不同硬度黑莓果实发育成熟进程中的解剖构造
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摘要
【目的】揭示黑莓果实发育成熟过程中与硬度相关的细胞形态结构特征。【方法】以生育期相似但硬度差异显著的黑莓品种‘Arapaho’和‘Boysen’为研究材料,采用石蜡切片法观察比较果实各发育阶段的细胞构造。【结果】两个品种果实开始着色时,外果皮和中果皮细胞排列整齐紧密,细胞结构清晰完整;果实着色至果面大部分变红过程中,‘Arapaho’中果皮细胞明显伸长变大,多呈卵圆形,细胞壁开始变薄但依然完整;果实完全变红过程中,两个品种果肉细胞均不断变大,至花后33 d局部果肉细胞的细胞壁裂解明显,细胞分界模糊;至果实完全变黑成熟时,‘Boysen’外果皮细胞及相邻一层中果皮细胞相对完整,而内部果肉细胞的细胞壁近乎完全裂解,‘Arapaho’果肉细胞细胞壁裂解程度明显低于‘Boysen’。【结论】两个不同硬度黑莓品种果实发育成熟前果肉细胞动态变化类似,成熟时中果皮细胞细胞壁裂解差异明显,与其果实硬度一致,表明黑莓成熟果实硬度主要由中果皮细胞的细胞壁裂解程度决定。
【Objective】This study aimed to reveal the characteristics of morphology and structure of blackberry fruit related to firmness during fruit development and ripening.【Methods】Soft-fruited( 'Boysen') and firm-fruited( 'Arapaho')blackberry cultivars which had similar growth periods but significant hardness differences as research materials were used to comparatively investigate the mesocarp cell structure characteristics of fruits at different developmental stages by paraffin section.【Result】At the initial phase of fruit coloring,the exocarp and mesocarp cells of the both cultivars were arranged neatly and tightly and the cell structures were clear and intact. During the stage of fruit coloring into majority red,pericarp cells of'Arapaho'fruits quickly elongated and became large as ovoid shapes,cell wall became thinner,but the cell morphology remained intact. During the process of fruit coloring into complete red,pulp cells of the both cultivars expanded continuously,cell walls of partial pulp cells dissociated obviously and the cell boundary exhibited ambiguity after 33 days after flowering. At the time of fruit ripening with full black,only exocarp cells and one layer of adjacent mesocarp cells were intact in'Boysen'fruits,and the other cells inside were almost completely lysed,the cell wall lysis extent in'Arapaho'fruit was really lower than that in 'Boysen'.【Conclusion】The changing dynamics of the pulp cells during fruit development in two blackberry cultivars with differential firmness were similar before ripening. At ripening stages,cell wall lysis degree of mesocarp cells of them demonstrated an obvious difference,which was coincided with the fruit firmness,implying a vital role of cell wall lysis degree of mesocarp cells in fruit firmness determination.
【Objective】This study aimed to reveal the characteristics of morphology and structure of blackberry fruit related to firmness during fruit development and ripening.【Methods】Soft-fruited( 'Boysen') and firm-fruited( 'Arapaho')blackberry cultivars which had similar growth periods but significant hardness differences as research materials were used to comparatively investigate the mesocarp cell structure characteristics of fruits at different developmental stages by paraffin section.【Result】At the initial phase of fruit coloring,the exocarp and mesocarp cells of the both cultivars were arranged neatly and tightly and the cell structures were clear and intact. During the stage of fruit coloring into majority red,pericarp cells of'Arapaho'fruits quickly elongated and became large as ovoid shapes,cell wall became thinner,but the cell morphology remained intact. During the process of fruit coloring into complete red,pulp cells of the both cultivars expanded continuously,cell walls of partial pulp cells dissociated obviously and the cell boundary exhibited ambiguity after 33 days after flowering. At the time of fruit ripening with full black,only exocarp cells and one layer of adjacent mesocarp cells were intact in'Boysen'fruits,and the other cells inside were almost completely lysed,the cell wall lysis extent in'Arapaho'fruit was really lower than that in 'Boysen'.【Conclusion】The changing dynamics of the pulp cells during fruit development in two blackberry cultivars with differential firmness were similar before ripening. At ripening stages,cell wall lysis degree of mesocarp cells of them demonstrated an obvious difference,which was coincided with the fruit firmness,implying a vital role of cell wall lysis degree of mesocarp cells in fruit firmness determination.
引文
[1]STRIK B C,CLARK J R,FINN C E,et al. Worldwide production of blackberries[J]. Acta Horticulturae,2008(777):209-218. DOI:10.17660/Acta Hortic.2008.777.31.
