猕猴桃果实采后冷害发生生理机制及调控作用
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摘要
猕猴桃果实皮薄、汁多、营养丰富，是人们喜爱的水果之一。猕猴桃为呼吸跃变型桨果，常温下货架期短易软烂，低温能延长其贮藏期，但猕猴桃果实对低温较敏感，冷藏条件下很容易诱发冷害。而冷害症状在低温状态下多不表现，待贮藏温度上升或脱离低温环境后，其冷害症状才逐步表现。冷害导致贮藏中和出库后货架期大量腐烂，这已成为猕猴桃果实采后冷链物流的最大障碍。目前猕猴桃果实采后研究多集中在生理变化特点及贮藏保鲜技术上，有关其果实冷害鲜有报道。因此研究猕猴桃果实冷害发生机理，并运用适当的调控措施减轻或抑制冷害，对改善和提高采后猕猴桃果实贮运技术水平，提高流通效益具有重要的理论和实践意义。
本论文研究不同采收成熟度、逐步降温和低温预贮（low temperature conditioning,LTC）两种物理降温方式以及外源腐胺（Put）化学处理对猕猴桃果实冷害发生的影响及其机制，为猕猴桃贮运技术体系的建立提供参考。研究结果分述如下：
1.研究了不同成熟度对‘徐香’猕猴桃果实冷害、品质、呼吸、乙烯、相对膜透性、丙二醛含量和抗氧化酶活性的影响。结果显示：（1）Ⅰ可溶性固形物（TSS）:4.5%～5.5%）、Ⅱ（TSS:6.5%～7.5%）和Ⅲ（TSS:8.0%～9.0%）3个不同采收成熟度的‘徐香’果实低温贮藏效果和冷害程度差异较大：采收Ⅰ果实失重率和腐烂率较高，较早表现木质化、水渍化等冷害症状，冷害率和冷害指数也较高，贮后货架期结束时硬度、可滴定酸、可溶性固性物和维生素C含量明显下降。而Ⅱ和Ⅲ两采收期出现木质化和水渍化等冷害症状较晚，冷害率和冷害指数较低，而且在贮后货架期结束时仍保持较高可溶性固形物和维生素C含量。与Ⅱ相比较，Ⅲ采收期果实的失重率和腐烂率较高，贮后货架期结束时果肉硬度、可滴定酸和维生素C含量较小，因此，‘徐香’果实适宜贮藏的采收期为Ⅱ。（2）猕猴桃果实发生冷害时呼吸速率、乙烯释放量和相对膜透性均异常增加。而Ⅱ采收期‘徐香’果实显著降低呼吸和乙烯释放，并保持较高的超氧化物歧化酶（SOD）、过氧化物酶（POD）、过氧化氢酶（CAT）和抗坏血酸过氧化物酶（APX）等抗氧化酶活性，抑制膜脂过氧化程度，从而减轻果实冷害的发生。这些结果说明Ⅱ采收成熟度果实冷害相对较低可能与保持较高的抗氧化酶活性有关。
2.首先研究了不同降温处理对‘徐香’果实冷害、失重率、腐烂率和品质的影响，发现逐步降温可显著降低‘徐香’猕猴桃冷害指数、冷害率、腐烂率和失重率，并保持较高的果肉硬度、可溶性固性物、可滴定酸和维生素C含量，其中逐步降温2（10℃→5℃2d→2℃2d→0±0.5℃）的效果最好。
在上述研究的基础上，选用逐步降温2探索逐步降温处理对减轻‘徐香’果实冷害的作用，通过研究逐步降温处理对‘徐香’果实呼吸、乙烯和活性氧代谢的影响，以揭示其减轻果实冷害的机理。结果显示：‘徐香’果实发生冷害时呼吸速率、乙烯释放量和相对膜透性均异常增加，SOD、CAT、POD、APX等酶活性降低，丙二醛（MDA）大量累积。逐步降温处理有效降低冷藏‘徐香’果实的冷害指数和冷害率，抑制呼吸速率和乙烯释放量的增加，保持较高的SOD、CAT、POD、APX酶活性，并降低O_2~(·—)和H_2O_2含量，显著减少膜脂过氧化产物MDA的积累，抑制相对膜透性的增加。这说明，逐步降温减轻‘徐香’果实冷害与活性氧清除酶活性的提高及活性氧积累的降低有关。
3.研究了不同LTC处理对‘海沃德’果实冷害、失重率、腐烂率和品质的影响，发现LTC可显著降低‘海沃德’冷害指数、冷害率、腐烂率和失重率，并保持较高的果肉硬度、可滴定酸、可溶性固性物和维生素C的含量，其中LTC5（12℃，3d）处理的效果最好。在此基础上，通过研究LTC5处理对‘海沃德’果实活性氧代谢、脂氧合酶（LOX）活性和內源激素的影响，以掲示LTC处理减轻‘海沃德’果实冷害的作用机制。结果表明，LTC处理保持较高的SOD、CAT、POD及APX抗氧化酶活性和较低的LOX酶活性，并降低O_2~(·—)生成速率和H_2O_2含量，抑制MDA和相对膜透性上升，同时积累较低GA_3，并保持较高ABA、IAA、ZR水平和ABA/IAA、ABA/GA_3比例，最终减轻‘海沃德’果实的冷害症状。这些结果说明，LTC处理可维持活性氧代谢平衡，积累较少的GA_3，并保持较高ABA、IAA、ZR水平和ABA/IAA、ABA/GA_3比例，从而减轻果实冷害的发生。
