细胞壁对果实质地影响机制的研究进展
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摘要
果实质地的改变严重影响采后果实的贮藏运输及其商品价值。而采后果实质地的改变通常认为是由于果实细胞初生壁和胞间层组织中的多糖降解,引起细胞壁超微结构改变所导致的。同时细胞壁多糖降解和多聚物化学键的改变,会进而引起细胞分离、软化、溶胀,改变细胞膨压,从而影响果实质地。多糖降解过程受到一系列细胞壁酶的协同作用,不同酶在果实质地变化的各阶段作用不同,而且各种酶活性变化在不同品种的果实中表现不同。本文主要对细胞壁多糖降解以及其相关酶对果实质地变化的影响进行了系统的分析与展望,总结了果实成熟软化中细胞壁的降解机理,为更好地进行果实贮藏、保鲜及加工提供理论依据。
Substantial changes have seriously affected the storage and transportation of postharvest fruits and their commercial value. Substantial changes in the consequences would be thought to be caused by the degradation of polysaccharides in the primary and intercellular layers of the fruit cells,which lead to the changes in the ultrastructure of the cell wall. At the same time,the degradation of cell wall polysaccharides and the changes of chemical bonds of the polymer would cause the cells to separate,soften and swell,and change the cell swell pressure,thereby effect the fruit texture. The degradation process of polysaccharides is synergistically affected by a series of cell wall enzymes. Different enzymes play different roles in various stages of fruit texture change,and the changes of various enzyme activities are different in fruits of different varieties. This paper systematically analyzed and prospects the degradation of cell wall polysaccharides and the effects of their related enzymes on fruit texture changes. The degradation mechanism of cell wall in fruit ripening and softening was summarized,which would provide a theoretical basis for better fruit storage,preservation and processing.
Substantial changes have seriously affected the storage and transportation of postharvest fruits and their commercial value. Substantial changes in the consequences would be thought to be caused by the degradation of polysaccharides in the primary and intercellular layers of the fruit cells,which lead to the changes in the ultrastructure of the cell wall. At the same time,the degradation of cell wall polysaccharides and the changes of chemical bonds of the polymer would cause the cells to separate,soften and swell,and change the cell swell pressure,thereby effect the fruit texture. The degradation process of polysaccharides is synergistically affected by a series of cell wall enzymes. Different enzymes play different roles in various stages of fruit texture change,and the changes of various enzyme activities are different in fruits of different varieties. This paper systematically analyzed and prospects the degradation of cell wall polysaccharides and the effects of their related enzymes on fruit texture changes. The degradation mechanism of cell wall in fruit ripening and softening was summarized,which would provide a theoretical basis for better fruit storage,preservation and processing.
引文
[1]Contador L,Shinya P,Infante R. Texture phenotyping in fresh fleshy fruit[J]. Scientia Horticulturae,2015,193:40-46.
[2]Dintwa E,Jancsók P,Mebatsion H K,et al. A finite element model for mechanical deformation of single tomato suspension cells[J]. Journal of Food Engineering,2011,103(3):265-272.
[3]Kendra M Blaker,James W. Olmstead. Cell wall composition of the skin and flesh tissue of crisp and standard texture southern highbush blueberry genotypes[J].Journal of Berry Research,2015,5(1):9-15.
[4]Liaotrakoon W,Buggenhout S V,Christiaens S,et al. An explorative study on the cell wall polysaccharides in the pulp and peel of dragon fruits(Hylocereus spp.)[J]. European Food Research & Technology,2013,237(3):341-351.
[5]Valérie Cornuault,Sara Pose,J Paul Knox. Extraction,texture analysis and polysaccharide epitope mapping data of sequential extracts of strawberry,apple,tomato and aubergine fruit parenchyma[J].Data in Brief,2018:314-320.
[6]王炜,李鹏霞,胡花丽,等.甘薯在贮藏期间细胞壁降解酶活性的变化[J].食品与发酵工业,2012,38(7):186-189.
[7]Nehanjali Parmar,Kunwar Harendra Singh,Deepika Sharma,et al. Genetic engineering strategies for biotic and abiotic stress tolerance and quality enhancement in horticultural crops:A comprehensive review[J]. Biotech,2017,7(4):239.
