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Background Kiwifruit are classified as climacteric since exogenous ethylene (or its analogue propylene) induces rapid ripening accompanied by ethylene production under positive feedback regulation. However, most of the ripening–associated changes (Phase 1 ripening) in kiwifruit during storage and on–vine occur largely in the absence of any detectable ethylene. This ripening behavior is often attributed to basal levels of system I ethylene, although it is suggested to be modulated by low temperature. Results To elucidate the mechanisms regulating Phase 1 ripening in kiwifruit, a comparative transcriptome analysis using fruit continuously exposed to propylene (at 20 °C), and during storage at 5 °C and 20 °C was conducted. Propylene exposure induced kiwifruit softening, reduction of titratable acidity (TA), increase in soluble solids content (SSC) and ethylene production within 5 days. During storage, softening and reduction of TA occurred faster in fruit at 5 °C compared to 20 °C although no endogenous ethylene production was detected. Transcriptome analysis revealed 3761 ripening–related differentially expressed genes (DEGs), of which 2742 were up–regulated by propylene while 1058 were up–regulated by low temperature. Propylene exclusively up–regulated 2112 DEGs including those associated with ethylene biosynthesis and ripening such as AcACS1 , AcACO2 , AcPL1 , AcXET1 , Acβ–GAL , AcAAT , AcERF6 and AcNAC7 . Similarly, low temperature exclusively up–regulated 467 DEGS including AcACO3 , AcPL2 , AcPMEi , AcADH , Acβ–AMY2 , AcGA2ox2 , AcNAC5 and AcbZIP2 among others. A considerable number of DEGs such as AcPG , AcEXP1 , AcXET2 , Acβ–AMY1 , AcGA2ox1 , AcNAC6 , AcMADS1 and AcbZIP1 were up–regulated by either propylene or low temperature. Frequent 1–MCP treatments failed to inhibit the accelerated ripening and up–regulation of associated DEGs by low temperature indicating that the changes were independent of ethylene. On–vine kiwifruit ripening proceeded in the absence of any detectable endogenous ethylene production, and coincided with increased expression of low temperature–responsive DEGs as well as the decrease in environmental temperature. Conclusions These results indicate that kiwifruit possess both ethylene−dependent and low temperature–modulated ripening mechanisms that are distinct and independent of each other. The current work provides a foundation for elaborating the control of these two ripening mechanisms in kiwifruit.

Background Methylation of nucleotides, notably in the forms of 5-methylcytosine (5mC) in DNA and N 6 -methyladenosine (m 6 A) in mRNA, carries important information for gene regulation. 5mC has been elucidated to participate in the regulation of fruit ripening, whereas the function of m 6 A in this process and the interplay between 5mC and m 6 A remain uncharacterized. Results Here, we show that mRNA m 6 A methylation exhibits dynamic changes similar to DNA methylation during tomato fruit ripening. RNA methylome analysis reveals that m 6 A methylation is a prevalent modification in the mRNA of tomato fruit, and the m 6 A sites are enriched around the stop codons and within the 3′ untranslated regions. In the fruit of the ripening-deficient epimutant Colorless non-ripening ( Cnr ) which harbors DNA hypermethylation, over 1100 transcripts display increased m 6 A levels, while only 134 transcripts show decreased m 6 A enrichment, suggesting a global increase in m 6 A. The m 6 A deposition is generally negatively correlated with transcript abundance. Further analysis demonstrates that the overall increase in m 6 A methylation in Cnr mutant fruit is associated with the decreased expression of RNA demethylase gene SlALKBH2 , which is regulated by DNA methylation. Interestingly, SlALKBH2 has the ability to bind the transcript of SlDML2 , a DNA demethylase gene required for tomato fruit ripening, and modulates its stability via m 6 A demethylation. Mutation of SlALKBH2 decreases the abundance of SlDML2 mRNA and delays fruit ripening. Conclusions Our study identifies a novel layer of gene regulation for key ripening genes and establishes an essential molecular link between DNA methylation and mRNA m 6 A methylation during fruit ripening.

