Fruit ripening in response to treatments with propylene, 1-methycyclopropene (1-MCP), and low temperature was characterized in ‘Sanuki Gold’ kiwifruit, Actinidia chinensis Planch. Propylene treatment immediately induced rapid fruit softening, increased AC-PG (polygalacturonase) and AC-EXP (expansin) mRNA accumulation, and stimulated an increase in the soluble solid concentration (SSC) and a decrease in titratable acidity (TA). After 3 d exposure to propylene, ethylene production and AC-PL (pectate lyase) mRNA accumulation were observed. 1-MCP treatment after 24 h exposure to propylene eliminated AC-PG mRNA accumulation and suppressed continued changes in SSC and TA. Application of 1-MCP at the start of the treatment, followed by continuous propylene exposure, markedly delayed fruit softening, and the expression of the cell wall-modifying genes, and changes in the SSC and TA, indicating that kiwifruit become insensitive to ethylene at least for 3 d following 1-MCP exposure. Surprisingly, significant fruit softening, mRNA accumulation of AC-PG, AC-PL, and AC-EXP, and decreased TA were observed without ethylene production in intact fruit stored at low temperature for 1 month, but not in fruit stored at room temperature. Repeated 1-MCP treatments (twice a week) failed to inhibit the changes that occurred in low temperature storage. These observations indicate that low temperature modulates the ripening of kiwifruit in an ethylene-independent manner, suggesting that kiwifruit ripening is inducible by either ethylene or low temperature signals.
BackgroundKiwifruit 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.ResultsTo 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.ConclusionsThese 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.Electronic supplementary materialThe online version of this article (10.1186/s12870-018-1264-y) contains supplementary material, which is available to authorized users.
The responses of three kiwifruit cultivars, Actinidia chinensis 'Sanuki Gold', A. chinensis 'Rainbow Red', and A. deliciosa 'Hayward' to various storage temperatures (0, 5, 10, 15, and 20°C) for 8 weeks were investigated. The rate of fruit which initiated ethylene production due to rot development increased with increases in storage temperature. Early-maturing cultivars, 'Rainbow Red' and 'Sanuki Gold' fruit stored at 5, 10, and 15°C showed drastic softening and a decrease in titratable acidity (TA) to an edible level within 4 weeks without detectable ethylene production, whereas fruit stored at 0 and 20°C maintained high firmness and TA even after 8 weeks unless they were infected with rot. A late-maturing cultivar, 'Hayward' fruit stored at 5 and 10°C softened more rapidly than when stored at 0, 15, or 20°C. Treatment with 1-Methylcyclopropene (1-MCP) did not suppress the low temperature modulated fruit ripening in any cultivars, indicating its independence from ethylene. These results suggest that 'Sanuki Gold' and 'Rainbow Red' are more sensitive to low temperatures compared to 'Hayward' and the sensitivity is involved in the determination of storage life and how early the fruit matures on the vine.
In order to extend the "eating window", the optimum ripening phase suitable for eating, the combination of treatment with propylene (an ethylene analog) and 1-methylcyclopropene (1-MCP; an ethylene inhibitor) was assessed in three kiwifruit cultivars: 'Rainbow Red' Actinidia chinensis, 'Sanuki Gold' A. chinensis, and 'Hayward' A. deliciosa. Propylene treatment initiated the ripening process by inducing fruit softening, increasing soluble solid content (SSC), and decreasing titratable acids (TA), with or without endogenous ethylene production, depending on the duration of exposure. Once endogenous ethylene was induced, it accelerated fruit ripening, resulting in an over-ripening phase and shortening of the "eating window". 'Rainbow Red' and 'Sanuki Gold' fruit treated with propylene continuously or for 48 h initiated endogenous ethylene production that led to an "eating window" lasting only 2 days (range of 3-5 days after the start of treatment), whereas it lasted for 7 days (range 3-10 days) in 'Hayward' fruit. Limited propylene treatment of the three cultivars for 24 h induced ripening without the detection of ethylene production, suggesting that the optimum ripening phase suitable for eating can be attained without endogenous ethylene production, resulting in a longer "eating window". 'Rainbow Red' and 'Sanuki Gold' fruit treated with propylene for 48 h followed by 1-MCP treatment had extended "eating window" and shelf-life, with the suppression of endogenous ethylene. These results illustrate the practicability of different durations of propylene treatment in facilitating kiwifruit ripening and the additional benefit of 1-MCP treatment to the extend shelf-life of new high-quality kiwifruit cultivars, 'Rainbow Red' and 'Sanuki Gold'.
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