[8]姚佳. 超高压下莴笋质构的变化及机制研究[D].北京:中国农业大学,2014.
[9]Lahaye M,Bouin C,Barbacci A,et al. Water and cell wall contributions to apple mechanical properties[J]. Food Chemistry,2018,268:3 86-394.
[10]Hangjun Chena,Shifeng Caob,Xiangjun Fanga,et al.Changes in fruit firmness,cell wall composition and cell wall degrading enzymes in postharvest blueberries during storage[J]. Scientia Horticulturae,2015,188:44-48.
[11]阚娟,刘涛,金昌海,等.硬溶质型桃果实成熟过程中细胞壁多糖降解特性及其相关酶研究[J].食品科学,2011,32(4):268-274.
[12]Jovyn K T Ng,Roswitha Schrder,Paul W Sutherland,et al. Cell wall structures leading to cultivar differences in softening rates develop early during apple(Malus×domestica)fruit growth[J].BMC Plant Biology,2013,13(1):183.
[13]Contador L,Shinya P,Infante R. Texture phenotyping in fresh fleshy fruit[J]. Scientia Horticulturae,2015,193:40-46.
[14]Rachel A Burton,Michael J Gidley,Geoffrey B Fincher. Heterogeneity in the chemistry,structure and function of plant cell walls[J].Nature Chemical Biology.2010,6(10):724-732.
[15]Kong Y,O’Neill M,Zhou G. Plant cell walls:Isolation and monosaccharide composition analysis[J].Methods in Molecular Biology,2018:313-319.
[16]Jin CH,Suo B,Kan J,et al.Changes in cell wall polysaccharide of harvested peach fruit during storage[J].Journal of Plant Physiology and Molecular Biology,2006,32(6):657-664.
[17]Sila D N,Buggenhout S V,Duvetter T,et al. Pectins in processed fruits and vegetables:Part II—Structure-Function relationships[J]. Comprehensive Reviews in Food Science and Food Safety,2010,8(2):86-104.
[18]Dyson R J,Band L R,Jensen O E. A model of crosslink kinetics in the expanding plant cell wall:Yield stress and enzyme action[J].Journal of Theoretical Biology,2012,307:125-136.
[19]Bidhendi Amir J,Geitmann Anja. Relating the mechanics of the primary plant cell wall to morphogenesis[J].Journal of Experimental Botany,2016,67(2):449-461.
[20]张保才,周奕华.植物细胞壁形成机制的新进展[J].中国科学:生命科学,2015,45(6):544-556.
[21]Philip J Harris,Bronwen G Smith. Plant cell walls and cell-wall polysaccharides:Structures,properties and uses in food products[J].International Journal of Food Science and Technology,2006,41(2):129-143.
[22]Justyna Cybulska,Els Vanstreels,Quang Tri Ho,et al. Mechanical characteristics of artificial cell walls[J].Journal of Food Engineering,2010,96(2):287-294.
[23]Jin CH,Suo B,Kan J,et al. Changes in cell wall polysaccharide of harvested peach fruit during storage[J].Journal of Plant Physiology and Molecular Biology,2006,32(6):657-664.
[24]Dong Y,Zhang S,Wang Y. Compositional changes in cell wall polyuronides and enzyme activities associated with melting/mealy textural property during ripening following long-term storage of ‘Comice’ and ‘d’Anjou’ pears[J]. Postharvest Biology and Technology,2018,135:131-140.
[25]Xiao M,Yi J,Bi J,et al. Modification of cell wall polysaccharides during drying process affects texture properties of apple chips[J]. Journal of Food Quality,2018,2018(3):1-11.
[26]Njoroge D M,Kinyanjui P K,Chigwedere C M,et al. Mechanistic insight into common bean pectic polysaccharide changes during storage,soaking and thermal treatment in relation to the hard-to-cook defect[J]. Food Research International,2016,81:39-49.
[27]Christiaens Stefanie,Van Buggenhout,Sandy Houben,et al. Process-structure-function relations of pectin in food[J].Critical Reviews in Food Science and Nutrition,2016,56(6):1021-1042.
[28]Munarin F,Tanzi M C,P Petrini. Advances in biomedical applications of pectin gels[J].International Journal of Biological Macromolecules,2012,51(4):681-689.
