‘中油桃9号’及其黄肉突变体花和叶片中类胡萝卜素代谢及相关基因的时空表达
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摘要
以‘中油桃9号’及其黄肉芽变的花和叶片为试材,采用HPLC法对类胡萝卜素的积累水平进行定性和定量分析,实时荧光定量PCR法对类胡萝卜素代谢关键基因的表达水平进行分析。结果表明:在花萼中,黄肉突变体的总类胡萝卜素水平为‘中油桃9号’的5.7倍,其中的β–胡萝卜素为‘中油桃9号’的19.6倍;除DXS和LCY-E外,突变体花萼中无论是合成类胡萝卜素的基因,还是降解类胡萝卜素基因的表达水平均显著高于‘中油桃9号’。在花瓣中,两个材料的类胡萝卜素总量无显著差异,但各类胡萝卜素成分含量存在显著差异;突变体DXS、HDR、PDS、LCY-E、CHY-B和CCD1的表达量显著高于‘中油桃9号’,ZDS、NCED1和CCD4的表达量显著低于‘中油桃9号’。在叶片中,‘中油桃9号’和黄肉突变体均以β–胡萝卜素、紫黄质、9–顺式–紫黄质和叶黄质为主,幼果期时两个材料的类胡萝卜素总量无显著差异,果实成熟后突变体的类胡萝卜素总量呈上升趋势,显著高于‘中油桃9号’;自花后55～80 d,仅CCD4的表达模式与类胡萝卜素的积累一致,因而认为CCD4的表达差异是两个材料后期叶片类胡萝卜素含量差异的主要原因。
Two nectarine cultivars,CN9(white-fleshed)and its yellow-fleshed mutant CN9 Y,were used as materials to investigate carotenoids metabolism and genes expression. Qualitative and quantitative levels of carotenoids were analyzed by using HPLC,the expression of genes related to carotenoid accumulation were detected via real-time qPCR method. The results showed that the total carotenoid and β-carotene content of CN9 Y nectarine were 5.7 and 19.6 times higher than that of CN9 respectively in calyxes. Carotenoid metabolism genes of both biosynthesis and degradation showed significant higher expression level in CN9 Y calyxes than in CN9,except DXS and LCY-E. In petals,the total carotenoid content was similar between CN9 and CN9 Y,but their carotenoid composition contents were different. The gene expression levels of DXS,HDR,PDS,LCY-E,CHY-B and CCD1 were higher in CN9 Y petals than in CN9,but ZDS,NCED1 and CCD4 were lower in CN9 Y than in CN9. In leaves,the carotenoid compositions of CN9 and CN9 Y were all dominated by β-carotene,violaxanthin,cis-9-violaxanthin and lutein,the total carotenoid content was similar between CN9 and CN9 Y during early fruit development stage,but CN9 Y obviously increased after fruits matured,however,CN9 was still stabilized. During 55–80 d after flowering,only the expression level of CCD4 was in accordance with carotenoid accumulation,it is suggested that the difference of CCD4 expression level may lead to the difference of carotenoid accumulation in leaves between the two materials in later stage.
Two nectarine cultivars,CN9(white-fleshed)and its yellow-fleshed mutant CN9 Y,were used as materials to investigate carotenoids metabolism and genes expression. Qualitative and quantitative levels of carotenoids were analyzed by using HPLC,the expression of genes related to carotenoid accumulation were detected via real-time qPCR method. The results showed that the total carotenoid and β-carotene content of CN9 Y nectarine were 5.7 and 19.6 times higher than that of CN9 respectively in calyxes. Carotenoid metabolism genes of both biosynthesis and degradation showed significant higher expression level in CN9 Y calyxes than in CN9,except DXS and LCY-E. In petals,the total carotenoid content was similar between CN9 and CN9 Y,but their carotenoid composition contents were different. The gene expression levels of DXS,HDR,PDS,LCY-E,CHY-B and CCD1 were higher in CN9 Y petals than in CN9,but ZDS,NCED1 and CCD4 were lower in CN9 Y than in CN9. In leaves,the carotenoid compositions of CN9 and CN9 Y were all dominated by β-carotene,violaxanthin,cis-9-violaxanthin and lutein,the total carotenoid content was similar between CN9 and CN9 Y during early fruit development stage,but CN9 Y obviously increased after fruits matured,however,CN9 was still stabilized. During 55–80 d after flowering,only the expression level of CCD4 was in accordance with carotenoid accumulation,it is suggested that the difference of CCD4 expression level may lead to the difference of carotenoid accumulation in leaves between the two materials in later stage.