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[3]吴文龙,王小敏,赵慧芳,等.黑莓品种“Chester”鲜果贮藏性能的研究[J].食品科学,2010,31(8):280-284.WU W L,WANG X M,ZHAO H F,et al. Storage quality of fresh‘Chester’blackberry fruits[J]. Food Science,2010,31(8):280-284.
[4]赵慧芳,王小敏,吴文龙,等.黑莓品种‘Boysen’鲜果贮藏特性的研究[J].植物资源与环境学报,2010,19(3):28-36.ZHAO H F,WANG X M,WU W L,et al. Study on storage property of fresh fruit of blackberry(Rubus spp.)cultivar‘Boysen’[J]. Journal of Plant Resources and Environment,2010,19(3):28-36.
[5]HARDENBURG R E,WATADA A E,WANG C Y. The commercial storage of fruits, vegetables, and florist and nursery stocks[M]//USDA:Agriculture Handbook. Washington DC:USDA,1986.
[6]BOWER C. Postharvest handling,storage,and treatment of fresh market berries[J]. Food Science and Technology, 2007,168:261.
[7]JOO M J,LEWANDOWSKI N,AURAS R,et al. Comparative shelf life study of blackberry fruit in bio-based and petroleumbased containers under retail storage conditions[J]. Food Chemistry,2011,126(4):1734-1740.
[8]ANTUNES L E C,DUARTE FILHO J,DE SOUZA C M. Postharvest conservation of blackberry fruits[J]. Pesquisa Agropecuária Brasileira,2003,38(3):413-419.
[9] PERKINS-VEAZIE P,COLLINS J K,CLARK J R. Shelf-life and quality of ‘Navaho’and ‘Shawnee’blackberry fruit stored under retail storage conditions[J]. Journal of Food Quality,1999,22(5):535-544. DOI:10. 1111/j. 1745-4557.1999.tb00184.
[10]PERKINS V,COLLINS J K. Quality of erect-type blackberry fruit after short intervals of controlled atmosphere storage[J]. Postharvest Biology and Technology,2002,25(2):235-239.DOI:10.1016/50925-5214102700025-x.
[11]ZHANG D P,LI M,WANG Y. Ultrastructural changes in the mesocarp cells of grape berry during its development[J]. Acta Botanica Sinica,1997,39(5):389-396,489-492.
[12]吴文龙,李维林,闾连飞,等.黑莓引种栽培与利用[M].南京:江苏科学技术出版社,2010:30-32.
[13]汤飞云,张春红,胡淑英,等.不同品种黑莓果实发育过程中硬度变化规律的调查与分析[J].江西农业学报,2012,24(10):9-11,14. DOI:10.3969/j.issn.1001-8581.2012.10.003.TANG F Y,ZHANG C H,HU S Y,et al. Changes in fruit firmness of different blackberry cultivars during development and ripening[J]. Acta Agriculturae Jiangxi,2012,24(10):9-11.
[14]汪贵斌,曹福亮,景茂,等.水分胁迫对银杏叶片叶肉细胞超微结构的影响[J].南京林业大学学报(自然科学版),2008,32(5):65-70. DOI:10.3969/j.issn.1000-2006.2008.05.015.WANG G B,CAO F L,JING M,et al. The effect of water stress on mesophyll cell ultrastructure of Ginkgo[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2008,32(5):65-70.
[15]樊汝汶,黄金生.黑杨派两个无性系叶小脉的超微结构[J].南京林业大学学报,1987,11(2):13-17. DOI:10.3969/j.jssn.1000-2006.1987.02.002.FAN R W,HUANG J S. Ultrastructure in the minor leaf vein of two clones of the aegeiros poplar[J]. Journal of Nanjing Forestry University,1987,11(2):13-17.
[16]张宏平,张晋元,吴国良.北方园艺果实组织结构研究进展[J].山西林业科技,2008(4):28-30,36.ZHANG H P,ZHANG J Y,WU G L. The research progress on tissue structure of fruits of deciduous species[J]. Shanxi Forestry Science and Technology,2008(4):28-30,36.
[17]CHALMERS D,ENDE B. A reappraisal of the growth and development of peach fruit[J]. Functional Plant Biology,1975,2(4):623-634.DOI:10.1071/pp9750623.
[18]杨德兴,戴京晶,庞向宇,等.猕猴桃衰老过程中PG,果胶质和细胞壁超微结构的变化[J].园艺学报,1993(4):341-345.YANG D X,DAI J J,PANG X Y,et al. Changes of polygalacturonase,pectin and ultrastructure of cell wall in kiwifruit during senescence[J]. Acta Horticulturae Sinica,1993(4):341-345.