[29]Mierczyńska J,Cybulska J,So?owiej B,et al. Effect of Ca2+,Fe2+ and Mg2+ on rheological properties of new food matrix made of modified cell wall polysaccharides from apple[J]. Carbohydrate Polymers,2015,133:547-555.
[30]Caffall Kerry Hosmer,Mohnen Debra. The structure,function,and biosynthesis of plant cell wall pectic polysaccharides[J].Carbohydrate Research,2009,344(14):1879-1900.
[31]Justyna Cybulska,Els Vanstreels,Quang Tri Ho,et al. Mechanical characteristics of artificial cell walls[J].Journal of Food Engineering,2010,96(2):287-294.
[32]Zykwinska Agata,Thibault Jean-Franois,Ralet Marie-Christine. Competitive binding of pectin and xyloglucan with primary cell wall cellulose[J].Carbohydrate Polymers,2008,74(4):957-961.
[33]Zhao Wenlin,Hussain Shehzad,Yan Shoulei,et al. Acetic acid pretreatment improves the hardness of cooked potato slices[J].Food Chemistry,2017,228:204-210.
[34]Cybulska J,Zdunek A,Psonkaantonczyk K M,et al. The relation of apple texture with cell wall nanostructure studied using an atomic force microscope[J]. Carbohydrate Polymers,2013,92(1):128-137.
[35]Cosgrove D J. Loosening of plant cell walls by expansins[J].Nature,2000,407(6802):321-326.
[36]David A Brummell,Mark H Harpster,Pamela Dunsmuir. Differential expression of expansin gene family members during growth and ripening of tomato fruit[J].Plant Molecular Biology,1999,39(1):161-169.
[37]Claudia M. Understanding the structure and function of bacterial expansins:A prerequisite towards practical applications for the bioenergy and agricultural industries[J]. Microbial Biotechnology,2016,9(6):727-736.
[38]王中凤.细胞壁分解酶与果实软化的关系研究进展[J]. 中国农学通报,2009,25(18):126-130.
[39]刘慧,陈复生,杨宏顺.影响桃果实质地的细胞壁降解酶的研究进展[J].食品与机械,2008(3):136-140.
[40]赵雯霖. 马铃薯及莲藕组织中多糖特性对其硬度影响机理研究[D].武汉:华中农业大学,2017.
[41]Rose J K,Saladié M,Catalá C.The plot thickens:New perspectives of primary cell wall modification[J].Current Opinion in Plant Biology,2004,7(3):296-301.
[42]陆胜民,金勇丰,张耀洲,等. 果实成熟过程中细胞壁组成的变化[J].植物生理学通讯,2001(3):246-249.
[43]Nitza L Díaz-Blanco,José R Rodríguez-Medina. Dosage rescue by UBC4 restores cell wall integrity in Saccharomyces cerevisiae lacking the myosin type II gene MYO1[J].Yeast,2007,24(4):343-355.
[44]Almeida Domingos P F,Huber Donald J. Polygalacturonase-mediated dissolution and depolymerization of pectins in solutions mimicking the pH and mineral composition of tomato fruit apoplast[J].Plant Science,2007,172(6):1087-1094.
[45]李春燕,张光伦,曾秀丽,等.细胞壁酶活性与甜橙果实质地的相关性研究[J].四川农业大学报,2006(1):73-76.
[46]Phan Thanh D,Wen Bo,West Gill,et al. Silencing of the major salt-dependent isoform of pectinesterase in tomato alters fruit softening(1)[J].Physiologia Plantarum,2007,144(4):1960-1967.
[47]Owino Willis Omondi,Nakano Ryohei,Kubo Yasutaka. Coordinated expression patterns of genes encoding cell wall modifying enzymes during ripening in distinct anatomical tissue regions of the fig(Ficus carica L.)fruit[J].Postharvest Biology and Technology,2004,32(3):253-261.
[48]Zhou Houcheng,Li Gang,Zhao Xia. Comparative analysis of pectate lyase in relation to softening in strawberry fruits[J]. Canadian Journal of Plant Science,2016,96(4):604-612.
[49]杨宏顺.果蔬气调冷藏下表皮和果胶超微结构与品质变化[D].上海:上海交通大学,2005.
