age is a key developmental determinant of the process along with other senescence inducing factors. At the cellular level, different hormones, signaling molecules, and transcription factors contribute to the regulation of senescence. This review summarizes the recent progress in understanding the complexity of the senescence process with primary focuses on perception and transduction of senescence signals as well as downstream regulatory events. Future directions in this field and potential applications of related techniques in crop improvement will be discussed.

Signal Transduction in Leaf Senescence: Progress and Perspective

Paris polyphylla Smith var. yunnanensis (Franch.) Hand.-Mazz. is a rhizomatous, herbaceous, perennial plant that is used as a medicinal plant with a variety of pharmacological activities. However, the functions of the green, leafy sepal of this plant are poorly understood. The main objectives of this study were to: (a) test the hypothesis that sepals make measurable contributions to fruit development and rhizome growth; and (b) investigate the allocation and partitioning of photosynthates produced by sepals and leaves to fruit and rhizome. Net photosynthetic rate, photosynthetic pigment composition and δ13C values were similar for sepals and leaves. Sepal-darkening and sepal-removal treatments resulted in smaller fruit size and decreased rhizome biomass, whereas fruit removal led to a decrease in calyx size and an increase in rhizome yield and saponin content. Fruit and seed mass were positively and linearly related to calyx size. These results indicate that photosynthates produced by sepals are involved in the fruit growth and seed development and that developing fruit and rhizomes compete for the photosynthates exported by leaves. We propose that the sepals of P. polyphylla function partly as leaves to compensate for reproductive costs. Fruit removal increased carbon partitioning to the rhizome and improved rhizome yield and quality, offering a useful strategy for the domestication of this valuable medicinal plant.

Function of leafy sepals in Paris polyphylla: photosynthate allocation and partitioning to the fruit and rhizome.

Senescence in perennials has not been extensively studied compared to that in annual plants, and the effects of delaying senescence on plant biomass and metabolic features in herbaceous perennials has also been poorly reported. We recently examined the effects of gibberellin A3 (GA3) on senescence of aerial parts of Paris polyphylla, which characterize the metabolic changes associated with senescence, and found antagonistic effects of GA and abscisic acid during this process. Rhizome yield and quality (saponin content) of Paris polyphylla were both improved by GA3-induced retardation of senescence. We propose that GA3-induced retardation of senescence increases green leaf area and prolongs the duration of photosynthesis, leading to increased rhizome yield. The increased saponin accumulation in GA3-treated plants may be explained by the longer growth phase and ensuing increased environmental stress.

Improving rhizome yield and quality of Paris polyphylla through gibberellic acid-induced retardation of senescence of aerial parts.

Rhizomes of the perennial herb Paris polyphylla have been used in traditional Chinese medicine for hundreds of years. Agricultural production of the rhizomes requires 7-10 years, which is too long to meet the demand of the medicinal industry. Therefore, studies on improving the yield of the herb and shortening the culturing period are imperative. The present work aimed to investigate the effect of sodium bisulfite (NaHSO (3)) on rhizome yield and quality, as well as some related metabolic features of P. polyphylla. The rhizome yield was improved by NaHSO (3) treatment in long-term experiments conducted during 2006 and 2007, with 2 mM NaHSO (3) giving the highest yield. HPLC analysis revealed that NaHSO (3) treatment increased the total saponin content (49 %), including three pennogenin glycosides and two diosgenin glycosides. In a short-term experiment, NaHSO (3) treatment resulted in an enhanced net photosynthetic rate (Pn) for about 4 days without significant changes in the chlorophyll or carotenoid content. The total soluble sugars and sucrose contents in the leaves also significantly increased after 2 mM NaHSO (3) treatment, whereas the starch content changed only slightly. The activities of the enzymes involved in ammonium assimilation (glutamine synthetase [GS] and glutamate dehydrogenase [GDH]) were not significantly influenced. In a long-term experiment, chlorophylls and carotenoids were not significantly affected, and neither was the starch content in leaves, but the total soluble sugars and sucrose contents in leaves increased significantly. The NaHSO (3) treatment significantly increased GS and GDH activities. These results indicate that NaHSO (3) treatment improved the rhizome yield in P. polyphylla, not only through enhancement of Pn but also by improving carbohydrate accumulation and ammonium assimilation. The increased saponin content after NaHSO (3) treatment was indicative of high rhizome quality.

Sodium bisulfite improves rhizome yield and quality in Paris polyphylla.

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Function of leafy sepals in Paris polyphylla: photosynthate allocation and partitioning to the fruit and rhizome.

Improving rhizome yield and quality of Paris polyphylla through gibberellic acid-induced retardation of senescence of aerial parts.

Sodium bisulfite improves rhizome yield and quality in Paris polyphylla.