红色砂梨花青苷生物合成相关基因分离及表达研究
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摘要
砂梨原产于我国,是东亚地区栽培最广的类型,以往栽培的砂梨品种主要以黄绿色和褐色(实际是木栓层)为主,红色砂梨品种相对较少。近年来,以原产云南的著名红色砂梨地方品种‘火把梨’与日本梨‘幸水’为亲本育成的‘美人酥’、‘满天红’以及从地方品种中筛选出的‘云红梨1号’等优良红色品种已在国内很多地方引种栽培,得到了消费者的认可,市场潜力巨大。然而,生产上,红色砂梨着色表现不佳,严重影响了果实特有外观品质的展现。因此,开展套袋及生态条件影响红色砂梨花青苷合成的机制研究,将为开发适合生产实际的着色调控技术,解决红色砂梨的着色不良问题,改善其外观品质,以及今后通过现代生物学技术改善红色砂梨着色情况提供重要理论支持。
本试验以红色砂梨中熟品种‘美人酥’和晚熟品种‘云红梨1号’果皮为材料,通过已报道植物序列设计特异或兼并引物和参考苹果EST序列两种途径,利用RACE技术,分离得到2个PAL成员,分别为PpPAL1和PpPAL2;4个CHS成员,分别为PpCHS1、PpCHS2、PpCHS3和PpCHS4；3个CHI成员,分别为PpCHI1、PpCHI2和PpCHI3;1个PpF3Hl成员；2个DFR成员,分别为PpDFR1和PpDFR2;1个PpANS1成员；2个UFGT成员,分别为PpUFGT1和PpUFGT2;已分离转录因子PpMYBl和PpMYB10,其中PpMYB10属于第十亚组的R2R3MYB类群。
套袋显著改善红色砂梨果实着色。本研究以红色砂梨‘美人酥’套袋果为材料,研究商业采收前15d去袋后果面着色过程中花青苷合成相关基因表达模式。PpPAL1、PpCHSl、PpCHI1、PpF3H1、PpDFRl、PpANSl和PpUFGT1受到光照诱导表达量迅速上升,去袋后1d表达量均达到最大值,明显提前于花青苷的合成高峰。PpCHSl、PpF3H1、PpANSl和PpUFGT1在去袋后的前5d内都保持相对较高的表达水平。7个结构基因被光照诱导后表达量都迅速上升,说明合成途径中的结构基因通过协同作用参与花青苷的合成。光质和光强对果实着色都具有重要影响,同一个果实的阳面和阴面着色完全不同,在‘美人酥’的阴面果皮几乎不合成花青苷,定量结果显示阴面果皮中7个结构基因表达量都低于阳面果皮,其中PpCHS1、PpUFGT1、PpF3H1和PpANSl等4个基因在阳面果皮表达量的最大值(1DABR)与阴面的高峰(1DABR)比值超过2.7。PpCHS1、PpF3H1、 PpANS1和PpUFGT1等在成熟果实果皮中的表达均显著高于其它器官,同时PpCHS1与PpANS1在淡红色幼叶中的表达也较高。与成熟果中花青苷含量呈正相关关系。在红色砂梨‘美人酥’去袋着色过程中,PpMYB10与花青苷合成途径中的结构基因协同表达,而PpMYB1没有表现出相应的协同表达现象。
花青苷合成相关的结构基因家族成员在序列和表达模式上均存在较大差异。在氨基酸水平上,PpCHS1和PpCHS2的序列同源性高达97%,而PpCHI1和PpCHI2最低,仅有23%。红色砂梨‘云红梨1号’去袋后,花青苷合成途径中的结构基因部分成员被诱导,包括(?)PpPAL1、PpCHS1、PpCHS2、PpCHI1、PpCHI3、PpF3H1、 PpDFR1、PpANS1、PpUFGT1和PpUFGT2。而PpPAL2、PpCHS3、PpCHS4和PpCHI2的表达没有明显变化,由此推测这些基因可能不是果皮花青苷合成的关键基因。被诱导的基因中PpPAL1、PpCHS1、PpCHI1、PpCHI3、PpF3HI1、pDFR1、 PpANS1和PpUFGT1的表达高峰都出现在去袋后1d,另外两个基因PpCHS2和PpUFGT2的转录水平则在去袋后3d达到最大值,表现出不同的表达模式,这些结构基因可能通过协同作用共同调控花青苷的合成。
果实的着色受基因型决定,环境条件影响着色的程度。位于云南的安宁气候特点为亚热带高原气候,伴随着高海拔,强辐射和昼夜温差大等特点；河南的荥阳则为北温带大陆性季风气候,伴随着低海拔,低辐射和昼夜温差小等特点。本试验比较了红色砂梨‘满天红’在两个不同生态区套袋处理果实着色及品质相关生理分子研究。商业采收时,安宁较荥阳具有更好的着色,果皮中花青苷和类胡萝卜素含量较后者高,叶绿素含量较后者低；单果重和果实的硬度前者较后者低；前者果肉中可溶性固形物和糖含量较后者高,淀粉含量较后者低；着色过程中前者果皮花青苷合成结构和调节基因的表达量均较后者高。统计分析显示可溶性糖和花青苷含量在安宁(r=0.96981)和荥阳(r=0.94433)都显著相关。安宁具有的特定生态因子,特别是高海拔,强辐射和昼夜温差大等特点有利于果实中糖积累和诱导果皮中花青苷合成结构和调节基因,尤其是PpCHS1、PpANS1、 PpUFGT1、PpMYB10和PpbHLH的表达,促进了花青苷的合成。
Chinese sand pear (Pyrus pyrifolia Nakai) originated from China and is widely cultivated in East Asia. Ripe sand pears usually appear green, yellow or brown (actually is phellem). Red sand pear also could be found, but not as common as others. In recent years, red skin sand pear, such as'Meirensu','Mantianhong'and'Yunhongli No.1', were well received by customers, showing a great potential of custom market. However, bad color development of red Chinese sand pear greatly influenced the fruit appearance. The study on anthocyanin synthesis mechanism of red Chinese sand pear affected by fruit bagging treatments and different habitats will be of great help to develop a new technology to product high quality fruits.