4.研究了不同浓度（1mmol·L~(-1)、2mmol·L~(-1)、4mmol·L~(-1)）Put处理对‘红阳’果实冷害、品质、腐烂率和失重率的影响，结果发现2mmol·L~(-1)Put处理可显著减少‘红阳’果肉木质化和水渍化等冷害症状的表现，减少果实失重率和腐烂率，并保持较高果肉硬度、可滴定酸和维生素C含量。在此基础上研究了2mmol·L~(-1)Put处理对‘红阳’呼吸、乙烯、活性氧代谢、ASA-GSH循环，內源多胺、LOX酶活性以及膜脂脂肪酸的影响，以掲示Put处理减轻‘红阳’果实冷害的机理。结果显示：
（1）猕猴桃果实的冷害与内源亚精胺（Spd）、精胺（Spm）含量的降低有关。内源Spd和Spm的积累水平与冷害的发生程度存在明显的负相关性，相关系数分别为R=–0.9255和R=–0.9133。而Put处理提高‘红阳’内源Put、Spd、Spm含量，说明外源Put诱导了內源多胺含量的增加，从而有助于减轻猕猴桃果实冷害。
（2）Put处理显著抑制了‘红阳’果实呼吸速率和乙烯释放速率，保持较高POD、SOD、CAT、APX、脱氢抗坏血酸还原酶（DHAR）、单脱氢抗坏血酸还原酶（MDHAR）、谷胱甘肽还原酶（GR）活性及还原型谷胱甘肽（GSH）、抗坏血酸（ASA）抗氧化物质含量，积累较少的氧化型谷胱甘肽（GSSG）和脱氢抗坏血酸（DHA），保持较高的GSH/GSSG和ASA/DHA比例，并抑制O_2~(·—)和H_2O_2积累，降低了果实中MDA含量和相对膜透性，减轻‘红阳’果实冷害。说明， Put处理提高‘红阳’果实抗冷性与保持较高的抗氧化物质含量和抗氧化酶活性以及降低膜脂过氧化程度密切相关。
（3）猕猴桃果实的冷害与LOX酶活性上升以及亚油酸、亚麻酸含量的降低密切相关。 Put处理抑制‘红阳’果实LOX活性上升，提高了亚油酸、亚麻酸不饱和脂肪酸相对含量，降低了棕榈酸和硬脂酸等饱和脂肪酸相对含量，抑制膜脂肪酸不饱和度和不饱和指数下降。说明，Put处理是通过降低‘红阳’果实LOX活性，抑制膜脂不饱和脂肪酸相对含量的下降，保持较高的膜脂肪酸不饱和度和不饱和指数，从而增强了‘红阳’果实抗冷性，减轻果实冷害的发生。
Kiwifruit as an important fruit is popular for its thin skin, juiceful texture and rich innutrient. The kiwifruit, as a typical kind of respiratory climactic fruit, undergoes rapidsoftening with severe decay after harvest, which shortens its shelf life. Cold storageeffectively inhibits fruit softening and extends the postharvest life of kiwifruits. However,kiwifruit is susceptible to chilling injury when protractedly stored at inaptitude temperatures.CI symptoms of kiwifruit are characterized by the appearance of concave spot in the skin,grainy tissue in the outer pericarp and the development of diffuse pitting in association with awater-soaked appearance. These symptoms become most severe after fruit is transferred toambient temperature. CI increase susceptibility to decay, which become a major obstacle topostharvest storage and transportation. At present, most study focus on storage of kiwifruit.Little information is available on the CI of kiwifruit. Thus, it is crucial to study themechanism and regulation of chilling injury in postharvest kiwifruit for storage andtransportation.
