红色砂梨着色诱导及其调控的分子生理机制
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摘要
红色砂梨(Pyrus pyrifolia Nakai)原产中国云南和四川南部,近年来,以原产云南著名红色砂梨地方品种‘火把梨’为亲本育成的优良品种已在全国各地引种栽培,得到了消费者的广泛认可,市场前景广阔。红色砂梨的相关研究多集中于着色机理上,着色调控方面的研究相对较少,难以满足生产上优质红色砂梨生产的需求。此外,本课题组的前期研究发现红色砂梨不同于西洋梨和苹果的着色规律,因此有必要对红色砂梨着色调控及其相关分子生理机制进行系统研究,为红色砂梨生产中的着色调控技术的开发提供理论支持。本研究以红色砂梨中熟品种‘满天红’和‘美人酥’,晚熟品种‘云红梨1号’为主要试材,系统研究了温度、光照、机械伤和化学药剂等试验处理对红色砂梨着色调控的效应及相关分子生理机制,同时探讨了采收成熟度对诱导红色砂梨着色的影响及相关分子生理机制,主要研究成果如下：
1、UV-B/白光诱导条件下,高温(27℃)诱导红色砂梨品种‘云红梨1号’果皮中花青苷积累的效应显著优于低温(17℃),这可能缘于高温较低温果皮具有更高的PAL酶活性以及(?)yMYB10和结构基因PpPAL、PpCHS、PpANS和PpUFGT的表达水平。
2、红色砂梨对UV-B/白光诱导的响应与苹果较为相似,而与西洋梨有显著差异。不同红色砂梨品种的最佳着色温度和可诱导着色的温度范围均有所不同,红色砂梨‘满天红’和‘美人酥’最佳着色温度分别为17℃和12℃。光强和光质都对红色砂梨着色具有重要影响,高光强有利于红色砂梨着色,UV-B和白光在红色砂梨果皮花青苷积累方面具有显著地互作增益效应。基于本研究的结果,我们认为低温和高强度的UV-B/白光环境有利于获得较佳着色的‘满天红’和‘美人酥’,同时可较好的维持果实内在品质。
3、机械伤处理能够有效促进‘满天红’果实在UV-B/白光诱导条件下受伤部位花青苷的积累,并伴随着该部位果实硬度显著下降,乙烯释放速率显著上升,花色苷合成相关基因PpF3H、PpANS、PpUFGT和PyMYB10表达量的显著上调,而对照果实仅发生了轻微着色,且未检测到乙烯的释放。推测乙烯可能参与了红色砂梨果实花色苷合成的调控,对花色苷合成具有促进作用。机械伤处理可以作为一种有效的研究方法用于果实着色调控机制的研究。
4、采收成熟度对诱导红色砂梨着色效应具有重要影响,且这种影响会随着果实临近商业采收期而减弱。成熟度较高果实中更高的花青苷的积累,可能与其果实具有更高的可溶性糖含量以及花青苷相关结构和调节基因尤其是PpCHS, PpF3H, PpANS, PpUFGT, PyMYB10和PpbHLH等的高量表达有关。商业成熟期前10d是红色砂梨采后诱导着色的最佳采收期。
5、在UV-B/白光的诱导条件下,不同药剂处理对红色砂梨‘满天红’着色及主要果实品质的调控效应不同。0.4m mol·L-1ABA和3mmol·L-1GA对‘满天红’果皮花青苷合成没有增效；2m mol·L-1ETH,0.12m mol·L-1GNT和2m mol·L-1MJ处理以及它们混合使用均能促进‘满天红’果皮花青苷著合成,ETH和MJ间还存在显著地互作增益现象,这些处理还能较好地维持果实硬度、可溶性固形物、可溶性糖和有机酸的含量等。
6、MJ处理显著提高了红色砂梨‘满天红’果皮中PpCHS2、PpCHI3、PpF3H, PpANS、PpDFR1、PpUFGT1、PpUFGT2和PyMYB10的表达水平,从而促进其果皮中花青苷的积累。曝光处理后,PpbHLH和PpWD40表达在对照中不受诱导,而在MJ处理中出现上调表达现象,推测这两个基因的上调表达也是MJ促进花青苷积累的原因之一。红色砂梨花青苷合成过程中,花青苷合成结构基因不同成员的表达模式和相对表达量的变化存在显著差异,这些成员的功能可能有所不同。
Red Chinese sand pears (Pyrus prifolia Nakai) native to Yunnan Province and southern part of Sichuan Province, Recently, with extensive cultivation of red Chinese sand pear cultivars newly released by hybrid breeding in China, the pears had been widely accepted by consumers, and are expected to be of value of Chinese production in the future. Poor fruit coloration has been a noticeable problem, however, there are short of the technique of coloration regulation on production. To date, little studies have been carried on coloration regulation of red Chinese sand pears on scientific research. What's more, Chinese sand pears and European pears(P. communis L.) differ considerably in pigmentation pattern. Therefore, an investigation of molecular physiological mechanism of coloration regulation of red Chinese sand pears need to be done, to improve the coloration of red Chinese sand pears. Three red Chinese sand pear cultivars,'Meirensu'(mid-maturing),'Mantianhong'(mid-maturing) and 'Yunhongli No.l'(late-maturing) were used as materials in this study. Color response and related molecular physiological mechanism of coloration regulation of red Chinese sand pears to post-harvest induced is influenced by temperature, light, maturity, wounded, chemicals was investigated systematically. The main results are as follows:
The efficacy of UV-B/visible irradiation was evaluated under high-(27℃) and low-temperature (17℃) conditions using red Chinese sand pear'Yunhongli No.1'as a model. The results showed that UV-B/visible irradiation was more effective in inducing anthocyanin synthesis in peel tissues and improving fruit coloration at27℃than at17℃. PAL activity was markedly higher at27℃than at17℃. Expression of PyMYB10and five anthocyanin structural genes, PpPAL, PpCHI, PpCHS, PpF3H, and PpANS, were also higher in fruit irradiated at27℃than in fruit irradiated at17℃. For PpUFGT, transcription reached a maximum at48and240hours after the onset of irratiation at27℃and17℃, respetively, but the peak value was lower at27℃than at 17℃. These results indicated that high temperatures (27℃) enhanced UV-B/visible irradiation induced postharvest anthocyanin accumulation in'Yunhongli No.1'pears by up-regulating PyMYB10and anthocyanin structural genes and increasing the activity of phenylalanine ammonia-lyase.
Color response of red Chinese sand pears to different temperatures treatment similar to'Gala' apple (Malus domestic Borkh), but differ considerably to European pears under postharvest artificial irradiation conditions. The optimum temperature of 'Mantianhong; and'Meirensu'was17℃and12℃, respectively. Light intensity (quantity) and light wavelength (quality) are important factors affecting anthocyanin synthesis in red Chinese sand pears. Higher light intensity was good for anthocyanin synthesis in red Chinese sand pears. Simultaneous irradiation with UV-B and white light had a synergistic effect. Based on the current results, we suggested that low temperature and high light intensity was good for red color development and obtaining fruit quality of'Mantianhong'and'Meirensu'
Wounded treatments decreased fruit firmness, increased ethylene production rate of fruit, enhanced post-harvest anthocyanin accumulation in injured part of red Chinese sand pears, and up-regulated the transcription level of anthocyanin biosynthetic related genes:PpF3H, PpANS, PpUFGT and PyMYB10, markedly. Ethylene production of control fruit was under the detection limit, and only a trace amount of anthocyanin was detected in control fruit. Results above indicated that ethylene might be involved in the regulation of anthocyanin accumulation in red Chinese sand pear, and improve the level of anthocyanin accumulation in red Chinese sand pear. It was suggested that wounded treatments could be used as a mean to better understand the regulation mechanism of anthocyanin biosynthesis in fruits.
Color response of'Mantianhong'pears to postharvest UV-B irradiation was affected by fruit maturity, and the effect of maturity on accumulation of anthocyanin became smaller near the commercial harvesting period. Higher accumulation of anthocyanin in mature fruits than in immature fruits resulted from significant increase of the anthocyanin structural and regulatory genes, especially for PpCHS, PpF3H, PpANS, PpUFGT, PyMYB10and PpbHLH in red Chinese sand pears. Based on the results above, we suggested that10DBH harvest was the optimal maturity for UV-B-Induced post-harvest anthocyanin accumulation of red Chinese sand pear.
Different chemicals has different efficiency in coloration induced and fruit quality regulating. No positive efficiency was observed in0.4m mol·L-1ABA and3m mol·L-1GA treatment in inducing anthocyanin accumulation of'Mantianhong'. However, alone2m mol·L-1ETH,0.12m mol· L-1GNT,2m mol·L-1MJ and mixture of these chemicals enhanced anthocyanin accumulation in red Chinese sand pear. Simultaneous irradiation with MJ and ETH had a synergistic effect in inducing anthocyanin accumulation of'Mantianhong'. All these treatments can obtain fruit quality of'Mantianhong'pear, such as fruit firmness, TSS, soluble sugar and organic acid.
