蜜橘乙烯利处理后果皮不同部位着色差异的研究
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摘要
以无核‘南丰蜜橘’为试材,对其采后乙烯利褪绿过程中果皮不同部位着色差异现象进行研究。结果表明,褪绿过程中,果蒂部位着色快于赤道部位,该现象可以归因于这两个部位果皮叶绿素含量和类胡萝卜素含量的差异。进一步分析表明,乙烯利处理后,果蒂部位叶绿素a和叶绿素b降解快于赤道部位;果蒂部位橙色色素物质(以β–隐黄质为主)的积累高于赤道部位,而赤道部位黄色色素(以叶黄素为主)的积累高于果蒂部位,导致果蒂部位橙色深于赤道部位。褪绿关键期(贮藏3d时),果蒂部位叶绿素a/b相互转化相关基因CitNYC、叶绿素降解关键基因CitChlase、类胡萝卜素生物合成调控基因CitPDS、类胡萝卜素合成途径下游基因CitCHYB、CitNCED1和CitNCED2的表达水平均显著高于赤道部位(P <0.05)。这些基因表达水平的差异可以在一定程度上解释乙烯利处理后蜜橘果蒂和赤道两个部位果皮色素的变化。
The gradient coloration of ethylene-degreened mandarin fruit was studied. The results indicated that the coloration process of stem end position of the peel was faster than the equator position during the degreening process. The difference of chlorophyll content and carotenoid content of these two positions could well explain this phenomenon. Further studies indicated that the degradation of chlorophyll a and chlorophyll b in the stem end position of ethylene-degreened mandarin fruit was faster than that in the equator position,more orange-colored carotenoids(mainly β-cryptoxanthin)were synthesized in the stem end position of ethylene-degreened mandarin fruit,while more yellow-colored carotenoids(mainly lutein)were synthesized in the equator position. The different expression levels of some regulatory genes in chlorophyll and carotenoid metabolism(including CitChlase,CitNYC,CitPDS,CitCHYB,CitNCED1 and CitNCED2)could explain the pigment changes of these positions to some extent.
The gradient coloration of ethylene-degreened mandarin fruit was studied. The results indicated that the coloration process of stem end position of the peel was faster than the equator position during the degreening process. The difference of chlorophyll content and carotenoid content of these two positions could well explain this phenomenon. Further studies indicated that the degradation of chlorophyll a and chlorophyll b in the stem end position of ethylene-degreened mandarin fruit was faster than that in the equator position,more orange-colored carotenoids(mainly β-cryptoxanthin)were synthesized in the stem end position of ethylene-degreened mandarin fruit,while more yellow-colored carotenoids(mainly lutein)were synthesized in the equator position. The different expression levels of some regulatory genes in chlorophyll and carotenoid metabolism(including CitChlase,CitNYC,CitPDS,CitCHYB,CitNCED1 and CitNCED2)could explain the pigment changes of these positions to some extent.
引文
Agócs A,Nagy V,SzabóZ,Márk L,Ohmacht R,Deli J.2007.Comparative study on the carotenoid composition of the peel and the pulp of different citrus species.Innovative Food Science&Emerging Technologies,8(3):390-394.
Bernardi J,Licciardello C,Russo M P,Luisa C M,Carletti G,Recupero G R,Marocco A.2010.Use of a custom array to study differentially expressed genes during blood orange(Citrus sinensis L.Osbeck)ripening.Journal of Plant Physiology,167(4):301-310.
Carmona L,Alquézar B,Marques V V,Pe?a L.2017.Anthocyanin biosynthesis and accumulation in blood oranges during postharvest storage at different low temperatures.Food Chemistry,237:7-14.
CrifòT,Petrone G,Cicero L L,Piero A R L.2012.Short cold storage enhances the anthocyanin contents and level of transcripts related to their biosynthesis in blood oranges.Journal of Agricultural and Food Chemistry,60(1):476-481.
CrifòT,Puglisi I,Petrone G,Recupero G R,Lo Piero A R.2011.Expression analysis in response to low temperature stress in blood oranges:implication of the flavonoid biosynthetic pathway.Gene,476(1):1-9.