Background Recently, DNA methylation was proposed to regulate fleshy fruit ripening. Fleshy fruits can be distinguished by their ripening process as climacteric fruits, such as tomatoes, or non-climacteric fruits, such as strawberries. Tomatoes undergo a global decrease in DNA methylation during ripening, due to increased expression of a DNA demethylase gene. The dynamics and biological relevance of DNA methylation during the ripening of non-climacteric fruits are unknown. Results Here, we generate single-base resolution maps of the DNA methylome in immature and ripe strawberry. We observe an overall loss of DNA methylation during strawberry fruit ripening. Thus, ripening-induced DNA hypomethylation occurs not only in climacteric fruit, but also in non-climacteric fruit. Application of a DNA methylation inhibitor causes an early ripening phenotype, suggesting that DNA hypomethylation is important for strawberry fruit ripening. The mechanisms underlying DNA hypomethylation during the ripening of tomato and strawberry are distinct. Unlike in tomatoes, DNA demethylase genes are not upregulated during the ripening of strawberries. Instead, genes involved in RNA-directed DNA methylation are downregulated during strawberry ripening. Further, ripening-induced DNA hypomethylation is associated with decreased siRNA levels, consistent with reduced RdDM activity. Therefore, we propose that a downregulation of RdDM contributes to DNA hypomethylation during strawberry ripening. Conclusions Our findings provide new insight into the DNA methylation dynamics during the ripening of non-climacteric fruit and suggest a novel function of RdDM in regulating an important process in plant development.

We review current knowledge of the pheophorbide a oxygenase/phyllobilin pathway of chlorophyll breakdown, with particular focus on its biochemistry and the transcriptional regulation of chlorophyll catabolic genes. Abstract Chlorophyll breakdown is one of the most obvious signs of leaf senescence and fruit ripening. The resulting yellowing of leaves can be observed every autumn, and the color change of fruits indicates their ripening state. During these processes, chlorophyll is broken down in a multistep pathway, now termed the 'PAO/phyllobilin' pathway, acknowledging the core enzymatic breakdown step catalysed by pheophorbide a oxygenase, which determines the basic linear tetrapyrrole structure of the products of breakdown that are now called 'phyllobilins'. This review provides an update on the PAO/phyllobilin pathway, and focuses on recent biochemical and molecular progress in understanding phyllobilin-modifying reactions as the basis for phyllobilin diversity, on the evolutionary diversity of the pathway, and on the transcriptional regulation of the pathway genes.

Coccinia grandis contains secondary metabolites, such as flavonoids, phenolic acids, terpenoids, alkaloids, sterols, and glycosides, which are known to have in vitro antioxidant, antidiabetic, anti-inflammatory, and antidyslipidemic activities. C. grandis fruits change dramatically during ripening, and the differences in the phytochemicals contribute to various uses. This study reports the phytochemical compounds and antioxidant activities during ripening of C. grandis for the first time. Characterizations were conducted on the physiologically active substances in C. grandis fruits at three ripening stages, and a total of 25 peaks were identified. Key phytochemicals in the ripening stages of C. grandis were identified, and the major substances that contributed to antioxidant properties were selected and quantitatively analyzed. Although the concentration of tiliroside increased during aging, hydroxycinnamic acid (chlorogenic and p-coumaric acids), flavonols (rutin), and triterpenes (cucurbitacins B and D) with antioxidant effects decreased. Therefore, phenolic compounds and cucurbitacins dominate immature C. grandis quantitatively. Regarding phytohormones, the gibberellin A4 content decreased as the fruits matured, but indoleacetic acid and salicylic acid increased with fruit maturity. The antioxidant capacities determined by DPPH and ABTS consistently decreased with increasing maturity. Accordingly, the extracts of immature C. grandis fruits have high levels of bioactive compounds and can be used to develop food additives and health supplements.