引文
Adami M,Franceschi P D,Brandi F,Liverani A,Giovannini D,Rosati C,Dondini L,Tartarini S.2013.Identifying a carotenoid cleavage dioxygenase(CCD4)gene controlling yellow/white fruit flesh color of peach.Plant Molecular Biology Reporter,31:1166-1175.
Alder A,Jamil M,Marzorati M,Bruno M,Vermathen M,Bigler P,Ghisla S,Bouwmeester H,Beyer P,Al-Babili S.2012.The path fromβ-carotene to carlactone,a strigolactone-like plant hormone.Science,335:1348-1351.
Andreaz D V,Ilaria M,Anna S,Fabio G,Salvatore C.2007.The antioxidant profile of three different peaches cultivars(Prunus persica)and their short-term effect on antioxidant status in human.European Food Research and Technology,225:167-172.
Arango J,Welsch R.2014.Carotene hydroxylase activity determines the levels of bothα-carotene and total carotenoids in orange carrots.Plant Cell,26:2223-2233.
Auldridge M E,Mccarty D R,Klee H J.2006.Plant carotenoid cleavage oxygenases and their apocarotenoid products.Current Opinion in Plant Biology,9:315-321.
Bartley G E,Scolnik P A.1995.Plant carotenoids:pigments for photoprotection,visual attraction,and human health.Plant Cell,7:1027-1038.
Beyer P,Albabili S,Ye X,Lucca P,Schaub P,Welsch R,Potrykus I.2002.Golden rice:introducing theβ-carotene biosynthesis pathway into rice endosperm by genetic engineering to defeat vitamin A deficiency.Journal of Nutrition,132:5065-5105.
Brandi F,Bar E,Mourgues F,Horváth G,Turcsi E,Giuliano G,Liverani A,Tartarini S,Lewinsohn E,Rosati C.2011.Study of‘Redhaven’peach and its white-fleshed mutant suggests a key role of CCD4 carotenoid dioxygenase in carotenoid and norisoprenoid volatile metabolism.BMC Plant Biology,11:1-14.
Cazzonelli C I,Pogson B J.2010.Source to sink:regulation of carotenoid biosynthesis in plants.Trends in Plant Science,15:266-274.
Falchi R,Vendramin E,Zanon L,Scalabrin S,Cipriani G,Verde I,Vizzotto G,Morgante M.2013.Three distinct mutational mechanisms acting on a single gene underpin the origin of yellow flesh in peach.The Plant Journal,76:175-187.
Gomez-Roldan V,Fermas S,Brewer P B,Puech-Pages V,Dun E A,Pillot J P,Letisse F,Matusova R,Danoun S,Portais J C,Bouwmeester H,Bécard G,Beveridge C A,Rameau C,Rochange S F.2008.Strigolactone inhibition of shoot branching.Nature,455:189-194.
Hannoufa A,Hossain Z.2012.Regulation of carotenoid accumulation in plants.Biocatalysis&Agricultural Biotechnology,1:198-202.
Kato M,Matsumoto H,Ikoma Y,Okuda H,Yano M.2006.The role of carotenoid cleavage dioxygenases in the regulation of carotenoid profiles during maturation in citrus fruit.Journal of Experimental Botany,57:2153-2164.
Kilcrease J,Collins A M,Richins R D,Timlin J A,O'Connell M A.2013.Multiple microscopic approaches demonstrate linkage between chromoplast architecture and carotenoid composition in diverse Capsicum annuum fruit.Plant Journal for Cell&Molecular Biology,76:1074-1083.
Liu H,Kishimoto S,Yamamizo C,Fukuta N,Ohmiya A.2013.Carotenoid accumulations and carotenogenic gene expressions in the petals of Eustoma grandiflorum.Plant Breeding,132:417-422.
Li L,Paolillo D J,Parthasarathy M V,Dimuzio E M,Garvin D F.2001.A novel gene mutation that confers abnormal patterns ofβ-carotene accumulation in cauliflower(Brassica oleracea var.botrytis).Plant Journal for Cell&Molecular Biology,26:59-67.
Li L,Yuan H.2013.Chromoplast biogenesis and carotenoid accumulation.Archives of Biochemistry and Biophysics,539:102-109.