Deng L L,Yuan Z Y,Xie J,Yao S X,Zeng K F.2017.Sensitivity to ethephon degreening treatment is altered by blue led light irradiation in mandarin fruit.Journal of Agricultural and Food Chemistry,65(30):6158-6168.
Eaks I L.1970.Respiratory response,ethylene production,and response to ethylene of citrus fruit during ontogeny.Plant Physiology,45(3):334-338.
Goldschmidt E E,Huberman M,Goren R.1993.Probing the role of endogenous ethylene in the degreening of citrus fruit with ethylene antagonists.Plant Growth Regulation,12(3):325-329.
Kato M,Matsumoto H,Ikoma Y,Kuniga T,Nakajima N,Yoshida T,Yano M.2007.Accumulation of carotenoids and expression of carotenoid biosynthetic genes and carotenoid cleavage dioxygenase genes during fruit maturation in the juice sacs of‘Tamami’,‘Kiyom’tangor,and‘Wilking’mandarin.Journal of the Japanese Society for Horticultural Science,76(2):103-111.
Katz E,Lagunes P M,Riov J,Weiss D,Goldschmidt E E.2004.Molecular and physiological evidence suggests the existence of a system II-like pathway of ethylene production in non-climacteric Citrus fruit.Planta,219(2):243-252.
Liu Xiao-zhong,Long Qin-xian,Xu Sheng-ji,Lai Yi.1984.A study on the degeeening of Valencia orange by benzoic acid,cinnamic acid and diethylaminoethanal.Journal of Southwest Agricultural College,4:71-80.(in Chinese)刘孝仲,龙钦贤,许生吉,赖毅.1984.苯甲酸,桂皮酸,二乙氨基乙醇对伏令夏橙退绿的试验.西南农学院学报,4:71-80.
Ma G,Zhang L,Kato M,Yamawaki K,Kiriiwa Y,Yahata M,Ikoma Y,Matsumoto H.2015.Effect of the combination of ethylene and red LEDlight irradiation on carotenoid accumulation and carotenogenic gene expression in the flavedo of citrus fruit.Postharvest Biology and Technology,99:99-104.
Matsumoto H,Ikoma Y,Kato M,Nakajima N,Hasegawa Y.2009.Effect of postharvest temperature and ethylene on carotenoid accumulation in the flavedo and juice sacs of Satsuma mandarin(Citrus unshiu Marc.)fruit.Journal of Agricultural and Food Chemistry,57(11):4724-4732.
Meng Xiang-chun,Gao Zi-xiang,Zhang Zhao-qi,Zhang Ai-yu.2011.Carotenoid accumulation and its regulation by ethylene in fruits of Valencia orange during its late development and re-greening stages.Scientia Agricultura Sinica,44(3):538-544.(in Chinese)孟祥春,高子祥,张昭其,张爱玉.2011.夏橙果实发育后期及返青期类胡萝卜素积累及乙烯的调控.中国农业科学,44(3):538-544.
Peng Gang.2013.Chlorophyll and carotenoid metabolism based fruit color-break in Citrus[Ph.D.Dissertation].Hangzhou:Zhejiang University.(in Chinese)彭刚.2013.柑橘果实转色的叶绿素和类胡萝卜素代谢基础[博士论文].杭州:浙江大学.
Rodrigo M J,Zacarias L.2007.Effect of postharvest ethylene treatment on carotenoid accumulation and the expression of carotenoid biosynthetic genes in the flavedo of orange(Citrus sinensis L.Osbeck)fruit.Postharvest Biology and Technology,43(1):14-22.
Shemer T A,Harpaz-Saad S,Belausov E,Lovat N,Krokhin O,Spicer V,Eyal Y.2008.Citrus chlorophyllase dynamics at ethylene-induced fruit color-break:a study of chlorophyllase expression,posttranslational processing kinetics,and in situ intracellular localization.Plant Physiology,148(1):108-118.
Shimokawa K,Sakanoshita A,Horiba K.1978.Ethylene-induced changes of chloroplast structure in Satsuma mandarin(Citrus unshiu Marc.).Plant and Cell Physiology,19:229-236.
Tian Shi-ping,Lei Fu-quan.1989.The application of ethephon on Valencia orange.Science and Technology of Sichuan Agriculture,(1):19.(in Chinese)田世平,雷复权.1989.乙烯利在夏橙果实上的应用.四川农业科技,(1):19.