In this paper，it was investigated that the effect of different harvested maturity, gradualcooling, low temperature conditioning (LTC) and exogenous putrescine (Put) on chillinginjury in kiwifruit and the possible mechanisms. The results were as follows:
1. The effects of different harvested maturities on chilling injury, fruit quality，respirationand ethylene production, relative membrane permeability, malonaldehyde (MDA) content andthe activities of antioxidant enzymes in kiwifruit (Actinidia deliciosa cv.‘Xuxiang’) duringcold storage were investigated. The results showed as follows:
(1) The storage qualities and chilling injury of kiwifruit were much difference amongⅠ (TSS:4.5%～5.5%), Ⅱ(TSS:6.5%～7.5%), Ⅲ(TSS:8.0%～9.0%) in low temperaturestorage. Compared to fruits of Ⅱ and Ⅲ, the fruits of Ⅰmaturity showed early CI symptomscharacterized by flesh grainy and water-soaked, maintained higher weight loss ratio, decayrate, chilling injury index and chilling injury incidence. In addition, the fruits ofⅠ maturityshowed a decrease in firmness, total soluble solids (TSS), titratable acidity (TA) and VCat theend of storage and shelf life. However, the fruits of Ⅲmaturity maintained higher weight lossratio, decay rate, and lower firmness, TA and VCthan these of Ⅱ maturity. Thus，the best harvest maturity grade for kiwifruit storage performed well was Ⅱmaturity.
(2) The respiration rate, ethylene production and the relative membrane penetrabilityabnormally increased under chilling stress. However, the fruits of Ⅱm aturity maintainedlower respiration and ethylene production, induced the increase in the activities of superoxidedismutase (SOD), ascorbate peroxidase (APX), catalase (CAT) and peroxidase (POD),reduced lipid peroxidation and alleviated chilling injury. These results indicated thatⅡmaturity retard chilling injury in ‘Xuxiang’ fruit which may be related to enhancing theactivities of antioxidant enzymes.
2. The ‘Xuxiang’ Kiwifruit were treated by direct cooling and gradual cooling todeterminate the effect of cooling pattern on chilling injury, weight loss ratio, decay rate andfruit quality. The gradual cooling treatment significantly reduced chilling injury index,chilling injury incidence, weight loss ratio and decay rate, maintained higher firmness, TSS,VCand TA. It was found that treatment2(10℃→5℃2d→2℃2d→0±0.5℃) was mosteffective in ‘Xuxiang’ fruit.