[50]Monika Chylińska,Monika Szymańska-Chargot,Kamil Deryo,et al. Changing of biochemical parameters and cell wall polysaccharides distribution during physiological development of tomato fruit[J].Plant Physiology and Biochemistry,2017:328-337.
[51]David A Brummell,Mark H Harpster. Cell wall metabolism in fruit softening and quality and its manipulation in transgenicplants[J].Plant Molecular Biology,2001,47(1-2):311-339.
[52]Ng J K,Schrder R,Brummell D A,et al. Lower cell wall pectin solubilisation and galactose loss during early fruit development in apple(Malus×domestica)cultivar‘Scifresh’are associated with slower softening rate[J].Journal of Plant Physiology,2015:129-137.
[53]Pea M J,Carpita N C. Loss of highly branched arabinans and debranching of rhamnogalacturonan I accompany loss of firm texture and cell separation during prolonged storage of apple[J].Physiologia Plantarum,2004,135(3):1305-1313.
[54]张娟,高滋艺,杨惠娟,等.‘秦冠’和‘富士’质地差异的解剖学观察及相关酶活性研究[J].西北农业学报,2015,24(10):88-94.
[55]Jo C Dumville,Stephen C Fry. Solubilisation of tomato fruit pectins by ascorbate:A possible non-enzymic mechanism of fruit softening[J].Planta,2003,217(6):951-961.
[2]Dintwa E,Jancsók P,Mebatsion H K,et al. A finite element model for mechanical deformation of single tomato suspension cells[J]. Journal of Food Engineering,2011,103(3):265-272.
[3]Kendra M Blaker,James W. Olmstead. Cell wall composition of the skin and flesh tissue of crisp and standard texture southern highbush blueberry genotypes[J].Journal of Berry Research,2015,5(1):9-15.
[4]Liaotrakoon W,Buggenhout S V,Christiaens S,et al. An explorative study on the cell wall polysaccharides in the pulp and peel of dragon fruits(Hylocereus spp.)[J]. European Food Research & Technology,2013,237(3):341-351.
[5]Valérie Cornuault,Sara Pose,J Paul Knox. Extraction,texture analysis and polysaccharide epitope mapping data of sequential extracts of strawberry,apple,tomato and aubergine fruit parenchyma[J].Data in Brief,2018:314-320.
[6]王炜,李鹏霞,胡花丽,等.甘薯在贮藏期间细胞壁降解酶活性的变化[J].食品与发酵工业,2012,38(7):186-189.
[7]Nehanjali Parmar,Kunwar Harendra Singh,Deepika Sharma,et al. Genetic engineering strategies for biotic and abiotic stress tolerance and quality enhancement in horticultural crops:A comprehensive review[J]. Biotech,2017,7(4):239.
[8]姚佳. 超高压下莴笋质构的变化及机制研究[D].北京:中国农业大学,2014.
[9]Lahaye M,Bouin C,Barbacci A,et al. Water and cell wall contributions to apple mechanical properties[J]. Food Chemistry,2018,268:3 86-394.
[10]Hangjun Chena,Shifeng Caob,Xiangjun Fanga,et al.Changes in fruit firmness,cell wall composition and cell wall degrading enzymes in postharvest blueberries during storage[J]. Scientia Horticulturae,2015,188:44-48.
[11]阚娟,刘涛,金昌海,等.硬溶质型桃果实成熟过程中细胞壁多糖降解特性及其相关酶研究[J].食品科学,2011,32(4):268-274.
[12]Jovyn K T Ng,Roswitha Schrder,Paul W Sutherland,et al. Cell wall structures leading to cultivar differences in softening rates develop early during apple(Malus×domestica)fruit growth[J].BMC Plant Biology,2013,13(1):183.
[13]Contador L,Shinya P,Infante R. Texture phenotyping in fresh fleshy fruit[J]. Scientia Horticulturae,2015,193:40-46.
[14]Rachel A Burton,Michael J Gidley,Geoffrey B Fincher. Heterogeneity in the chemistry,structure and function of plant cell walls[J].Nature Chemical Biology.2010,6(10):724-732.