Li Pei-rong,Zhang Shu-jiang,Zhang Shi-fan,Li Fei,Zhang Hui,Liu Xin-yan,Wu Jian,Wang Xiao-wu,Sun Ri-fei.2014.Analysis of the carotenoids pigments and orange inner leaf gene in Chinese cabbage(Brassica rapa).Acta Horticulturae Sinica,41(3):469-478.(in Chinese)李佩荣,张淑江,章时蕃,李菲,张慧,刘新艳,武剑,王晓武,孙日飞.2014.大白菜橘红心类胡萝卜素组分及其基因分析.园艺学报,41(3):469-478.
Liu Qing.2008.Molecular mechanism for the altered traits of the red flesh bud sport of‘Anliu’sweet orange[Ph.D.Dissertation].Wuhan:Huangzhong Agricultural University.(in chinese)刘庆.2008.‘暗柳’甜橙红色突变体性状形成的分子机理研究[博士论文].武汉:华中农业大学.
Livak K J,Schmittgen T D.2001.Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method.Methods,25:402-408.
Lopez-Juez E,Pyke K A.2005.Plastids unleashed:their development and their integration in plant development.The International Journal of Developmental Biology,49:557-577.
Lu S,Van Eck J,Zhou X,Lopez A B,O'Halloran D M,Cosman K M,Conlin B J,Paolillo D J,Garvin D F,Vrebalov J,Kochian L V,Küpper H,Earle E D,Cao J,Li L.2006.The cauliflower Or gene encodes a DnaJ cysteine-rich domain-containing protein that mediates high levels ofβ-carotene accumulation.Plant Cell,18:3594-3605.
Ma J J,Li J,Zhao J B,Zhou H,Ren F,Wang L,Gu C,Liao L,Han Y P.2014.Inactivation of a gene encoding carotenoid cleavage dioxygenase(CCD4)leads to carotenoid-based yellow coloration of fruit flesh and leaf midvein in peach.Plant Molecular Biology Reporter,32:246-257.
Marty I,Bureau S,Sarkissian G,Gouble B,Audergon J M,Albagnac G.2005.Ethylene regulation of carotenoid accumulation and carotenogenic gene expression in colour-contrasted apricot varieties(Prunus armeniaca).Journal of Experimental Botany,56:1877-1886.
Moehs C P,Tian Li,Osteryoung K W,DellaPenna D.2001.Analysis of carotenoid biosynthetic gene expression during marigold petal development.Plant Molecular Biology,45:281-293.
Nisar N,Li L,Lu S,Khin N C,Pogson B J.2015.Carotenoid metabolism in plants.Molecular Plant,8:68-82.
Nogueira M,Mora L,Enfissi E M A,Bramley P M,Fraser P D.2013.Subchromoplast sequestration of carotenoids affects regulatory mechanisms in tomato lines expressing different carotenoid gene combinations.Plant Cell,25:4560-4579.
Ohmiya A,Kishimoto S,Aida R,Yoshioka S,Sumitomo K.2006.Carotenoid cleavage dioxygenase(CmCCD4)contributes to white color formation in chrysanthemum petals.Plant Physiology,142:1193-1201.
Ruyter-Spira C,Al-Babili S,Krol S V D,Bouwmeester H.2013.The biology of strigolactones.Trends in Plant Science,18:72-83.
Song Cheng-xiu,Zhang Li-yi,Zhang Cai-xia,Tian Yi,Cong Pei-hua.2017.Differential expression analysis of the fruits of yellow-fleshed apple and exploits genes involved in carotenoid pigmentation.Acta Horticulturae Sinica,44(2):205-214.(in Chinese)宋成秀,张利义,张彩霞,田义,丛佩华.2017.黄肉苹果转色期前后类胡萝卜素合成相关差异表达基因鉴定.园艺学报,44(2):205-214.
Tan B C,Joseph L M,Deng W T,Liu L J,Li Q B,Cline K,McCarty D R.2003.Molecular characterization of the Arabidopsis 9-cis epoxycarotenoid dioxygenase gene family.Plant Journal for Cell&Molecular Biology,35:44-56.
Toledo-Ortiz G,Huq E,Rodríguez-Concepción M.2010.From the cover:direct regulation of phytoene synthase gene expression and carotenoid biosynthesis by phytochrome-interacting factors.Proceedings of the National Academy of Sciences of USA,107:11626-11631.