Two red Chinese sand pear cultivars,'Meirensu'(mid-maturing) and'Yunhongli No.1'(late-maturing) in different maturation periods, were used as materials in this study. Two ways were used to isolate the gene sequences, one is used specific primers or degenerate primers which were designed according to plant sequences from available databases, the other is used specific primers based on malus EST database. Combined with RACE methods, two PALS (PpPALl and PpPAL2), four CHSS (PpCHSl, PpCHS2, PpCHS3and PpCHS4), three CHIS (PpCHI1, PpCHI2and PpCH13), one F3H (PpF3H1), two DFRs(PpDFRl and PpDFR2), one ANS (PpANSl) and two UFGTs(PpUFGT1and PpUFGT2) were isolated from red chinese sand pear. Transcription factors PpMYB1and PpMYB10were cloned, and sequence homology analysis showed PpMYB10belong to the subgroup10R2R3MYBs.
The coloration of red Chinese sand pears was significantly improved by fruit bagging treatments. Red Chinese sand pear'Meirensu'was used for fruit bagging treatments. Bags were removed15days prior to harvest and the fruits were completely re-exposed to light until harvest. Structural genes and transcription factors that involved in anthocyanin synthesis were analyzed during fruit coloration after the bags removal. Following bag removal, the expression levels of PpPAL1, PpCHSl, PpCHIl, PpF3Hl, PpDFR1, PpANSl and PpUFGTl were markedly enhanced in the sunlit side of 'Meirensu' fruit; PpCHS1, PpF3H1, PpANSl and PpUFGT1maintained high expression levels in5days after the bags removal (DABG). All seven structural genes were enhanced by sunlight, indicated both genes were involved in the anthocyanin biosynthesis. Light intensity and quality are important for fruit coloration. Anthocyanin amounts increased linearly in the skin of the sunlit side of'Meirensu' fruits. However, anthocyanin content was barely detected in the skin of the shaded side. The transcript level of anthocyanin biosynthetic genes was markedly up-regulated in the skin of sunlit side of'Meirensu', but much lower in the skin from the shaded side. The expression level of PpCHSl, PpUFGT1, PpF3H1and PpANSl (1DABR) in sunlit side as compared to shaded side is above2.7folds. PpCHS1, PpF3Hl, PpANSl and PpUFGTl expressed at higher level in the skin of mature fruits than in other tissues, and PpCHSl, PpANSl also showed high expression in young leaves with faint red. The expression level was coordinated with anthocyanin level in different organs. The expression pattern of PpMYB10was similar to other structural genes during the coloration of 'Meirensu' after the bags removal. However, PpMYB1showed less coordination with other genes.
