冷藏条件下白沙枇杷与红沙枇杷果皮活性氧代谢研究
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摘要
为探讨红沙枇杷与白沙枇杷冷藏耐储性差异的原因,为枇杷采后生理和保鲜技术研究提供参考,以白沙枇杷"白玉"和红沙枇杷"鸡蛋红"为材料,在6℃的冷藏条件下,测定了果皮中氧自由基产生速率(oxygenfree radical production rate, SPR)、超氧化物歧化酶(SOD)、过氧化物酶(POD)和过氧化氢酶(CAT)等保护酶活性,以及膜脂过氧化伤害产物(MDA)的含量等活性氧(reactive oxygen species, ROS)代谢相关指标的变化规律。表明,随着冷藏的进程,冷藏的前10 d,红沙枇杷果皮SPR高于白沙枇杷,10 d后红沙枇杷果皮SPR有所下降而白沙枇杷果皮则大幅上升,导致10 d后红沙枇杷果皮SPR低于白沙枇杷;冷藏后红沙枇杷果皮ROS相关酶(SOD, POD和CAT)活性高于白沙枇杷果皮,即其ROS清除能力高于白沙枇杷果皮;在冷藏5 d后白沙枇杷果皮膜脂过氧化伤害产物MDA含量持续上升,而红沙枇杷果皮则维持在较低水平并低于白沙枇杷果皮,说明白沙枇杷果皮膜脂过氧化程度较高。分析认为冷藏初期红沙枇杷果皮较高的氧自由基产生速率激活了ROS清除系统,导致红沙枇杷果皮具有更高的ROS清除能力;冷藏后期白沙枇杷果皮则出现了氧自由基和MDA的积累,暗示其膜脂过氧化的发生和果实内外环境的恶化。红沙枇杷和白沙枇杷果皮ROS代谢的差异与冷藏耐储性相关。
To reveal the factors causing the differences between white and red flesh loquat storability during cold storage, and to provide a reference for postharvest physiology and preservation technology of loquat fruits, the white flesh loquat(cv. Baiyu) and red flesh loquat(cv. Jidanhong) were used as materials, the ROS-related indices of superoxide production rate(SPR), the protective enzymes activity of Superoxide dismutase(SOD), peroxidase(POD) catalase(CAT) and the fruit pericarp reactive oxygen species(ROS) metabolism indexes between them determined. were determined under 6℃ cold storage. The results have showed that SPR before the first ten days of cold storage was higher in the white flesh loquat pericarp than that in the red one. While the red flesh loquat pericarp SPR declined slightly and the white one rose greatly, which resulted in the higher SPR in the red one than that in the white one after 10 d time point. Higher pericarp antioxidant enzyme activities of SOD, POD and CAT inthe red flesh loquat pericarp than in the white one were observed after 5 d time point in cold storage conditions,which means the higher ROS-scavenging capability of the red flesh loquat pericarp than the white one in them.MDA content was constantly increasing in the white flesh loquat pericarp while the content remained at a relatively low level in the red one after 5 d time point in cold storage conditions. This indicated a higher membrane lipid peroxidation degree in the white flesh loquat pericarp than in the red one. It was concluded that the high oxygen free radical production rate production in the red flesh loquat pericarp at the early stage of cold storage activated ROS scavenging system, and which led to higher ROS scavenging ability in the red flesh loquat pericarp than in the white one. The accumulation of oxygen free radical production and MDA content in the white flesh loquat pericarp in the later stage of cold storage indicated a high degree of membrane lipid peroxidation and the environmental deterioration of extra and intracellular environment of the fruits. It could conclude that the metabolism differences correlated with storability of white and red flesh loquat pericarp during cold storage.