Yamauchi N,Akiyama Y,Kako S,Hashinaga F.1997.Chlorophyll degradation in Wase satsuma mandarin(Citrus unshiu Marc.)fruit with on-tree maturation and ethylene treatment.Scientia Horticulturae,71:35-42.
Yuan Z Y,Deng L L,Yin B F,Yao S X,Zeng K F.2017.Effects of blue LED light irradiation on pigment metabolism of ethephon-degreened mandarin fruit.Postharvest Biology and Technology,134:45-54.
Yun Z,Jin S,Ding Y D,Wang Z,Gao H J,Pan Z Y,Xu J,Cheng Y J,Deng X X.2012.Comparative transcriptomics and proteomics analysis of citrus fruit,to improve understanding of the effect of low temperature on maintaining fruit quality during lengthy post-harvest storage.Journal of Experimental Botany,2012,63(8):2873-2893.
Zhang Y J,Wang X J,Wu J X,Chen S Y,Chen H,Chai L J,Li H L.2014.Comparative transcriptome analyses between a spontaneous late-ripening sweet orange mutant and its wild type suggest the functions of ABA,sucrose and JA during citrus fruit ripening.PLoS ONE,9(12):e116056.
Zhou J Y,Sun C D,Zhang L L,Dai X,Xu C J,Chen K S.2010.Preferential accumulation of orange-colored carotenoids in ponkan(Citrus reticulata)fruit peel following postharvest application of ethylene or ethephon.Scientia Horticulturae,126(2):229-235.
Bernardi J,Licciardello C,Russo M P,Luisa C M,Carletti G,Recupero G R,Marocco A.2010.Use of a custom array to study differentially expressed genes during blood orange(Citrus sinensis L.Osbeck)ripening.Journal of Plant Physiology,167(4):301-310.
Carmona L,Alquézar B,Marques V V,Pe?a L.2017.Anthocyanin biosynthesis and accumulation in blood oranges during postharvest storage at different low temperatures.Food Chemistry,237:7-14.
CrifòT,Petrone G,Cicero L L,Piero A R L.2012.Short cold storage enhances the anthocyanin contents and level of transcripts related to their biosynthesis in blood oranges.Journal of Agricultural and Food Chemistry,60(1):476-481.
CrifòT,Puglisi I,Petrone G,Recupero G R,Lo Piero A R.2011.Expression analysis in response to low temperature stress in blood oranges:implication of the flavonoid biosynthetic pathway.Gene,476(1):1-9.
Deng L L,Yuan Z Y,Xie J,Yao S X,Zeng K F.2017.Sensitivity to ethephon degreening treatment is altered by blue led light irradiation in mandarin fruit.Journal of Agricultural and Food Chemistry,65(30):6158-6168.
Eaks I L.1970.Respiratory response,ethylene production,and response to ethylene of citrus fruit during ontogeny.Plant Physiology,45(3):334-338.
Goldschmidt E E,Huberman M,Goren R.1993.Probing the role of endogenous ethylene in the degreening of citrus fruit with ethylene antagonists.Plant Growth Regulation,12(3):325-329.
Kato M,Matsumoto H,Ikoma Y,Kuniga T,Nakajima N,Yoshida T,Yano M.2007.Accumulation of carotenoids and expression of carotenoid biosynthetic genes and carotenoid cleavage dioxygenase genes during fruit maturation in the juice sacs of‘Tamami’,‘Kiyom’tangor,and‘Wilking’mandarin.Journal of the Japanese Society for Horticultural Science,76(2):103-111.
Katz E,Lagunes P M,Riov J,Weiss D,Goldschmidt E E.2004.Molecular and physiological evidence suggests the existence of a system II-like pathway of ethylene production in non-climacteric Citrus fruit.Planta,219(2):243-252.
Liu Xiao-zhong,Long Qin-xian,Xu Sheng-ji,Lai Yi.1984.A study on the degeeening of Valencia orange by benzoic acid,cinnamic acid and diethylaminoethanal.Journal of Southwest Agricultural College,4:71-80.(in Chinese)刘孝仲,龙钦贤,许生吉,赖毅.1984.苯甲酸,桂皮酸,二乙氨基乙醇对伏令夏橙退绿的试验.西南农学院学报,4:71-80.