There were family members in structural genes that involved in anthocyanin synthesis. They showed many differences in sequences and expression patterns. The highest percentage for amino acid sequences was found between PpCHSl and PpCHS2(97%), while the lowest one was PpCHIl and PpCHI2(23%). The expression levels of PpPAL1, PpCHSl, PpCHS2, PpCHI1, PpCHI3, PpF3Hl, PpDFR1, PpANS1, PpUFGT1and PpUFGT2was markedly enhanced in the peel of 'Yunhongli No.1'fruit after the bags removal. Other structural genes including PpPAL2, PpCHS3, PpCHS4and PpCHI2showed no response, indicating that they may not involved in anthocyanin synthesis during fruit coloration. The expression pattern of these genes that involved in anthocyanin synthesis shows different ways, transcript levels of PpPAL1, PpCHS1, PpCHI1, PpCHI3, PpF3H1, PpDFR1,PpANSl and PpUFGTl peaked at1DABR. However, PpCHS2and PpUFGT2peaked at3DABR. All these genes might be involved in anthocyanin biosynthesis by being coordinately expressed during anthocyanin accumulation.
The formation of anthocyanin was not only genetically determined, but also might be influenced by many environmental factors. Anning is located in Yunnan province, having a warm, high irradiant climate and high day and night temperature difference, while Xingyang is located in Henan province, with a hot, low irradiant climate and low day and night temperature difference. At harvest, red Chinese sand pear'Mantianhong'in Anning showed better coloration, higher concentrations of anthocyanin, carotenoid, total soluble solid (TSS) and soluble sugar but lower chlorophyll in pear peel, fruit weight, firmness and content of starch than those in Xingyang. Significantly increasing in the anthocyanin structural and regulatory genes, especially for PpCHS, PpANS, PpUFGT, PpMYB10and PpbHLH in red Chinese sand pears were found in materials sampled in Anning. The statistical analysis indicated that soluble sugar content was significantly associated with anthocyanin accumulation in both Anning (r=0.96981) and Xingyang (r=0.94433) habitats. Those results indicated high altitude, high-irradiant and high temperature differences in Anning habit were more conducive to the color development and inner quality of red Chinese sand pear, which was probably resulted in the higher concentration of sugar and expression of anthocyanin structural and regulatory genes, especially for the expression of PpCHS, PpANS, PpUFGT, PpMYB10and PpbHLH.
本试验以红色砂梨中熟品种‘美人酥’和晚熟品种‘云红梨1号’果皮为材料,通过已报道植物序列设计特异或兼并引物和参考苹果EST序列两种途径,利用RACE技术,分离得到2个PAL成员,分别为PpPAL1和PpPAL2;4个CHS成员,分别为PpCHS1、PpCHS2、PpCHS3和PpCHS4；3个CHI成员,分别为PpCHI1、PpCHI2和PpCHI3;1个PpF3Hl成员；2个DFR成员,分别为PpDFR1和PpDFR2;1个PpANS1成员；2个UFGT成员,分别为PpUFGT1和PpUFGT2;已分离转录因子PpMYBl和PpMYB10,其中PpMYB10属于第十亚组的R2R3MYB类群。