MJ treatment enhanced anthocyanin accumulation in peel of'Mantianhong'pear by up-regulating the transcription level of anthocyanin biosynthetic related genes: PpCHS2, PpCHI3, PpF3H, PpANS, PpDFRl, PpUFGT1PpUFGT2and PyMYB10O, markedly. After irradiation, PpbHLH and PpWD40were up-regulated in MJ treatment, but not in control. We suggested that higher transcription level of PpbHLH and PpWD40in MJ treatment than control is one of the reasons of enhancing anthocyanin accumulation in peel of'Mantianhong'pear. During anthocyanin accumulation of red Chinese sand pear, the transcription level of different anthocyanin structural gene family members was significantly different, and we presumed that the function of these genes was different.
1、UV-B/白光诱导条件下,高温(27℃)诱导红色砂梨品种‘云红梨1号’果皮中花青苷积累的效应显著优于低温(17℃),这可能缘于高温较低温果皮具有更高的PAL酶活性以及(?)yMYB10和结构基因PpPAL、PpCHS、PpANS和PpUFGT的表达水平。
2、红色砂梨对UV-B/白光诱导的响应与苹果较为相似,而与西洋梨有显著差异。不同红色砂梨品种的最佳着色温度和可诱导着色的温度范围均有所不同,红色砂梨‘满天红’和‘美人酥’最佳着色温度分别为17℃和12℃。光强和光质都对红色砂梨着色具有重要影响,高光强有利于红色砂梨着色,UV-B和白光在红色砂梨果皮花青苷积累方面具有显著地互作增益效应。基于本研究的结果,我们认为低温和高强度的UV-B/白光环境有利于获得较佳着色的‘满天红’和‘美人酥’,同时可较好的维持果实内在品质。
3、机械伤处理能够有效促进‘满天红’果实在UV-B/白光诱导条件下受伤部位花青苷的积累,并伴随着该部位果实硬度显著下降,乙烯释放速率显著上升,花色苷合成相关基因PpF3H、PpANS、PpUFGT和PyMYB10表达量的显著上调,而对照果实仅发生了轻微着色,且未检测到乙烯的释放。推测乙烯可能参与了红色砂梨果实花色苷合成的调控,对花色苷合成具有促进作用。机械伤处理可以作为一种有效的研究方法用于果实着色调控机制的研究。
4、采收成熟度对诱导红色砂梨着色效应具有重要影响,且这种影响会随着果实临近商业采收期而减弱。成熟度较高果实中更高的花青苷的积累,可能与其果实具有更高的可溶性糖含量以及花青苷相关结构和调节基因尤其是PpCHS, PpF3H, PpANS, PpUFGT, PyMYB10和PpbHLH等的高量表达有关。商业成熟期前10d是红色砂梨采后诱导着色的最佳采收期。
5、在UV-B/白光的诱导条件下,不同药剂处理对红色砂梨‘满天红’着色及主要果实品质的调控效应不同。0.4m mol·L-1ABA和3mmol·L-1GA对‘满天红’果皮花青苷合成没有增效；2m mol·L-1ETH,0.12m mol·L-1GNT和2m mol·L-1MJ处理以及它们混合使用均能促进‘满天红’果皮花青苷著合成,ETH和MJ间还存在显著地互作增益现象,这些处理还能较好地维持果实硬度、可溶性固形物、可溶性糖和有机酸的含量等。
6、MJ处理显著提高了红色砂梨‘满天红’果皮中PpCHS2、PpCHI3、PpF3H, PpANS、PpDFR1、PpUFGT1、PpUFGT2和PyMYB10的表达水平,从而促进其果皮中花青苷的积累。曝光处理后,PpbHLH和PpWD40表达在对照中不受诱导,而在MJ处理中出现上调表达现象,推测这两个基因的上调表达也是MJ促进花青苷积累的原因之一。红色砂梨花青苷合成过程中,花青苷合成结构基因不同成员的表达模式和相对表达量的变化存在显著差异,这些成员的功能可能有所不同。
Red Chinese sand pears (Pyrus prifolia Nakai) native to Yunnan Province and southern part of Sichuan Province, Recently, with extensive cultivation of red Chinese sand pear cultivars newly released by hybrid breeding in China, the pears had been widely accepted by consumers, and are expected to be of value of Chinese production in the future. Poor fruit coloration has been a noticeable problem, however, there are short of the technique of coloration regulation on production. To date, little studies have been carried on coloration regulation of red Chinese sand pears on scientific research. What's more, Chinese sand pears and European pears(P. communis L.) differ considerably in pigmentation pattern. Therefore, an investigation of molecular physiological mechanism of coloration regulation of red Chinese sand pears need to be done, to improve the coloration of red Chinese sand pears. Three red Chinese sand pear cultivars,'Meirensu'(mid-maturing),'Mantianhong'(mid-maturing) and 'Yunhongli No.l'(late-maturing) were used as materials in this study. Color response and related molecular physiological mechanism of coloration regulation of red Chinese sand pears to post-harvest induced is influenced by temperature, light, maturity, wounded, chemicals was investigated systematically. The main results are as follows:
The efficacy of UV-B/visible irradiation was evaluated under high-(27℃) and low-temperature (17℃) conditions using red Chinese sand pear'Yunhongli No.1'as a model. The results showed that UV-B/visible irradiation was more effective in inducing anthocyanin synthesis in peel tissues and improving fruit coloration at27℃than at17℃. PAL activity was markedly higher at27℃than at17℃. Expression of PyMYB10and five anthocyanin structural genes, PpPAL, PpCHI, PpCHS, PpF3H, and PpANS, were also higher in fruit irradiated at27℃than in fruit irradiated at17℃. For PpUFGT, transcription reached a maximum at48and240hours after the onset of irratiation at27℃and17℃, respetively, but the peak value was lower at27℃than at 17℃. These results indicated that high temperatures (27℃) enhanced UV-B/visible irradiation induced postharvest anthocyanin accumulation in'Yunhongli No.1'pears by up-regulating PyMYB10and anthocyanin structural genes and increasing the activity of phenylalanine ammonia-lyase.