Umehara M,Hanada A,Yoshida S,Akiyama K,Arite T,Takeda-Kamiya N,Magome H,Kamiya Y,Shirasu K,Yoneyama K,Kyozuka J,Yamaguchi S.2008.Inhibition of shoot branching by new terpenoid plant hormones.Nature,455:195-200.
Vaio C D,Graziani G,Marra L,Cascone A,Ritieni A.2008.Antioxidant capacities,carotenoids and polyphenols evaluation of fresh and refrigerated peach and nectarine cultivars from Italy.European Food Research and Technology,227:1225-1231.
Vallabhaneni R,Bradbury L M,Wurtzel E T.2010.The carotenoid dioxygenase gene family in maize,sorghum,and rice.Archives of Biochemistry and Biophysics,504:104-111.
Wang Hui,Ou Cheng-gang,Zhuang Fei-yun,Zhao Zhi-wei,Ma Zhen-guo.2014.Relationship of carotenoid accumulation and transcript of main genes in carotenoid biosynthesis in carrot.Acta Horticulturae Sinica,41(12):2513-2520.(in Chinese)王慧,欧承刚,庄飞云,赵志伟,马振国.2014.胡萝卜中类胡萝卜素积累与主要合成基因转录水平相关性分析.园艺学报,41(12):2513-2520.
Xu Jian-lan,Ma Rui-juan,Yu Ming-liang,Shen Zhi-jun,Yin Shou-fang.2010.Changes of flesh pigment contents of blood-flesh peach during fruit development.Jiangsu Journal of Agricultural Science,26:1347-1351.(in Chinese)许建兰,马瑞娟,俞明亮,沈志军,殷守防.2010.红肉桃果实发育过程中果肉色素含量的变化.江苏农业学报,26:1347-1351.
Yamamizo C,Kishimoto S,Ohmiya A.2010.Carotenoid composition and carotenogenic gene expression during ipomoea petal development.Journal of Experimental Botony,61:709-719.
Yan Shao-bin,Zhang Yu-yan,Ma Rui-juan,Yu Ming-liang.2013.Changes of carotenoids composition of yellow peach during fruit development.Journal of Fruit Science,30:260-266.(in Chinese)颜少宾,张妤艳,马瑞娟,俞明亮.2013.黄肉桃果实发育阶段类胡萝卜素的变化.果树学报,30:260-266.
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Zhu Yun-qin,Zeng Wen-fang,Lu Zhen-hua,Niu Liang,Cui Guo-chao,Wang Zhing-qiang.2015.Carotenoid metabolism and gene expression analysis of‘CN9’nectarine and its yellow flesh mutant‘CN9Y’.Acta Horticulturae Sinica,42(4):623-632.(in Chinese)朱运钦,曾文芳,鲁振华,牛良,催国朝,王志强.2015.‘中油桃9号’及其黄肉芽变的类胡萝卜素代谢和基因表达分析.园艺学报,42(4):623-632.
Alder A,Jamil M,Marzorati M,Bruno M,Vermathen M,Bigler P,Ghisla S,Bouwmeester H,Beyer P,Al-Babili S.2012.The path fromβ-carotene to carlactone,a strigolactone-like plant hormone.Science,335:1348-1351.
Andreaz D V,Ilaria M,Anna S,Fabio G,Salvatore C.2007.The antioxidant profile of three different peaches cultivars(Prunus persica)and their short-term effect on antioxidant status in human.European Food Research and Technology,225:167-172.
Arango J,Welsch R.2014.Carotene hydroxylase activity determines the levels of bothα-carotene and total carotenoids in orange carrots.Plant Cell,26:2223-2233.
Auldridge M E,Mccarty D R,Klee H J.2006.Plant carotenoid cleavage oxygenases and their apocarotenoid products.Current Opinion in Plant Biology,9:315-321.
Bartley G E,Scolnik P A.1995.Plant carotenoids:pigments for photoprotection,visual attraction,and human health.Plant Cell,7:1027-1038.
Beyer P,Albabili S,Ye X,Lucca P,Schaub P,Welsch R,Potrykus I.2002.Golden rice:introducing theβ-carotene biosynthesis pathway into rice endosperm by genetic engineering to defeat vitamin A deficiency.Journal of Nutrition,132:5065-5105.