套袋显著改善红色砂梨果实着色。本研究以红色砂梨‘美人酥’套袋果为材料,研究商业采收前15d去袋后果面着色过程中花青苷合成相关基因表达模式。PpPAL1、PpCHSl、PpCHI1、PpF3H1、PpDFRl、PpANSl和PpUFGT1受到光照诱导表达量迅速上升,去袋后1d表达量均达到最大值,明显提前于花青苷的合成高峰。PpCHSl、PpF3H1、PpANSl和PpUFGT1在去袋后的前5d内都保持相对较高的表达水平。7个结构基因被光照诱导后表达量都迅速上升,说明合成途径中的结构基因通过协同作用参与花青苷的合成。光质和光强对果实着色都具有重要影响,同一个果实的阳面和阴面着色完全不同,在‘美人酥’的阴面果皮几乎不合成花青苷,定量结果显示阴面果皮中7个结构基因表达量都低于阳面果皮,其中PpCHS1、PpUFGT1、PpF3H1和PpANSl等4个基因在阳面果皮表达量的最大值(1DABR)与阴面的高峰(1DABR)比值超过2.7。PpCHS1、PpF3H1、 PpANS1和PpUFGT1等在成熟果实果皮中的表达均显著高于其它器官,同时PpCHS1与PpANS1在淡红色幼叶中的表达也较高。与成熟果中花青苷含量呈正相关关系。在红色砂梨‘美人酥’去袋着色过程中,PpMYB10与花青苷合成途径中的结构基因协同表达,而PpMYB1没有表现出相应的协同表达现象。
花青苷合成相关的结构基因家族成员在序列和表达模式上均存在较大差异。在氨基酸水平上,PpCHS1和PpCHS2的序列同源性高达97%,而PpCHI1和PpCHI2最低,仅有23%。红色砂梨‘云红梨1号’去袋后,花青苷合成途径中的结构基因部分成员被诱导,包括(?)PpPAL1、PpCHS1、PpCHS2、PpCHI1、PpCHI3、PpF3H1、 PpDFR1、PpANS1、PpUFGT1和PpUFGT2。而PpPAL2、PpCHS3、PpCHS4和PpCHI2的表达没有明显变化,由此推测这些基因可能不是果皮花青苷合成的关键基因。被诱导的基因中PpPAL1、PpCHS1、PpCHI1、PpCHI3、PpF3HI1、pDFR1、 PpANS1和PpUFGT1的表达高峰都出现在去袋后1d,另外两个基因PpCHS2和PpUFGT2的转录水平则在去袋后3d达到最大值,表现出不同的表达模式,这些结构基因可能通过协同作用共同调控花青苷的合成。
果实的着色受基因型决定,环境条件影响着色的程度。位于云南的安宁气候特点为亚热带高原气候,伴随着高海拔,强辐射和昼夜温差大等特点；河南的荥阳则为北温带大陆性季风气候,伴随着低海拔,低辐射和昼夜温差小等特点。本试验比较了红色砂梨‘满天红’在两个不同生态区套袋处理果实着色及品质相关生理分子研究。商业采收时,安宁较荥阳具有更好的着色,果皮中花青苷和类胡萝卜素含量较后者高,叶绿素含量较后者低；单果重和果实的硬度前者较后者低；前者果肉中可溶性固形物和糖含量较后者高,淀粉含量较后者低；着色过程中前者果皮花青苷合成结构和调节基因的表达量均较后者高。统计分析显示可溶性糖和花青苷含量在安宁(r=0.96981)和荥阳(r=0.94433)都显著相关。安宁具有的特定生态因子,特别是高海拔,强辐射和昼夜温差大等特点有利于果实中糖积累和诱导果皮中花青苷合成结构和调节基因,尤其是PpCHS1、PpANS1、 PpUFGT1、PpMYB10和PpbHLH的表达,促进了花青苷的合成。
Chinese sand pear (Pyrus pyrifolia Nakai) originated from China and is widely cultivated in East Asia. Ripe sand pears usually appear green, yellow or brown (actually is phellem). Red sand pear also could be found, but not as common as others. In recent years, red skin sand pear, such as'Meirensu','Mantianhong'and'Yunhongli No.1', were well received by customers, showing a great potential of custom market. However, bad color development of red Chinese sand pear greatly influenced the fruit appearance. The study on anthocyanin synthesis mechanism of red Chinese sand pear affected by fruit bagging treatments and different habitats will be of great help to develop a new technology to product high quality fruits.
Two red Chinese sand pear cultivars,'Meirensu'(mid-maturing) and'Yunhongli No.1'(late-maturing) in different maturation periods, were used as materials in this study. Two ways were used to isolate the gene sequences, one is used specific primers or degenerate primers which were designed according to plant sequences from available databases, the other is used specific primers based on malus EST database. Combined with RACE methods, two PALS (PpPALl and PpPAL2), four CHSS (PpCHSl, PpCHS2, PpCHS3and PpCHS4), three CHIS (PpCHI1, PpCHI2and PpCH13), one F3H (PpF3H1), two DFRs(PpDFRl and PpDFR2), one ANS (PpANSl) and two UFGTs(PpUFGT1and PpUFGT2) were isolated from red chinese sand pear. Transcription factors PpMYB1and PpMYB10were cloned, and sequence homology analysis showed PpMYB10belong to the subgroup10R2R3MYBs.