On the basis of above results, the gradual cooling2in ‘Xuxiang’ fruit was chosen toinvestigate the effects of gradual cooling on chilling injury and the influence on active oxygenmetabolism. The results showed that the respiration rate, ethylene production, relativemembrane permeability and malonaldehyde (MDA) contents increased, while the activities ofCAT, SOD, POD and APX decreased during the development of CI symptom. However，thegradual cooling treatment significantly alleviated chilling injury, reduced respiration andethylene production, and inhibited the accumulation of MDA and the increase in relativemembrane permeability. Moreover，the gradual cooling treatment maintained higher activitiesof SOD, CAT, APX and POD than direct cooling fruit during the storage, and kept lowerlevels of superoxide anion production rate (O_2~(·—)) and H_2O_2content. These results indicatedthat gradual cooling treatment retard chilling injury in ‘Xuxiang’ fruit which may be related toenhancing the activities of active oxygen scavenging enzymes and reducing the accumulationof active oxygen.
3. The effects of LTC on chilling injury, weight loss ratio, decay rate and fruit quality inkiwifruit (Actinidia deliciosa cv.‘Hayward’) during cold storage were investigated. Thetreatment with LTC was more effective in reducing chilling injury index and chilling injuryincidence, while maintaining higher level of firmness, TSS, TA and VC. We found that LTC5(12℃，3d) treatment was the optimal treatment to alleviate chilling injury and maintainhigher quality of kiwifruit. Thus LTC5treatment was chosen to to test the effects of LTCtreatment on chilling injury and the possible mechanisms in kiwifruit. The results showed thatLTC treatment inhibited the accumulation of MDA and the increase in relative membrane permeability. Furthermore, LTC treatment decreased accumulation of H_2O_2and O_2~(·—)and theactivity in lipoxygenase (LOX), while increased the activities of CAT, SOD, POD and APX inkiwifruit. The treatment also maintained higher levels of the endogenous ABA, IAA and ZR,lower GA_3levels, and higher ratio of ABA/GA_3and ABA/IAA. These results indicated thatthe effect of LTC treatment on alleviating CI of kiwifruit may be due to its ability to improveantioxidant enzyme activities and reduce ROS accumulation, maintain higher levels of theendogenous ABA, IAA and ZR, lower GA_3levels, and higher ratios of ABA/GA_3andABA/IAA.
4. Kiwifruit (Actinidia chinensis cv.‘Hongyang’) were treated with differentconcentrations of Put (1mmol·L~(-1)，2mmol·L~(-1)，4mmol·L~(-1)) to investigate the effect onchilling injury, weight loss ratio, decay rate and fruit quality. We found that treatment with2mmol·L~(-1)Put treatment was most effective in reducing in CI symptoms of grainy andwater-soaked, decreasing weight loss ratio and decay rate and maintaining higher level offirmness, TA and VC. Thus，2mmol·L~(-1)Put treatment was chosen to investigate the possiblemechanisms of alleviating chilling injury. The results showed as follows:(1) Chilling injurywas related to the reduction of endogenesis Spd and Spm of kiwifruit. The accumulation levelof endogenesis Spm and Spd and chilling injury index appeared remarkable negativecorrelation, R=–0.9255and R=–0.9133. However, exogenous Put treatment increased thecontents of endogenesis Put、Spd and Spm. These results indicated that exogenous Puttreatment reducing in chilling injury may be related to improving endogenous polyaminecontent.
(2) Exogenous Put treatment alleviated chilling injury, inhibited respiration andethylene production, maintained higher level of GSH and ASA and lower level of GSSG andDHA, and kept higher ratio of GSH/GSSG and ASA/DHA. The activities of SOD、POD、CAT、APX、DHAR、MDHAR、GR were also induced by exogenous Put treatment. In addition,this treatment inhibited the accumulation of MDA, H_2O_2and O_2~(·—)and the increase inrelative membrane permeability. These results suggested that the effect of exogenous Puttreatment on alleviating chilling injury of kiwifruit may be attributed to its ability to enhanceantioxidant enzyme activities and the content of antioxidants and decrease lipid peroxidation.