Nanobubbles have many potential applications depending on their types. The long-term stability of different gas nanobubbles is necessary to be studied considering their applications. In the present study, five kinds of nanobubbles (N2, O2, Ar + 8%H2, air and CO2) in deionized water and a salt aqueous solution were prepared by the hydrodynamic cavitation method. The mean size and zeta potential of the nanobubbles were measured by a light scattering system, while the pH and Eh of the nanobubble suspensions were measured as a function of time. The nanobubble stability was predicted and discussed by the total potential energies between two bubbles by the extended Derjaguin–Landau–Verwey–Overbeek (DLVO) theory. The nanobubbles, except CO2, in deionized water showed a long-term stability for 60 days, while they were not stable in the 1 mM (milli mol/L) salt aqueous solution. During the 60 days, the bubble size gradually increased and decreased in deionized water. This size change was discussed by the Ostwald ripening effect coupled with the bubble interaction evaluated by the extended DLVO theory. On the other hand, CO2 nanobubbles in deionized water were not stable and disappeared after 5 days, while the CO2 nanobubbles in 1 mM of NaCl and CaCl2 aqueous solution became stable for 2 weeks. The floating and disappearing phenomena of nanobubbles were estimated and discussed by calculating the relationship between the terminal velocity of the floating bubble and bubble size.

Tomato ( L.) belongs to the Solanaceae family and is the second most important fruit or vegetable crop next to potato ( L.). It is cultivated for fresh fruit and processed products. Tomatoes contain many health-promoting compounds including vitamins, carotenoids, and phenolic compounds. In addition to its economic and nutritional importance, tomatoes have become the model for the study of fleshy fruit development. Tomato is a climacteric fruit and dramatic metabolic changes occur during its fruit development. In this review, we provide an overview of our current understanding of tomato fruit metabolism. We begin by detailing the genetic and hormonal control of fruit development and ripening, after which we document the primary metabolism of tomato fruits, with a special focus on sugar, organic acid, and amino acid metabolism. Links between primary and secondary metabolic pathways are further highlighted by the importance of pigments, flavonoids, and volatiles for tomato fruit quality. Finally, as tomato plants are sensitive to several abiotic stresses, we briefly summarize the effects of adverse environmental conditions on tomato fruit metabolism and quality.

• The regulatory network of R2R3 MYB transcription factors in anthocyanin biosynthesis is not fully understood in blue-coloured berries containing delphinidin compounds. • We used blue berries of bilberry (Vaccinium myrtillus) to comprehensively characterise flavonoid-regulating R2R3 MYBs, which revealed a new type of co-regulation in anthocyanin biosynthesis between members of MYBA-, MYBPA1- and MYBPA2-subgroups. • VmMYBA1, VmMYBPA1.1 and VmMYBPA2.2 expression was elevated at berry ripening and by abscisic acid treatment. Additionally, VmMYBA1 and VmMYBPA1.1 expression was strongly downregulated in a white berry mutant. Complementation and transient overexpression assays confirmed VmMYBA1 and VmMYBA2 to induce anthocyanin accumulation. Promoter activation assays showed that VmMYBA1, VmMYBPA1.1 and VmMYBPA2.2 had similar activity towards dihydroflavonol 4-reductase (DFR) and anthocyanidin synthase (ANS), but differential regulation activity for UDP-glucose flavonoid 3-O-glucosyltransferase (UFGT) and flavonoid 3′5′-hydroxylase (F3′5′H) promoters. Silencing of VmMYBPA1.1 in berries led to the downregulation of key anthocyanin and delphinidin biosynthesis genes. Functional analyses of other MYBPA regulators, and a member of novel MYBPA3 subgroup, associated them with proanthocyanidin biosynthesis and F3′5′H expression. • The existence of 18 flavonoid-regulating MYBs indicated gene duplication, which may have enabled functional diversification among MYBA, MYBPA1 and MYBPA2 subgroups. Our results provide new insights into the intricate regulation of the complex anthocyanin profile found in blue-coloured berries involving regulation of both cyanidin and delphinidin branches.

Living cells use phase separation and concentration gradients to organize chemical compartments in space. Here, we present a theoretical study of droplet dynamics in gradient systems. We derive the corresponding growth law of droplets and find that droplets exhibit a drift velocity and position dependent growth. As a consequence, the dissolution boundary moves through the system, thereby segregating droplets to one end. We show that for steep enough gradients, the ripening leads to a transient arrest of droplet growth that is induced by a narrowing of the droplet size distribution.