Brandi F,Bar E,Mourgues F,Horváth G,Turcsi E,Giuliano G,Liverani A,Tartarini S,Lewinsohn E,Rosati C.2011.Study of‘Redhaven’peach and its white-fleshed mutant suggests a key role of CCD4 carotenoid dioxygenase in carotenoid and norisoprenoid volatile metabolism.BMC Plant Biology,11:1-14.
Cazzonelli C I,Pogson B J.2010.Source to sink:regulation of carotenoid biosynthesis in plants.Trends in Plant Science,15:266-274.
Falchi R,Vendramin E,Zanon L,Scalabrin S,Cipriani G,Verde I,Vizzotto G,Morgante M.2013.Three distinct mutational mechanisms acting on a single gene underpin the origin of yellow flesh in peach.The Plant Journal,76:175-187.
Gomez-Roldan V,Fermas S,Brewer P B,Puech-Pages V,Dun E A,Pillot J P,Letisse F,Matusova R,Danoun S,Portais J C,Bouwmeester H,Bécard G,Beveridge C A,Rameau C,Rochange S F.2008.Strigolactone inhibition of shoot branching.Nature,455:189-194.
Hannoufa A,Hossain Z.2012.Regulation of carotenoid accumulation in plants.Biocatalysis&Agricultural Biotechnology,1:198-202.
Kato M,Matsumoto H,Ikoma Y,Okuda H,Yano M.2006.The role of carotenoid cleavage dioxygenases in the regulation of carotenoid profiles during maturation in citrus fruit.Journal of Experimental Botany,57:2153-2164.
Kilcrease J,Collins A M,Richins R D,Timlin J A,O'Connell M A.2013.Multiple microscopic approaches demonstrate linkage between chromoplast architecture and carotenoid composition in diverse Capsicum annuum fruit.Plant Journal for Cell&Molecular Biology,76:1074-1083.
Liu H,Kishimoto S,Yamamizo C,Fukuta N,Ohmiya A.2013.Carotenoid accumulations and carotenogenic gene expressions in the petals of Eustoma grandiflorum.Plant Breeding,132:417-422.
Li L,Paolillo D J,Parthasarathy M V,Dimuzio E M,Garvin D F.2001.A novel gene mutation that confers abnormal patterns ofβ-carotene accumulation in cauliflower(Brassica oleracea var.botrytis).Plant Journal for Cell&Molecular Biology,26:59-67.
Li L,Yuan H.2013.Chromoplast biogenesis and carotenoid accumulation.Archives of Biochemistry and Biophysics,539:102-109.
Li Pei-rong,Zhang Shu-jiang,Zhang Shi-fan,Li Fei,Zhang Hui,Liu Xin-yan,Wu Jian,Wang Xiao-wu,Sun Ri-fei.2014.Analysis of the carotenoids pigments and orange inner leaf gene in Chinese cabbage(Brassica rapa).Acta Horticulturae Sinica,41(3):469-478.(in Chinese)李佩荣,张淑江,章时蕃,李菲,张慧,刘新艳,武剑,王晓武,孙日飞.2014.大白菜橘红心类胡萝卜素组分及其基因分析.园艺学报,41(3):469-478.
Liu Qing.2008.Molecular mechanism for the altered traits of the red flesh bud sport of‘Anliu’sweet orange[Ph.D.Dissertation].Wuhan:Huangzhong Agricultural University.(in chinese)刘庆.2008.‘暗柳’甜橙红色突变体性状形成的分子机理研究[博士论文].武汉:华中农业大学.
Livak K J,Schmittgen T D.2001.Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCT method.Methods,25:402-408.
Lopez-Juez E,Pyke K A.2005.Plastids unleashed:their development and their integration in plant development.The International Journal of Developmental Biology,49:557-577.
Lu S,Van Eck J,Zhou X,Lopez A B,O'Halloran D M,Cosman K M,Conlin B J,Paolillo D J,Garvin D F,Vrebalov J,Kochian L V,Küpper H,Earle E D,Cao J,Li L.2006.The cauliflower Or gene encodes a DnaJ cysteine-rich domain-containing protein that mediates high levels ofβ-carotene accumulation.Plant Cell,18:3594-3605.