(3) Chilling injury was related to increase of LOX activity and the decrease of linoleicacid (C18:2) and linolenic acid (C18:3). Exogenous Put treatment inhibited increased of LOXactivity, maintained higher relative contents of linoleic acid (C18:2) and linolenic acid(C18:3), and showed the lower relative contents of palmitic acid (C16:0) and stearic acid(C18:0). In addition, this treatment inhibited the decrease in index of unsaturated fatty acidsand unsaturated degree of fatty acids. These results suggested that the enhancement of chilling tolerance and the alleviation of chilling injury in kiwifruit by exogenous Put treatment may bedue to the decrease of LOX activity, inhibiting decrease in relative content of unsaturated fattyacids of membrane lipids and maintaining higher unsaturated degree of fatty acids.
本论文研究不同采收成熟度、逐步降温和低温预贮（low temperature conditioning,LTC）两种物理降温方式以及外源腐胺（Put）化学处理对猕猴桃果实冷害发生的影响及其机制，为猕猴桃贮运技术体系的建立提供参考。研究结果分述如下：
1.研究了不同成熟度对‘徐香’猕猴桃果实冷害、品质、呼吸、乙烯、相对膜透性、丙二醛含量和抗氧化酶活性的影响。结果显示：（1）Ⅰ可溶性固形物（TSS）:4.5%～5.5%）、Ⅱ（TSS:6.5%～7.5%）和Ⅲ（TSS:8.0%～9.0%）3个不同采收成熟度的‘徐香’果实低温贮藏效果和冷害程度差异较大：采收Ⅰ果实失重率和腐烂率较高，较早表现木质化、水渍化等冷害症状，冷害率和冷害指数也较高，贮后货架期结束时硬度、可滴定酸、可溶性固性物和维生素C含量明显下降。而Ⅱ和Ⅲ两采收期出现木质化和水渍化等冷害症状较晚，冷害率和冷害指数较低，而且在贮后货架期结束时仍保持较高可溶性固形物和维生素C含量。与Ⅱ相比较，Ⅲ采收期果实的失重率和腐烂率较高，贮后货架期结束时果肉硬度、可滴定酸和维生素C含量较小，因此，‘徐香’果实适宜贮藏的采收期为Ⅱ。（2）猕猴桃果实发生冷害时呼吸速率、乙烯释放量和相对膜透性均异常增加。而Ⅱ采收期‘徐香’果实显著降低呼吸和乙烯释放，并保持较高的超氧化物歧化酶（SOD）、过氧化物酶（POD）、过氧化氢酶（CAT）和抗坏血酸过氧化物酶（APX）等抗氧化酶活性，抑制膜脂过氧化程度，从而减轻果实冷害的发生。这些结果说明Ⅱ采收成熟度果实冷害相对较低可能与保持较高的抗氧化酶活性有关。
2.首先研究了不同降温处理对‘徐香’果实冷害、失重率、腐烂率和品质的影响，发现逐步降温可显著降低‘徐香’猕猴桃冷害指数、冷害率、腐烂率和失重率，并保持较高的果肉硬度、可溶性固性物、可滴定酸和维生素C含量，其中逐步降温2（10℃→5℃2d→2℃2d→0±0.5℃）的效果最好。
在上述研究的基础上，选用逐步降温2探索逐步降温处理对减轻‘徐香’果实冷害的作用，通过研究逐步降温处理对‘徐香’果实呼吸、乙烯和活性氧代谢的影响，以揭示其减轻果实冷害的机理。结果显示：‘徐香’果实发生冷害时呼吸速率、乙烯释放量和相对膜透性均异常增加，SOD、CAT、POD、APX等酶活性降低，丙二醛（MDA）大量累积。逐步降温处理有效降低冷藏‘徐香’果实的冷害指数和冷害率，抑制呼吸速率和乙烯释放量的增加，保持较高的SOD、CAT、POD、APX酶活性，并降低O_2~(·—)和H_2O_2含量，显著减少膜脂过氧化产物MDA的积累，抑制相对膜透性的增加。这说明，逐步降温减轻‘徐香’果实冷害与活性氧清除酶活性的提高及活性氧积累的降低有关。
3.研究了不同LTC处理对‘海沃德’果实冷害、失重率、腐烂率和品质的影响，发现LTC可显著降低‘海沃德’冷害指数、冷害率、腐烂率和失重率，并保持较高的果肉硬度、可滴定酸、可溶性固性物和维生素C的含量，其中LTC5（12℃，3d）处理的效果最好。在此基础上，通过研究LTC5处理对‘海沃德’果实活性氧代谢、脂氧合酶（LOX）活性和內源激素的影响，以掲示LTC处理减轻‘海沃德’果实冷害的作用机制。结果表明，LTC处理保持较高的SOD、CAT、POD及APX抗氧化酶活性和较低的LOX酶活性，并降低O_2~(·—)生成速率和H_2O_2含量，抑制MDA和相对膜透性上升，同时积累较低GA_3，并保持较高ABA、IAA、ZR水平和ABA/IAA、ABA/GA_3比例，最终减轻‘海沃德’果实的冷害症状。这些结果说明，LTC处理可维持活性氧代谢平衡，积累较少的GA_3，并保持较高ABA、IAA、ZR水平和ABA/IAA、ABA/GA_3比例，从而减轻果实冷害的发生。