[15]Kong Y,O’Neill M,Zhou G. Plant cell walls:Isolation and monosaccharide composition analysis[J].Methods in Molecular Biology,2018:313-319.
[16]Jin CH,Suo B,Kan J,et al.Changes in cell wall polysaccharide of harvested peach fruit during storage[J].Journal of Plant Physiology and Molecular Biology,2006,32(6):657-664.
[17]Sila D N,Buggenhout S V,Duvetter T,et al. Pectins in processed fruits and vegetables:Part II—Structure-Function relationships[J]. Comprehensive Reviews in Food Science and Food Safety,2010,8(2):86-104.
[18]Dyson R J,Band L R,Jensen O E. A model of crosslink kinetics in the expanding plant cell wall:Yield stress and enzyme action[J].Journal of Theoretical Biology,2012,307:125-136.
[19]Bidhendi Amir J,Geitmann Anja. Relating the mechanics of the primary plant cell wall to morphogenesis[J].Journal of Experimental Botany,2016,67(2):449-461.
[20]张保才,周奕华.植物细胞壁形成机制的新进展[J].中国科学:生命科学,2015,45(6):544-556.
[21]Philip J Harris,Bronwen G Smith. Plant cell walls and cell-wall polysaccharides:Structures,properties and uses in food products[J].International Journal of Food Science and Technology,2006,41(2):129-143.
[22]Justyna Cybulska,Els Vanstreels,Quang Tri Ho,et al. Mechanical characteristics of artificial cell walls[J].Journal of Food Engineering,2010,96(2):287-294.
[23]Jin CH,Suo B,Kan J,et al. Changes in cell wall polysaccharide of harvested peach fruit during storage[J].Journal of Plant Physiology and Molecular Biology,2006,32(6):657-664.
[24]Dong Y,Zhang S,Wang Y. Compositional changes in cell wall polyuronides and enzyme activities associated with melting/mealy textural property during ripening following long-term storage of ‘Comice’ and ‘d’Anjou’ pears[J]. Postharvest Biology and Technology,2018,135:131-140.
[25]Xiao M,Yi J,Bi J,et al. Modification of cell wall polysaccharides during drying process affects texture properties of apple chips[J]. Journal of Food Quality,2018,2018(3):1-11.
[26]Njoroge D M,Kinyanjui P K,Chigwedere C M,et al. Mechanistic insight into common bean pectic polysaccharide changes during storage,soaking and thermal treatment in relation to the hard-to-cook defect[J]. Food Research International,2016,81:39-49.
[27]Christiaens Stefanie,Van Buggenhout,Sandy Houben,et al. Process-structure-function relations of pectin in food[J].Critical Reviews in Food Science and Nutrition,2016,56(6):1021-1042.
[28]Munarin F,Tanzi M C,P Petrini. Advances in biomedical applications of pectin gels[J].International Journal of Biological Macromolecules,2012,51(4):681-689.
[29]Mierczyńska J,Cybulska J,So?owiej B,et al. Effect of Ca2+,Fe2+ and Mg2+ on rheological properties of new food matrix made of modified cell wall polysaccharides from apple[J]. Carbohydrate Polymers,2015,133:547-555.
[30]Caffall Kerry Hosmer,Mohnen Debra. The structure,function,and biosynthesis of plant cell wall pectic polysaccharides[J].Carbohydrate Research,2009,344(14):1879-1900.
[31]Justyna Cybulska,Els Vanstreels,Quang Tri Ho,et al. Mechanical characteristics of artificial cell walls[J].Journal of Food Engineering,2010,96(2):287-294.
[32]Zykwinska Agata,Thibault Jean-Franois,Ralet Marie-Christine. Competitive binding of pectin and xyloglucan with primary cell wall cellulose[J].Carbohydrate Polymers,2008,74(4):957-961.
[33]Zhao Wenlin,Hussain Shehzad,Yan Shoulei,et al. Acetic acid pretreatment improves the hardness of cooked potato slices[J].Food Chemistry,2017,228:204-210.
[34]Cybulska J,Zdunek A,Psonkaantonczyk K M,et al. The relation of apple texture with cell wall nanostructure studied using an atomic force microscope[J]. Carbohydrate Polymers,2013,92(1):128-137.