Ma G,Zhang L,Kato M,Yamawaki K,Kiriiwa Y,Yahata M,Ikoma Y,Matsumoto H.2015.Effect of the combination of ethylene and red LEDlight irradiation on carotenoid accumulation and carotenogenic gene expression in the flavedo of citrus fruit.Postharvest Biology and Technology,99:99-104.
Matsumoto H,Ikoma Y,Kato M,Nakajima N,Hasegawa Y.2009.Effect of postharvest temperature and ethylene on carotenoid accumulation in the flavedo and juice sacs of Satsuma mandarin(Citrus unshiu Marc.)fruit.Journal of Agricultural and Food Chemistry,57(11):4724-4732.
Meng Xiang-chun,Gao Zi-xiang,Zhang Zhao-qi,Zhang Ai-yu.2011.Carotenoid accumulation and its regulation by ethylene in fruits of Valencia orange during its late development and re-greening stages.Scientia Agricultura Sinica,44(3):538-544.(in Chinese)孟祥春,高子祥,张昭其,张爱玉.2011.夏橙果实发育后期及返青期类胡萝卜素积累及乙烯的调控.中国农业科学,44(3):538-544.
Peng Gang.2013.Chlorophyll and carotenoid metabolism based fruit color-break in Citrus[Ph.D.Dissertation].Hangzhou:Zhejiang University.(in Chinese)彭刚.2013.柑橘果实转色的叶绿素和类胡萝卜素代谢基础[博士论文].杭州:浙江大学.
Rodrigo M J,Zacarias L.2007.Effect of postharvest ethylene treatment on carotenoid accumulation and the expression of carotenoid biosynthetic genes in the flavedo of orange(Citrus sinensis L.Osbeck)fruit.Postharvest Biology and Technology,43(1):14-22.
Shemer T A,Harpaz-Saad S,Belausov E,Lovat N,Krokhin O,Spicer V,Eyal Y.2008.Citrus chlorophyllase dynamics at ethylene-induced fruit color-break:a study of chlorophyllase expression,posttranslational processing kinetics,and in situ intracellular localization.Plant Physiology,148(1):108-118.
Shimokawa K,Sakanoshita A,Horiba K.1978.Ethylene-induced changes of chloroplast structure in Satsuma mandarin(Citrus unshiu Marc.).Plant and Cell Physiology,19:229-236.
Tian Shi-ping,Lei Fu-quan.1989.The application of ethephon on Valencia orange.Science and Technology of Sichuan Agriculture,(1):19.(in Chinese)田世平,雷复权.1989.乙烯利在夏橙果实上的应用.四川农业科技,(1):19.
Yamauchi N,Akiyama Y,Kako S,Hashinaga F.1997.Chlorophyll degradation in Wase satsuma mandarin(Citrus unshiu Marc.)fruit with on-tree maturation and ethylene treatment.Scientia Horticulturae,71:35-42.
Yuan Z Y,Deng L L,Yin B F,Yao S X,Zeng K F.2017.Effects of blue LED light irradiation on pigment metabolism of ethephon-degreened mandarin fruit.Postharvest Biology and Technology,134:45-54.
Yun Z,Jin S,Ding Y D,Wang Z,Gao H J,Pan Z Y,Xu J,Cheng Y J,Deng X X.2012.Comparative transcriptomics and proteomics analysis of citrus fruit,to improve understanding of the effect of low temperature on maintaining fruit quality during lengthy post-harvest storage.Journal of Experimental Botany,2012,63(8):2873-2893.
Zhang Y J,Wang X J,Wu J X,Chen S Y,Chen H,Chai L J,Li H L.2014.Comparative transcriptome analyses between a spontaneous late-ripening sweet orange mutant and its wild type suggest the functions of ABA,sucrose and JA during citrus fruit ripening.PLoS ONE,9(12):e116056.
Zhou J Y,Sun C D,Zhang L L,Dai X,Xu C J,Chen K S.2010.Preferential accumulation of orange-colored carotenoids in ponkan(Citrus reticulata)fruit peel following postharvest application of ethylene or ethephon.Scientia Horticulturae,126(2):229-235.