To reveal the factors causing the differences between white and red flesh loquat storability during cold storage, and to provide a reference for postharvest physiology and preservation technology of loquat fruits, the white flesh loquat(cv. Baiyu) and red flesh loquat(cv. Jidanhong) were used as materials, the ROS-related indices of superoxide production rate(SPR), the protective enzymes activity of Superoxide dismutase(SOD), peroxidase(POD) catalase(CAT) and the fruit pericarp reactive oxygen species(ROS) metabolism indexes between them determined. were determined under 6℃ cold storage. The results have showed that SPR before the first ten days of cold storage was higher in the white flesh loquat pericarp than that in the red one. While the red flesh loquat pericarp SPR declined slightly and the white one rose greatly, which resulted in the higher SPR in the red one than that in the white one after 10 d time point. Higher pericarp antioxidant enzyme activities of SOD, POD and CAT inthe red flesh loquat pericarp than in the white one were observed after 5 d time point in cold storage conditions,which means the higher ROS-scavenging capability of the red flesh loquat pericarp than the white one in them.MDA content was constantly increasing in the white flesh loquat pericarp while the content remained at a relatively low level in the red one after 5 d time point in cold storage conditions. This indicated a higher membrane lipid peroxidation degree in the white flesh loquat pericarp than in the red one. It was concluded that the high oxygen free radical production rate production in the red flesh loquat pericarp at the early stage of cold storage activated ROS scavenging system, and which led to higher ROS scavenging ability in the red flesh loquat pericarp than in the white one. The accumulation of oxygen free radical production and MDA content in the white flesh loquat pericarp in the later stage of cold storage indicated a high degree of membrane lipid peroxidation and the environmental deterioration of extra and intracellular environment of the fruits. It could conclude that the metabolism differences correlated with storability of white and red flesh loquat pericarp during cold storage.
引文
Cai C.,Chen K.S.,Xu W.P.,Zhang W.S.,Li X.,and Ferguson I.,2006a,Effect of 1-MCP on postharvest quality of loquat fruit,Postharvest Biology and Technology,40(2):155-162
Cai C.,Li X.,and Chen K.S.,2006b,Acetylsalicylic acid alleviates chilling injury of postharvest loquat(Eriobotrya japonica Lindl.)fruit,European Food Research and Technology,223(4):533-539
Cai C.,Xu C.J.,Li X.,Ferguson I.,and Chen K.S.,2006c,Accumulation of lignin in relation to change in activities of lignification enzymes in loquat fruit flesh after harvest,Postharvest Biology and Technology,40(2):163-169
Cai X.Q.,Hu F.P.,Chen W.,Lin N.,and Lin J.Q.,2010,Efect of endophytic bacterium Bacillus subtilis on controlling litchi downy blight and its physiochemical mechanism,Redai Zuowu Xuebao(Chinese Journal of Tropical Crops),31(2):241-247(蔡学清,胡方平,陈炜,林娜,林建强,2010,内生枯草芽孢杆菌BS-2防治荔枝霜疫病及其生化机理,热带作物学报,31(2):241-247)
Cao S.F.,Zheng Y.H.,Wang K.T.,Jin P.,and Rui H.J.,2009,Methyl jasmonate reduces chilling injury and enhances antioxidant enzyme activity in postharvest loquat fruit,Food Chemistry,115(4):1458-1463
Cao S.F.,Zheng Y.H.,Wang K.T.,Rui H.J.,and Tang S.S.,2010,Effect of methyl jasmonate on cell wall modification of loquat fruit in relation to chilling injury after harvest,Food Chemistry,118(3):641-647