Color response of red Chinese sand pears to different temperatures treatment similar to'Gala' apple (Malus domestic Borkh), but differ considerably to European pears under postharvest artificial irradiation conditions. The optimum temperature of 'Mantianhong; and'Meirensu'was17℃and12℃, respectively. Light intensity (quantity) and light wavelength (quality) are important factors affecting anthocyanin synthesis in red Chinese sand pears. Higher light intensity was good for anthocyanin synthesis in red Chinese sand pears. Simultaneous irradiation with UV-B and white light had a synergistic effect. Based on the current results, we suggested that low temperature and high light intensity was good for red color development and obtaining fruit quality of'Mantianhong'and'Meirensu'
Wounded treatments decreased fruit firmness, increased ethylene production rate of fruit, enhanced post-harvest anthocyanin accumulation in injured part of red Chinese sand pears, and up-regulated the transcription level of anthocyanin biosynthetic related genes:PpF3H, PpANS, PpUFGT and PyMYB10, markedly. Ethylene production of control fruit was under the detection limit, and only a trace amount of anthocyanin was detected in control fruit. Results above indicated that ethylene might be involved in the regulation of anthocyanin accumulation in red Chinese sand pear, and improve the level of anthocyanin accumulation in red Chinese sand pear. It was suggested that wounded treatments could be used as a mean to better understand the regulation mechanism of anthocyanin biosynthesis in fruits.
Color response of'Mantianhong'pears to postharvest UV-B irradiation was affected by fruit maturity, and the effect of maturity on accumulation of anthocyanin became smaller near the commercial harvesting period. Higher accumulation of anthocyanin in mature fruits than in immature fruits resulted from significant increase of the anthocyanin structural and regulatory genes, especially for PpCHS, PpF3H, PpANS, PpUFGT, PyMYB10and PpbHLH in red Chinese sand pears. Based on the results above, we suggested that10DBH harvest was the optimal maturity for UV-B-Induced post-harvest anthocyanin accumulation of red Chinese sand pear.
Different chemicals has different efficiency in coloration induced and fruit quality regulating. No positive efficiency was observed in0.4m mol·L-1ABA and3m mol·L-1GA treatment in inducing anthocyanin accumulation of'Mantianhong'. However, alone2m mol·L-1ETH,0.12m mol· L-1GNT,2m mol·L-1MJ and mixture of these chemicals enhanced anthocyanin accumulation in red Chinese sand pear. Simultaneous irradiation with MJ and ETH had a synergistic effect in inducing anthocyanin accumulation of'Mantianhong'. All these treatments can obtain fruit quality of'Mantianhong'pear, such as fruit firmness, TSS, soluble sugar and organic acid.
MJ treatment enhanced anthocyanin accumulation in peel of'Mantianhong'pear by up-regulating the transcription level of anthocyanin biosynthetic related genes: PpCHS2, PpCHI3, PpF3H, PpANS, PpDFRl, PpUFGT1PpUFGT2and PyMYB10O, markedly. After irradiation, PpbHLH and PpWD40were up-regulated in MJ treatment, but not in control. We suggested that higher transcription level of PpbHLH and PpWD40in MJ treatment than control is one of the reasons of enhancing anthocyanin accumulation in peel of'Mantianhong'pear. During anthocyanin accumulation of red Chinese sand pear, the transcription level of different anthocyanin structural gene family members was significantly different, and we presumed that the function of these genes was different.
引文
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