Ma J J,Li J,Zhao J B,Zhou H,Ren F,Wang L,Gu C,Liao L,Han Y P.2014.Inactivation of a gene encoding carotenoid cleavage dioxygenase(CCD4)leads to carotenoid-based yellow coloration of fruit flesh and leaf midvein in peach.Plant Molecular Biology Reporter,32:246-257.
Marty I,Bureau S,Sarkissian G,Gouble B,Audergon J M,Albagnac G.2005.Ethylene regulation of carotenoid accumulation and carotenogenic gene expression in colour-contrasted apricot varieties(Prunus armeniaca).Journal of Experimental Botany,56:1877-1886.
Moehs C P,Tian Li,Osteryoung K W,DellaPenna D.2001.Analysis of carotenoid biosynthetic gene expression during marigold petal development.Plant Molecular Biology,45:281-293.
Nisar N,Li L,Lu S,Khin N C,Pogson B J.2015.Carotenoid metabolism in plants.Molecular Plant,8:68-82.
Nogueira M,Mora L,Enfissi E M A,Bramley P M,Fraser P D.2013.Subchromoplast sequestration of carotenoids affects regulatory mechanisms in tomato lines expressing different carotenoid gene combinations.Plant Cell,25:4560-4579.
Ohmiya A,Kishimoto S,Aida R,Yoshioka S,Sumitomo K.2006.Carotenoid cleavage dioxygenase(CmCCD4)contributes to white color formation in chrysanthemum petals.Plant Physiology,142:1193-1201.
Ruyter-Spira C,Al-Babili S,Krol S V D,Bouwmeester H.2013.The biology of strigolactones.Trends in Plant Science,18:72-83.
Song Cheng-xiu,Zhang Li-yi,Zhang Cai-xia,Tian Yi,Cong Pei-hua.2017.Differential expression analysis of the fruits of yellow-fleshed apple and exploits genes involved in carotenoid pigmentation.Acta Horticulturae Sinica,44(2):205-214.(in Chinese)宋成秀,张利义,张彩霞,田义,丛佩华.2017.黄肉苹果转色期前后类胡萝卜素合成相关差异表达基因鉴定.园艺学报,44(2):205-214.
Tan B C,Joseph L M,Deng W T,Liu L J,Li Q B,Cline K,McCarty D R.2003.Molecular characterization of the Arabidopsis 9-cis epoxycarotenoid dioxygenase gene family.Plant Journal for Cell&Molecular Biology,35:44-56.
Toledo-Ortiz G,Huq E,Rodríguez-Concepción M.2010.From the cover:direct regulation of phytoene synthase gene expression and carotenoid biosynthesis by phytochrome-interacting factors.Proceedings of the National Academy of Sciences of USA,107:11626-11631.
Umehara M,Hanada A,Yoshida S,Akiyama K,Arite T,Takeda-Kamiya N,Magome H,Kamiya Y,Shirasu K,Yoneyama K,Kyozuka J,Yamaguchi S.2008.Inhibition of shoot branching by new terpenoid plant hormones.Nature,455:195-200.
Vaio C D,Graziani G,Marra L,Cascone A,Ritieni A.2008.Antioxidant capacities,carotenoids and polyphenols evaluation of fresh and refrigerated peach and nectarine cultivars from Italy.European Food Research and Technology,227:1225-1231.
Vallabhaneni R,Bradbury L M,Wurtzel E T.2010.The carotenoid dioxygenase gene family in maize,sorghum,and rice.Archives of Biochemistry and Biophysics,504:104-111.
Wang Hui,Ou Cheng-gang,Zhuang Fei-yun,Zhao Zhi-wei,Ma Zhen-guo.2014.Relationship of carotenoid accumulation and transcript of main genes in carotenoid biosynthesis in carrot.Acta Horticulturae Sinica,41(12):2513-2520.(in Chinese)王慧,欧承刚,庄飞云,赵志伟,马振国.2014.胡萝卜中类胡萝卜素积累与主要合成基因转录水平相关性分析.园艺学报,41(12):2513-2520.
Xu Jian-lan,Ma Rui-juan,Yu Ming-liang,Shen Zhi-jun,Yin Shou-fang.2010.Changes of flesh pigment contents of blood-flesh peach during fruit development.Jiangsu Journal of Agricultural Science,26:1347-1351.(in Chinese)许建兰,马瑞娟,俞明亮,沈志军,殷守防.2010.红肉桃果实发育过程中果肉色素含量的变化.江苏农业学报,26:1347-1351.
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