4.研究了不同浓度（1mmol·L~(-1)、2mmol·L~(-1)、4mmol·L~(-1)）Put处理对‘红阳’果实冷害、品质、腐烂率和失重率的影响，结果发现2mmol·L~(-1)Put处理可显著减少‘红阳’果肉木质化和水渍化等冷害症状的表现，减少果实失重率和腐烂率，并保持较高果肉硬度、可滴定酸和维生素C含量。在此基础上研究了2mmol·L~(-1)Put处理对‘红阳’呼吸、乙烯、活性氧代谢、ASA-GSH循环，內源多胺、LOX酶活性以及膜脂脂肪酸的影响，以掲示Put处理减轻‘红阳’果实冷害的机理。结果显示：
（1）猕猴桃果实的冷害与内源亚精胺（Spd）、精胺（Spm）含量的降低有关。内源Spd和Spm的积累水平与冷害的发生程度存在明显的负相关性，相关系数分别为R=–0.9255和R=–0.9133。而Put处理提高‘红阳’内源Put、Spd、Spm含量，说明外源Put诱导了內源多胺含量的增加，从而有助于减轻猕猴桃果实冷害。
（2）Put处理显著抑制了‘红阳’果实呼吸速率和乙烯释放速率，保持较高POD、SOD、CAT、APX、脱氢抗坏血酸还原酶（DHAR）、单脱氢抗坏血酸还原酶（MDHAR）、谷胱甘肽还原酶（GR）活性及还原型谷胱甘肽（GSH）、抗坏血酸（ASA）抗氧化物质含量，积累较少的氧化型谷胱甘肽（GSSG）和脱氢抗坏血酸（DHA），保持较高的GSH/GSSG和ASA/DHA比例，并抑制O_2~(·—)和H_2O_2积累，降低了果实中MDA含量和相对膜透性，减轻‘红阳’果实冷害。说明， Put处理提高‘红阳’果实抗冷性与保持较高的抗氧化物质含量和抗氧化酶活性以及降低膜脂过氧化程度密切相关。
（3）猕猴桃果实的冷害与LOX酶活性上升以及亚油酸、亚麻酸含量的降低密切相关。 Put处理抑制‘红阳’果实LOX活性上升，提高了亚油酸、亚麻酸不饱和脂肪酸相对含量，降低了棕榈酸和硬脂酸等饱和脂肪酸相对含量，抑制膜脂肪酸不饱和度和不饱和指数下降。说明，Put处理是通过降低‘红阳’果实LOX活性，抑制膜脂不饱和脂肪酸相对含量的下降，保持较高的膜脂肪酸不饱和度和不饱和指数，从而增强了‘红阳’果实抗冷性，减轻果实冷害的发生。
Kiwifruit as an important fruit is popular for its thin skin, juiceful texture and rich innutrient. The kiwifruit, as a typical kind of respiratory climactic fruit, undergoes rapidsoftening with severe decay after harvest, which shortens its shelf life. Cold storageeffectively inhibits fruit softening and extends the postharvest life of kiwifruits. However,kiwifruit is susceptible to chilling injury when protractedly stored at inaptitude temperatures.CI symptoms of kiwifruit are characterized by the appearance of concave spot in the skin,grainy tissue in the outer pericarp and the development of diffuse pitting in association with awater-soaked appearance. These symptoms become most severe after fruit is transferred toambient temperature. CI increase susceptibility to decay, which become a major obstacle topostharvest storage and transportation. At present, most study focus on storage of kiwifruit.Little information is available on the CI of kiwifruit. Thus, it is crucial to study themechanism and regulation of chilling injury in postharvest kiwifruit for storage andtransportation.