Chen F.H.,Zhang M.Z.,and Wu G.B.,2014,Study of lignification's delaying and its relationship with energy metabolism in loquat fruits after nitric oxide fumigation,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),47(12):2425-2434(陈发河,张美姿,吴光斌,2014,NO处理延缓采后枇杷果实木质化劣变及其与能量代谢的关系,中国农业科学,47(12):2425-2434)
Chen H.J.,Gao H.Y.,Mao J.L.,Chen W.X.,and Zhou Y.J.,2007,Changes in the storage characteristics and quality of loquat fruits for different cultivars under the condition of low temperature,Zhejiang Nongye Xuebao(Acta Agriculturae Zhejiangensis),19(4):280-283(陈杭君,郜海燕,毛金林,陈文烜,周拥军,2007,低温条件下不同品种枇杷果实的贮藏特性与品质变化,浙江农业学报,19(4):280-283)
Chen W.,Shi L.Y.,Su X.G.,and Yang Z.F.,2013,Research on postharvest quality in loquat fruit of different varieties during low temperature storage,Shipin Anquan Zhiliang Jiance Xuebao(Journal of Food Safety and Quality),4(6):1743-1749(陈伟,施丽愉,苏新国,杨震峰,2013,不同品种枇杷果实采后低温贮藏品质研究,食品安全质量检测学报,4(6):1743-1749)
Fang Q.,Qiao Y.J.,Wang H.H.,and Zhang S.L.,2006,Effect of storage Temperatuer on Baiyu loquat quality,Zhongguo Shipin Gongye(China Food Industry),21(2):46-48(方强,乔勇进,王海宏,张绍铃,2006,贮藏温度对白玉枇杷保鲜效果的影响,中国食品工业,21(2):46-48)
Fridovich I.,1975,Superoxide dismutases,Annual review of biochemistry,44(1):147-159
Gao J.F.,ed.,2006,Plant physiology experiment instruction,Higher Education Press,Beijing,China,pp.142-143,210-211,214-215(高俊凤,主编,2006,植物生理学实验指导,高等教育出版社,中国,北京,pp.142-143,210-211,214-215)
Jin W.W.,Xu C.J.,Li X.,Zhang B.,Wang P.,Allan A.C.,and Chen K.S.,2009,Expression of ROP/RAC GTPase genes in postharvest loquat fruit in association with senescence and cold regulated lignification.Postharvest Biology and Technology,54(1):9-14
Li W.L.,Li M.F.,He J.H.,Zhao X.Q.,Feng S.,Zhou H.L.,and Li S.P.,2015,Research progress on banana functional genes involved in cold resistance,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),13(5):1185-1192(李卫亮,李茂富,贺军虎,赵小青,冯顺,周海兰,李绍鹏,2015,香蕉抗寒相关功能基因研究进展,分子植物育种,13(5):1185-1192)
Lin J.C.,Lin H.T.,Zheng H.,Ye D.Y.,and Lin Y.F.,2014,Cell wall metabolism of pericarp and ultra-structure of fruit in relation to storability of harvested loquats,Redai Zuowu Xuebao(Chinese Journal of Tropical Crops),35(1):166-171(林建城,林河通,郑红,叶登云,林艺芬,2014,采后枇杷果皮细胞壁代谢和果实超微结构与贮藏性的关系,热带作物学报,35(1):166-171)
Lin S.Q.,2007,World loquat production and research with special reference to China,Acta Horticulturae,750:37-43
O'Brien J.A.,Daudi A.,Butt V.S.,and Bolwell G.P.,2012,Reactive oxygen species and their role in plant defence and cell wall metabolism,Planta,236(3):765-779
Pandhair V.,and Sekhon B.S.,2006,Reactive oxygen species and antioxidants in plants:an overview,Journal of Plant Biochemistry and Biotechnology,15(2):71-78
Pareek S.,Benkeblia N.,Janick J.,Cao S.F.,and Yahia E.M.,2014,Postharvest physiology and technology,of loquat(Eriobotrya japonica Lindl.)fruit,Journal of the Science of Food and Agriculture,94(8):1495-1504
Scandalios J.G.,1993,Oxygen stress and superoxide dismutases,Plant Physiology,101(1):7-12
Shao X.,and Tu K.,2014,Hot air treatment improved the chilling resistance of loquat fruit under cold storage,Journal of Food Processing and Preservation,38(2):694-703
Wang P.,Zhang B.,Li X.,Xu C.J.,Yin X.R.,Shan L.L.,Ferguson I.,and Chen K.S.,2010,Ethylene signal transduction elements involved in chilling injury in non-climacteric loquat fruit,Journal of Experimental Botany,61(1):179-190