[19]刘愚,吴敦肃,吴有梅,等.乙烯对番茄成熟过程中果皮细胞核超微结构的影响[J].植物生理学报,1990(1):43-48.LIU Y,WU D S,WU Y M,et al. Effects of ethylene on the ultrastructure of nucleus in tomato pericarp cell during ripening[J].Plant Physiology Journal,1990(1):43-48.
[20]PLATT-ALOIA K A,THOMSON W W. Ultrastructure of the mesocarp of mature avocado fruit and changes associated with ripening[J]. Annals of Botany,1981,48(4):451-466.
[21]BEN-ARIE R,KISLEV N,FRENKEL C. Ultrastructural changes in the cell walls of ripening apple and pear fruit[J]. Plant Physiology,1979,64(2):197-202.
[22]张海新,宁久丽,及华.果实采后品质和生理变化研究进展[J].河北农业科学,2010,14(2):54-56.ZHANG H X,NING J L,JI H. Research advances on quality and physiological changes of postharvest fruit[J]. Journal of Hebei Agricultural Sciences,2010,14(2):54-56.
[23]马之胜,王越辉,贾云云,等.桃果实果胶、可溶性糖、可滴定酸含量和果实大小与果实硬度关系的研究[J].江西农业学报,2008,20(10):45-46.MA Z S,WANG Y H,JIA Y Y,et al. Relation between pectin,soluble sugar,titratable acid,fruit size and fruit firmness of peach[J]. Acta Agriculturae Jiangxi,2008,20(10):45-46.
[24]贺军民. GA3和钙处理对草莓采后硬度变化、细胞膜透性及丙二醛含量的影响[J].陕西农业科学,1998,44(3):26-27.
[25]张大鹏,李珉,王毅.葡萄果实发育过程中果肉细胞超微结构的观察[J].植物学报,1997,39(5):389-396.ZHANG D P,LI M,WANG Y. Ultrastructural changes in the mesocarp cells of grape berry during its development[J]. Acta Botanica Sinica,1997,39(5):389-396.
[26]陈睿.植物衰老机理研究进展探讨[J].绿色科技,2012(5):6-9. DOI:10.3969/j.issn.1674-9944.2012.05.005.CHEN R. Research progress on the mechanism of plant senescence[J]. Journal of Green Science and Technology,2012(5):6-9.
[27]李银,张辉,骆建敏,等.蟠桃果实发育成熟过程中果肉细胞超微结构的变化[J].新疆农业科学,2011,48(6):1006-1010.LI Y,ZHANG H,LUO J M,et al. Ultrastructural change of flat peach fruit during fruit development and ripening[J]. Xinjiang Agricultural Sciences,2011,48(6):1006-1010.
[28]卢芳华,余望,肖华山.草莓果实薄壁细胞超微结构的电镜观察[J].福建分析测试,2001,10(1):1383-1385.LU F H,YU W,XIAO H S. TEM observation on the ultra-microstructure of the thin wall cell in the strawberry[J]. Fujian Analysis&Testing,2001,10(1):1383-1385.
[29]张春红,汤飞云,王小敏,等.不同黑莓品种果实形态和硬度分析及其解剖结构和微形态特征观察[J].植物资源与环境学报,2013,22(3):88-94.ZHANG C H,TANG F Y,WANG X M,et al. Analyses on morphology and firmness and observations on anatomical structure and micromorphological characteristics of fruit of different cultivars of blackberry(Rubus spp.)[J]. Journal of Plant Resources and Environment,2013,22(3):88-94.
[30]熊振豪,张春红,李维林,等.黑莓果实转色至成熟进程中相关激素含量的变化[J].北方园艺,2018(12):13-19.XIONG Z H,ZHANG C H,LI W L,et al. Changes of plant hormones during fruit development and ripening process in blackberry fruits[J]. Northern Horticulture,2018(12):13-19.
[31]张春红,熊振豪,吴文龙,等.黑莓果实成熟期木质素合成CAD酶及其基因表达[J].南京林业大学学报(自然科学版),2019,43(1):141-148. DOI:10. 3969/j. issn. 1000-2006.201805004.ZHANG C H,XIONG Z H,WU W L,et al. CAD enzyme activity and gene expression in connection with lignin development and ripening process of blackberry[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2019,43(1):141-148.
[2]BEATTIE J,CROZIER A,DUTHIE G. Potential health benefits of berries[J]. Current Nutrition&Food Science,2005,1(1):71-86.DOI:10.2174/1573401052953294.