The coloration of red Chinese sand pears was significantly improved by fruit bagging treatments. Red Chinese sand pear'Meirensu'was used for fruit bagging treatments. Bags were removed15days prior to harvest and the fruits were completely re-exposed to light until harvest. Structural genes and transcription factors that involved in anthocyanin synthesis were analyzed during fruit coloration after the bags removal. Following bag removal, the expression levels of PpPAL1, PpCHSl, PpCHIl, PpF3Hl, PpDFR1, PpANSl and PpUFGTl were markedly enhanced in the sunlit side of 'Meirensu' fruit; PpCHS1, PpF3H1, PpANSl and PpUFGT1maintained high expression levels in5days after the bags removal (DABG). All seven structural genes were enhanced by sunlight, indicated both genes were involved in the anthocyanin biosynthesis. Light intensity and quality are important for fruit coloration. Anthocyanin amounts increased linearly in the skin of the sunlit side of'Meirensu' fruits. However, anthocyanin content was barely detected in the skin of the shaded side. The transcript level of anthocyanin biosynthetic genes was markedly up-regulated in the skin of sunlit side of'Meirensu', but much lower in the skin from the shaded side. The expression level of PpCHSl, PpUFGT1, PpF3H1and PpANSl (1DABR) in sunlit side as compared to shaded side is above2.7folds. PpCHS1, PpF3Hl, PpANSl and PpUFGTl expressed at higher level in the skin of mature fruits than in other tissues, and PpCHSl, PpANSl also showed high expression in young leaves with faint red. The expression level was coordinated with anthocyanin level in different organs. The expression pattern of PpMYB10was similar to other structural genes during the coloration of 'Meirensu' after the bags removal. However, PpMYB1showed less coordination with other genes.
There were family members in structural genes that involved in anthocyanin synthesis. They showed many differences in sequences and expression patterns. The highest percentage for amino acid sequences was found between PpCHSl and PpCHS2(97%), while the lowest one was PpCHIl and PpCHI2(23%). The expression levels of PpPAL1, PpCHSl, PpCHS2, PpCHI1, PpCHI3, PpF3Hl, PpDFR1, PpANS1, PpUFGT1and PpUFGT2was markedly enhanced in the peel of 'Yunhongli No.1'fruit after the bags removal. Other structural genes including PpPAL2, PpCHS3, PpCHS4and PpCHI2showed no response, indicating that they may not involved in anthocyanin synthesis during fruit coloration. The expression pattern of these genes that involved in anthocyanin synthesis shows different ways, transcript levels of PpPAL1, PpCHS1, PpCHI1, PpCHI3, PpF3H1, PpDFR1,PpANSl and PpUFGTl peaked at1DABR. However, PpCHS2and PpUFGT2peaked at3DABR. All these genes might be involved in anthocyanin biosynthesis by being coordinately expressed during anthocyanin accumulation.
The formation of anthocyanin was not only genetically determined, but also might be influenced by many environmental factors. Anning is located in Yunnan province, having a warm, high irradiant climate and high day and night temperature difference, while Xingyang is located in Henan province, with a hot, low irradiant climate and low day and night temperature difference. At harvest, red Chinese sand pear'Mantianhong'in Anning showed better coloration, higher concentrations of anthocyanin, carotenoid, total soluble solid (TSS) and soluble sugar but lower chlorophyll in pear peel, fruit weight, firmness and content of starch than those in Xingyang. Significantly increasing in the anthocyanin structural and regulatory genes, especially for PpCHS, PpANS, PpUFGT, PpMYB10and PpbHLH in red Chinese sand pears were found in materials sampled in Anning. The statistical analysis indicated that soluble sugar content was significantly associated with anthocyanin accumulation in both Anning (r=0.96981) and Xingyang (r=0.94433) habitats. Those results indicated high altitude, high-irradiant and high temperature differences in Anning habit were more conducive to the color development and inner quality of red Chinese sand pear, which was probably resulted in the higher concentration of sugar and expression of anthocyanin structural and regulatory genes, especially for the expression of PpCHS, PpANS, PpUFGT, PpMYB10and PpbHLH.
引文
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