Yang Q.Z.,Zhang Z.K.,Rao J.P.,Wang Y.P.,Sun Z.Y.,Ma Q.S.,and Dong X.Q.,2013,Low-temperature conditioning induces chilling tolerance in'Hayward'kiwifruit by enhancing antioxidant enzyme activity and regulating en-dogenous hormones levels,Journal of the Science of Food&Agriculture,93(15):3691-3699
Yang S.L.,Sun C.D.,Wang P.,Shan L.L.,Cai C.,Zhang B.,Zhang W.S.,Li X.,Ferguson I.,and Chen K.S.,2008,Expression of expansion genes during postharvest lignification and softening of'Luoyangqing'and'Baisha'loquat fruit under different storage conditions,Postharvest Biology and Technology,49(1):46-53
Zhang M.Z.,Wu G.B.,and Chen F.H.,2014,Effect of nitric oxide fumigation on lignification of loquat fruits during cold storage,Shipin Kexue(Food Science),35(16):232-237(张美姿,吴光斌,陈发河,2014,NO熏蒸处理对冷藏枇杷果实木质化劣变的影响,食品科学,35(16):232-237)
Zhang Z.L.,Qu W.J.,and Li X.F.,eds.,2009,Plant physiology experiment instruction,4ed editon,Higher Education Press,Beijing,China,pp.100,219-220,223-224(张志良,瞿伟菁,李小方,主编,2009,植物生理学实验指导,第4版,高等教育出版社,中国,北京,pp.100,219-220,223-224)
Zheng Y.H.,Su X.G.,Li S.Y.,and Xi Y.F.,2000,Quality,ac tive oxygen and polyamines metabolic changes in cold-stored loquat fruits as affected by postharvest SO2treatment,Zhiwu Shengli Xuebao(Acta Phytophysiologica Sinica),26(5):397-401(郑永华,苏新国,李三玉,席玙芳,2000,SO2对冷藏枇杷果实品质及活性氧和多胺代谢的影响,植物生理学报,26(5):397-401)
Cai C.,Li X.,and Chen K.S.,2006b,Acetylsalicylic acid alleviates chilling injury of postharvest loquat(Eriobotrya japonica Lindl.)fruit,European Food Research and Technology,223(4):533-539
Cai C.,Xu C.J.,Li X.,Ferguson I.,and Chen K.S.,2006c,Accumulation of lignin in relation to change in activities of lignification enzymes in loquat fruit flesh after harvest,Postharvest Biology and Technology,40(2):163-169
Cai X.Q.,Hu F.P.,Chen W.,Lin N.,and Lin J.Q.,2010,Efect of endophytic bacterium Bacillus subtilis on controlling litchi downy blight and its physiochemical mechanism,Redai Zuowu Xuebao(Chinese Journal of Tropical Crops),31(2):241-247(蔡学清,胡方平,陈炜,林娜,林建强,2010,内生枯草芽孢杆菌BS-2防治荔枝霜疫病及其生化机理,热带作物学报,31(2):241-247)
Cao S.F.,Zheng Y.H.,Wang K.T.,Jin P.,and Rui H.J.,2009,Methyl jasmonate reduces chilling injury and enhances antioxidant enzyme activity in postharvest loquat fruit,Food Chemistry,115(4):1458-1463
Cao S.F.,Zheng Y.H.,Wang K.T.,Rui H.J.,and Tang S.S.,2010,Effect of methyl jasmonate on cell wall modification of loquat fruit in relation to chilling injury after harvest,Food Chemistry,118(3):641-647
Chen F.H.,Zhang M.Z.,and Wu G.B.,2014,Study of lignification's delaying and its relationship with energy metabolism in loquat fruits after nitric oxide fumigation,Zhongguo Nongye Kexue(Scientia Agricultura Sinica),47(12):2425-2434(陈发河,张美姿,吴光斌,2014,NO处理延缓采后枇杷果实木质化劣变及其与能量代谢的关系,中国农业科学,47(12):2425-2434)
Chen H.J.,Gao H.Y.,Mao J.L.,Chen W.X.,and Zhou Y.J.,2007,Changes in the storage characteristics and quality of loquat fruits for different cultivars under the condition of low temperature,Zhejiang Nongye Xuebao(Acta Agriculturae Zhejiangensis),19(4):280-283(陈杭君,郜海燕,毛金林,陈文烜,周拥军,2007,低温条件下不同品种枇杷果实的贮藏特性与品质变化,浙江农业学报,19(4):280-283)
Chen W.,Shi L.Y.,Su X.G.,and Yang Z.F.,2013,Research on postharvest quality in loquat fruit of different varieties during low temperature storage,Shipin Anquan Zhiliang Jiance Xuebao(Journal of Food Safety and Quality),4(6):1743-1749(陈伟,施丽愉,苏新国,杨震峰,2013,不同品种枇杷果实采后低温贮藏品质研究,食品安全质量检测学报,4(6):1743-1749)
Fang Q.,Qiao Y.J.,Wang H.H.,and Zhang S.L.,2006,Effect of storage Temperatuer on Baiyu loquat quality,Zhongguo Shipin Gongye(China Food Industry),21(2):46-48(方强,乔勇进,王海宏,张绍铃,2006,贮藏温度对白玉枇杷保鲜效果的影响,中国食品工业,21(2):46-48)
Fridovich I.,1975,Superoxide dismutases,Annual review of biochemistry,44(1):147-159