In this paper，it was investigated that the effect of different harvested maturity, gradualcooling, low temperature conditioning (LTC) and exogenous putrescine (Put) on chillinginjury in kiwifruit and the possible mechanisms. The results were as follows:
1. The effects of different harvested maturities on chilling injury, fruit quality，respirationand ethylene production, relative membrane permeability, malonaldehyde (MDA) content andthe activities of antioxidant enzymes in kiwifruit (Actinidia deliciosa cv.‘Xuxiang’) duringcold storage were investigated. The results showed as follows:
(1) The storage qualities and chilling injury of kiwifruit were much difference amongⅠ (TSS:4.5%～5.5%), Ⅱ(TSS:6.5%～7.5%), Ⅲ(TSS:8.0%～9.0%) in low temperaturestorage. Compared to fruits of Ⅱ and Ⅲ, the fruits of Ⅰmaturity showed early CI symptomscharacterized by flesh grainy and water-soaked, maintained higher weight loss ratio, decayrate, chilling injury index and chilling injury incidence. In addition, the fruits ofⅠ maturityshowed a decrease in firmness, total soluble solids (TSS), titratable acidity (TA) and VCat theend of storage and shelf life. However, the fruits of Ⅲmaturity maintained higher weight lossratio, decay rate, and lower firmness, TA and VCthan these of Ⅱ maturity. Thus，the best harvest maturity grade for kiwifruit storage performed well was Ⅱmaturity.
(2) The respiration rate, ethylene production and the relative membrane penetrabilityabnormally increased under chilling stress. However, the fruits of Ⅱm aturity maintainedlower respiration and ethylene production, induced the increase in the activities of superoxidedismutase (SOD), ascorbate peroxidase (APX), catalase (CAT) and peroxidase (POD),reduced lipid peroxidation and alleviated chilling injury. These results indicated thatⅡmaturity retard chilling injury in ‘Xuxiang’ fruit which may be related to enhancing theactivities of antioxidant enzymes.
2. The ‘Xuxiang’ Kiwifruit were treated by direct cooling and gradual cooling todeterminate the effect of cooling pattern on chilling injury, weight loss ratio, decay rate andfruit quality. The gradual cooling treatment significantly reduced chilling injury index,chilling injury incidence, weight loss ratio and decay rate, maintained higher firmness, TSS,VCand TA. It was found that treatment2(10℃→5℃2d→2℃2d→0±0.5℃) was mosteffective in ‘Xuxiang’ fruit.
On the basis of above results, the gradual cooling2in ‘Xuxiang’ fruit was chosen toinvestigate the effects of gradual cooling on chilling injury and the influence on active oxygenmetabolism. The results showed that the respiration rate, ethylene production, relativemembrane permeability and malonaldehyde (MDA) contents increased, while the activities ofCAT, SOD, POD and APX decreased during the development of CI symptom. However，thegradual cooling treatment significantly alleviated chilling injury, reduced respiration andethylene production, and inhibited the accumulation of MDA and the increase in relativemembrane permeability. Moreover，the gradual cooling treatment maintained higher activitiesof SOD, CAT, APX and POD than direct cooling fruit during the storage, and kept lowerlevels of superoxide anion production rate (O_2~(·—)) and H_2O_2content. These results indicatedthat gradual cooling treatment retard chilling injury in ‘Xuxiang’ fruit which may be related toenhancing the activities of active oxygen scavenging enzymes and reducing the accumulationof active oxygen.