[3]吴文龙,王小敏,赵慧芳,等.黑莓品种“Chester”鲜果贮藏性能的研究[J].食品科学,2010,31(8):280-284.WU W L,WANG X M,ZHAO H F,et al. Storage quality of fresh‘Chester’blackberry fruits[J]. Food Science,2010,31(8):280-284.
[4]赵慧芳,王小敏,吴文龙,等.黑莓品种‘Boysen’鲜果贮藏特性的研究[J].植物资源与环境学报,2010,19(3):28-36.ZHAO H F,WANG X M,WU W L,et al. Study on storage property of fresh fruit of blackberry(Rubus spp.)cultivar‘Boysen’[J]. Journal of Plant Resources and Environment,2010,19(3):28-36.
[5]HARDENBURG R E,WATADA A E,WANG C Y. The commercial storage of fruits, vegetables, and florist and nursery stocks[M]//USDA:Agriculture Handbook. Washington DC:USDA,1986.
[6]BOWER C. Postharvest handling,storage,and treatment of fresh market berries[J]. Food Science and Technology, 2007,168:261.
[7]JOO M J,LEWANDOWSKI N,AURAS R,et al. Comparative shelf life study of blackberry fruit in bio-based and petroleumbased containers under retail storage conditions[J]. Food Chemistry,2011,126(4):1734-1740.
[8]ANTUNES L E C,DUARTE FILHO J,DE SOUZA C M. Postharvest conservation of blackberry fruits[J]. Pesquisa Agropecuária Brasileira,2003,38(3):413-419.
[9] PERKINS-VEAZIE P,COLLINS J K,CLARK J R. Shelf-life and quality of ‘Navaho’and ‘Shawnee’blackberry fruit stored under retail storage conditions[J]. Journal of Food Quality,1999,22(5):535-544. DOI:10. 1111/j. 1745-4557.1999.tb00184.
[10]PERKINS V,COLLINS J K. Quality of erect-type blackberry fruit after short intervals of controlled atmosphere storage[J]. Postharvest Biology and Technology,2002,25(2):235-239.DOI:10.1016/50925-5214102700025-x.
[11]ZHANG D P,LI M,WANG Y. Ultrastructural changes in the mesocarp cells of grape berry during its development[J]. Acta Botanica Sinica,1997,39(5):389-396,489-492.
[12]吴文龙,李维林,闾连飞,等.黑莓引种栽培与利用[M].南京:江苏科学技术出版社,2010:30-32.
[13]汤飞云,张春红,胡淑英,等.不同品种黑莓果实发育过程中硬度变化规律的调查与分析[J].江西农业学报,2012,24(10):9-11,14. DOI:10.3969/j.issn.1001-8581.2012.10.003.TANG F Y,ZHANG C H,HU S Y,et al. Changes in fruit firmness of different blackberry cultivars during development and ripening[J]. Acta Agriculturae Jiangxi,2012,24(10):9-11.
[14]汪贵斌,曹福亮,景茂,等.水分胁迫对银杏叶片叶肉细胞超微结构的影响[J].南京林业大学学报(自然科学版),2008,32(5):65-70. DOI:10.3969/j.issn.1000-2006.2008.05.015.WANG G B,CAO F L,JING M,et al. The effect of water stress on mesophyll cell ultrastructure of Ginkgo[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2008,32(5):65-70.
[15]樊汝汶,黄金生.黑杨派两个无性系叶小脉的超微结构[J].南京林业大学学报,1987,11(2):13-17. DOI:10.3969/j.jssn.1000-2006.1987.02.002.FAN R W,HUANG J S. Ultrastructure in the minor leaf vein of two clones of the aegeiros poplar[J]. Journal of Nanjing Forestry University,1987,11(2):13-17.
[16]张宏平,张晋元,吴国良.北方园艺果实组织结构研究进展[J].山西林业科技,2008(4):28-30,36.ZHANG H P,ZHANG J Y,WU G L. The research progress on tissue structure of fruits of deciduous species[J]. Shanxi Forestry Science and Technology,2008(4):28-30,36.
[17]CHALMERS D,ENDE B. A reappraisal of the growth and development of peach fruit[J]. Functional Plant Biology,1975,2(4):623-634.DOI:10.1071/pp9750623.
[18]杨德兴,戴京晶,庞向宇,等.猕猴桃衰老过程中PG,果胶质和细胞壁超微结构的变化[J].园艺学报,1993(4):341-345.YANG D X,DAI J J,PANG X Y,et al. Changes of polygalacturonase,pectin and ultrastructure of cell wall in kiwifruit during senescence[J]. Acta Horticulturae Sinica,1993(4):341-345.