Gao J.F.,ed.,2006,Plant physiology experiment instruction,Higher Education Press,Beijing,China,pp.142-143,210-211,214-215(高俊凤,主编,2006,植物生理学实验指导,高等教育出版社,中国,北京,pp.142-143,210-211,214-215)
Jin W.W.,Xu C.J.,Li X.,Zhang B.,Wang P.,Allan A.C.,and Chen K.S.,2009,Expression of ROP/RAC GTPase genes in postharvest loquat fruit in association with senescence and cold regulated lignification.Postharvest Biology and Technology,54(1):9-14
Li W.L.,Li M.F.,He J.H.,Zhao X.Q.,Feng S.,Zhou H.L.,and Li S.P.,2015,Research progress on banana functional genes involved in cold resistance,Fenzi Zhiwu Yuzhong(Molecular Plant Breeding),13(5):1185-1192(李卫亮,李茂富,贺军虎,赵小青,冯顺,周海兰,李绍鹏,2015,香蕉抗寒相关功能基因研究进展,分子植物育种,13(5):1185-1192)
Lin J.C.,Lin H.T.,Zheng H.,Ye D.Y.,and Lin Y.F.,2014,Cell wall metabolism of pericarp and ultra-structure of fruit in relation to storability of harvested loquats,Redai Zuowu Xuebao(Chinese Journal of Tropical Crops),35(1):166-171(林建城,林河通,郑红,叶登云,林艺芬,2014,采后枇杷果皮细胞壁代谢和果实超微结构与贮藏性的关系,热带作物学报,35(1):166-171)
Lin S.Q.,2007,World loquat production and research with special reference to China,Acta Horticulturae,750:37-43
O'Brien J.A.,Daudi A.,Butt V.S.,and Bolwell G.P.,2012,Reactive oxygen species and their role in plant defence and cell wall metabolism,Planta,236(3):765-779
Pandhair V.,and Sekhon B.S.,2006,Reactive oxygen species and antioxidants in plants:an overview,Journal of Plant Biochemistry and Biotechnology,15(2):71-78
Pareek S.,Benkeblia N.,Janick J.,Cao S.F.,and Yahia E.M.,2014,Postharvest physiology and technology,of loquat(Eriobotrya japonica Lindl.)fruit,Journal of the Science of Food and Agriculture,94(8):1495-1504
Scandalios J.G.,1993,Oxygen stress and superoxide dismutases,Plant Physiology,101(1):7-12
Shao X.,and Tu K.,2014,Hot air treatment improved the chilling resistance of loquat fruit under cold storage,Journal of Food Processing and Preservation,38(2):694-703
Wang P.,Zhang B.,Li X.,Xu C.J.,Yin X.R.,Shan L.L.,Ferguson I.,and Chen K.S.,2010,Ethylene signal transduction elements involved in chilling injury in non-climacteric loquat fruit,Journal of Experimental Botany,61(1):179-190
Yang Q.Z.,Zhang Z.K.,Rao J.P.,Wang Y.P.,Sun Z.Y.,Ma Q.S.,and Dong X.Q.,2013,Low-temperature conditioning induces chilling tolerance in'Hayward'kiwifruit by enhancing antioxidant enzyme activity and regulating en-dogenous hormones levels,Journal of the Science of Food&Agriculture,93(15):3691-3699
Yang S.L.,Sun C.D.,Wang P.,Shan L.L.,Cai C.,Zhang B.,Zhang W.S.,Li X.,Ferguson I.,and Chen K.S.,2008,Expression of expansion genes during postharvest lignification and softening of'Luoyangqing'and'Baisha'loquat fruit under different storage conditions,Postharvest Biology and Technology,49(1):46-53
Zhang M.Z.,Wu G.B.,and Chen F.H.,2014,Effect of nitric oxide fumigation on lignification of loquat fruits during cold storage,Shipin Kexue(Food Science),35(16):232-237(张美姿,吴光斌,陈发河,2014,NO熏蒸处理对冷藏枇杷果实木质化劣变的影响,食品科学,35(16):232-237)
Zhang Z.L.,Qu W.J.,and Li X.F.,eds.,2009,Plant physiology experiment instruction,4ed editon,Higher Education Press,Beijing,China,pp.100,219-220,223-224(张志良,瞿伟菁,李小方,主编,2009,植物生理学实验指导,第4版,高等教育出版社,中国,北京,pp.100,219-220,223-224)
Zheng Y.H.,Su X.G.,Li S.Y.,and Xi Y.F.,2000,Quality,ac tive oxygen and polyamines metabolic changes in cold-stored loquat fruits as affected by postharvest SO2treatment,Zhiwu Shengli Xuebao(Acta Phytophysiologica Sinica),26(5):397-401(郑永华,苏新国,李三玉,席玙芳,2000,SO2对冷藏枇杷果实品质及活性氧和多胺代谢的影响,植物生理学报,26(5):397-401)