3. The effects of LTC on chilling injury, weight loss ratio, decay rate and fruit quality inkiwifruit (Actinidia deliciosa cv.‘Hayward’) during cold storage were investigated. Thetreatment with LTC was more effective in reducing chilling injury index and chilling injuryincidence, while maintaining higher level of firmness, TSS, TA and VC. We found that LTC5(12℃，3d) treatment was the optimal treatment to alleviate chilling injury and maintainhigher quality of kiwifruit. Thus LTC5treatment was chosen to to test the effects of LTCtreatment on chilling injury and the possible mechanisms in kiwifruit. The results showed thatLTC treatment inhibited the accumulation of MDA and the increase in relative membrane permeability. Furthermore, LTC treatment decreased accumulation of H_2O_2and O_2~(·—)and theactivity in lipoxygenase (LOX), while increased the activities of CAT, SOD, POD and APX inkiwifruit. The treatment also maintained higher levels of the endogenous ABA, IAA and ZR,lower GA_3levels, and higher ratio of ABA/GA_3and ABA/IAA. These results indicated thatthe effect of LTC treatment on alleviating CI of kiwifruit may be due to its ability to improveantioxidant enzyme activities and reduce ROS accumulation, maintain higher levels of theendogenous ABA, IAA and ZR, lower GA_3levels, and higher ratios of ABA/GA_3andABA/IAA.
4. Kiwifruit (Actinidia chinensis cv.‘Hongyang’) were treated with differentconcentrations of Put (1mmol·L~(-1)，2mmol·L~(-1)，4mmol·L~(-1)) to investigate the effect onchilling injury, weight loss ratio, decay rate and fruit quality. We found that treatment with2mmol·L~(-1)Put treatment was most effective in reducing in CI symptoms of grainy andwater-soaked, decreasing weight loss ratio and decay rate and maintaining higher level offirmness, TA and VC. Thus，2mmol·L~(-1)Put treatment was chosen to investigate the possiblemechanisms of alleviating chilling injury. The results showed as follows:(1) Chilling injurywas related to the reduction of endogenesis Spd and Spm of kiwifruit. The accumulation levelof endogenesis Spm and Spd and chilling injury index appeared remarkable negativecorrelation, R=–0.9255and R=–0.9133. However, exogenous Put treatment increased thecontents of endogenesis Put、Spd and Spm. These results indicated that exogenous Puttreatment reducing in chilling injury may be related to improving endogenous polyaminecontent.
(2) Exogenous Put treatment alleviated chilling injury, inhibited respiration andethylene production, maintained higher level of GSH and ASA and lower level of GSSG andDHA, and kept higher ratio of GSH/GSSG and ASA/DHA. The activities of SOD、POD、CAT、APX、DHAR、MDHAR、GR were also induced by exogenous Put treatment. In addition,this treatment inhibited the accumulation of MDA, H_2O_2and O_2~(·—)and the increase inrelative membrane permeability. These results suggested that the effect of exogenous Puttreatment on alleviating chilling injury of kiwifruit may be attributed to its ability to enhanceantioxidant enzyme activities and the content of antioxidants and decrease lipid peroxidation.
(3) Chilling injury was related to increase of LOX activity and the decrease of linoleicacid (C18:2) and linolenic acid (C18:3). Exogenous Put treatment inhibited increased of LOXactivity, maintained higher relative contents of linoleic acid (C18:2) and linolenic acid(C18:3), and showed the lower relative contents of palmitic acid (C16:0) and stearic acid(C18:0). In addition, this treatment inhibited the decrease in index of unsaturated fatty acidsand unsaturated degree of fatty acids. These results suggested that the enhancement of chilling tolerance and the alleviation of chilling injury in kiwifruit by exogenous Put treatment may bedue to the decrease of LOX activity, inhibiting decrease in relative content of unsaturated fattyacids of membrane lipids and maintaining higher unsaturated degree of fatty acids.
引文
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