[19]刘愚,吴敦肃,吴有梅,等.乙烯对番茄成熟过程中果皮细胞核超微结构的影响[J].植物生理学报,1990(1):43-48.LIU Y,WU D S,WU Y M,et al. Effects of ethylene on the ultrastructure of nucleus in tomato pericarp cell during ripening[J].Plant Physiology Journal,1990(1):43-48.
[20]PLATT-ALOIA K A,THOMSON W W. Ultrastructure of the mesocarp of mature avocado fruit and changes associated with ripening[J]. Annals of Botany,1981,48(4):451-466.
[21]BEN-ARIE R,KISLEV N,FRENKEL C. Ultrastructural changes in the cell walls of ripening apple and pear fruit[J]. Plant Physiology,1979,64(2):197-202.
[22]张海新,宁久丽,及华.果实采后品质和生理变化研究进展[J].河北农业科学,2010,14(2):54-56.ZHANG H X,NING J L,JI H. Research advances on quality and physiological changes of postharvest fruit[J]. Journal of Hebei Agricultural Sciences,2010,14(2):54-56.
[23]马之胜,王越辉,贾云云,等.桃果实果胶、可溶性糖、可滴定酸含量和果实大小与果实硬度关系的研究[J].江西农业学报,2008,20(10):45-46.MA Z S,WANG Y H,JIA Y Y,et al. Relation between pectin,soluble sugar,titratable acid,fruit size and fruit firmness of peach[J]. Acta Agriculturae Jiangxi,2008,20(10):45-46.
[24]贺军民. GA3和钙处理对草莓采后硬度变化、细胞膜透性及丙二醛含量的影响[J].陕西农业科学,1998,44(3):26-27.
[25]张大鹏,李珉,王毅.葡萄果实发育过程中果肉细胞超微结构的观察[J].植物学报,1997,39(5):389-396.ZHANG D P,LI M,WANG Y. Ultrastructural changes in the mesocarp cells of grape berry during its development[J]. Acta Botanica Sinica,1997,39(5):389-396.
[26]陈睿.植物衰老机理研究进展探讨[J].绿色科技,2012(5):6-9. DOI:10.3969/j.issn.1674-9944.2012.05.005.CHEN R. Research progress on the mechanism of plant senescence[J]. Journal of Green Science and Technology,2012(5):6-9.
[27]李银,张辉,骆建敏,等.蟠桃果实发育成熟过程中果肉细胞超微结构的变化[J].新疆农业科学,2011,48(6):1006-1010.LI Y,ZHANG H,LUO J M,et al. Ultrastructural change of flat peach fruit during fruit development and ripening[J]. Xinjiang Agricultural Sciences,2011,48(6):1006-1010.
[28]卢芳华,余望,肖华山.草莓果实薄壁细胞超微结构的电镜观察[J].福建分析测试,2001,10(1):1383-1385.LU F H,YU W,XIAO H S. TEM observation on the ultra-microstructure of the thin wall cell in the strawberry[J]. Fujian Analysis&Testing,2001,10(1):1383-1385.
[29]张春红,汤飞云,王小敏,等.不同黑莓品种果实形态和硬度分析及其解剖结构和微形态特征观察[J].植物资源与环境学报,2013,22(3):88-94.ZHANG C H,TANG F Y,WANG X M,et al. Analyses on morphology and firmness and observations on anatomical structure and micromorphological characteristics of fruit of different cultivars of blackberry(Rubus spp.)[J]. Journal of Plant Resources and Environment,2013,22(3):88-94.
[30]熊振豪,张春红,李维林,等.黑莓果实转色至成熟进程中相关激素含量的变化[J].北方园艺,2018(12):13-19.XIONG Z H,ZHANG C H,LI W L,et al. Changes of plant hormones during fruit development and ripening process in blackberry fruits[J]. Northern Horticulture,2018(12):13-19.
[31]张春红,熊振豪,吴文龙,等.黑莓果实成熟期木质素合成CAD酶及其基因表达[J].南京林业大学学报(自然科学版),2019,43(1):141-148. DOI:10. 3969/j. issn. 1000-2006.201805004.ZHANG C H,XIONG Z H,WU W L,et al. CAD enzyme activity and gene expression in connection with lignin development and ripening process of blackberry[J]. Journal of Nanjing Forestry University(Natural Sciences Edition),2019,43(1):141-148.
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