[35]Cosgrove D J. Loosening of plant cell walls by expansins[J].Nature,2000,407(6802):321-326.
[36]David A Brummell,Mark H Harpster,Pamela Dunsmuir. Differential expression of expansin gene family members during growth and ripening of tomato fruit[J].Plant Molecular Biology,1999,39(1):161-169.
[37]Claudia M. Understanding the structure and function of bacterial expansins:A prerequisite towards practical applications for the bioenergy and agricultural industries[J]. Microbial Biotechnology,2016,9(6):727-736.
[38]王中凤.细胞壁分解酶与果实软化的关系研究进展[J]. 中国农学通报,2009,25(18):126-130.
[39]刘慧,陈复生,杨宏顺.影响桃果实质地的细胞壁降解酶的研究进展[J].食品与机械,2008(3):136-140.
[40]赵雯霖. 马铃薯及莲藕组织中多糖特性对其硬度影响机理研究[D].武汉:华中农业大学,2017.
[41]Rose J K,Saladié M,Catalá C.The plot thickens:New perspectives of primary cell wall modification[J].Current Opinion in Plant Biology,2004,7(3):296-301.
[42]陆胜民,金勇丰,张耀洲,等. 果实成熟过程中细胞壁组成的变化[J].植物生理学通讯,2001(3):246-249.
[43]Nitza L Díaz-Blanco,José R Rodríguez-Medina. Dosage rescue by UBC4 restores cell wall integrity in Saccharomyces cerevisiae lacking the myosin type II gene MYO1[J].Yeast,2007,24(4):343-355.
[44]Almeida Domingos P F,Huber Donald J. Polygalacturonase-mediated dissolution and depolymerization of pectins in solutions mimicking the pH and mineral composition of tomato fruit apoplast[J].Plant Science,2007,172(6):1087-1094.
[45]李春燕,张光伦,曾秀丽,等.细胞壁酶活性与甜橙果实质地的相关性研究[J].四川农业大学报,2006(1):73-76.
[46]Phan Thanh D,Wen Bo,West Gill,et al. Silencing of the major salt-dependent isoform of pectinesterase in tomato alters fruit softening(1)[J].Physiologia Plantarum,2007,144(4):1960-1967.
[47]Owino Willis Omondi,Nakano Ryohei,Kubo Yasutaka. Coordinated expression patterns of genes encoding cell wall modifying enzymes during ripening in distinct anatomical tissue regions of the fig(Ficus carica L.)fruit[J].Postharvest Biology and Technology,2004,32(3):253-261.
[48]Zhou Houcheng,Li Gang,Zhao Xia. Comparative analysis of pectate lyase in relation to softening in strawberry fruits[J]. Canadian Journal of Plant Science,2016,96(4):604-612.
[49]杨宏顺.果蔬气调冷藏下表皮和果胶超微结构与品质变化[D].上海:上海交通大学,2005.
[50]Monika Chylińska,Monika Szymańska-Chargot,Kamil Deryo,et al. Changing of biochemical parameters and cell wall polysaccharides distribution during physiological development of tomato fruit[J].Plant Physiology and Biochemistry,2017:328-337.
[51]David A Brummell,Mark H Harpster. Cell wall metabolism in fruit softening and quality and its manipulation in transgenicplants[J].Plant Molecular Biology,2001,47(1-2):311-339.
[52]Ng J K,Schrder R,Brummell D A,et al. Lower cell wall pectin solubilisation and galactose loss during early fruit development in apple(Malus×domestica)cultivar‘Scifresh’are associated with slower softening rate[J].Journal of Plant Physiology,2015:129-137.
[53]Pea M J,Carpita N C. Loss of highly branched arabinans and debranching of rhamnogalacturonan I accompany loss of firm texture and cell separation during prolonged storage of apple[J].Physiologia Plantarum,2004,135(3):1305-1313.
[54]张娟,高滋艺,杨惠娟,等.‘秦冠’和‘富士’质地差异的解剖学观察及相关酶活性研究[J].西北农业学报,2015,24(10):88-94.
[55]Jo C Dumville,Stephen C Fry. Solubilisation of tomato fruit pectins by ascorbate:A possible non-enzymic mechanism of fruit softening[J].Planta,2003,217(6):951-961.