中国原产完全甜柿自然脱涩机理研究
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摘要
完全甜柿不需人工脱涩即可脆食,较涩柿的商品价值更高,因而是柿育种主要目标。但完全甜柿资源稀少,分布狭窄,遗传一致度高,世界范围内遗传改良计划已面临近交退化的困境。近年来,在我国中部大别山区发现以‘罗田甜柿’为代表的新的完全甜柿类型,但其遗传背景以及作为完全甜柿育种亲本的价值尚不清楚。可溶性单宁是柿果实呈现涩味的主要物质。本研究以我国原产完全甜柿为试材,并与非完全甜柿和日本原产完全甜柿作比较,探讨单宁细胞发育模式以及单宁生物合成和凝固的关键基因表达及其与果实脱涩的关系,以期为完全甜柿自然脱涩机理的研究及其遗传改良提供科学依据。主要研究结果如下：
1.供试我国原产完全甜柿单宁细胞在果实发育早期(武汉地区7月10日左右)停止膨大,最大面积小于30×103μm2,单宁细胞内单宁积累持续到果实发育中后期(武汉地区9月10日左右),在果实发育整个过程中伴随着可溶性单宁向不溶性单宁的转化,且在果实发育后期(武汉地区9月到10月上旬)转化尤为明显。可能说明我国原产完全甜柿单宁细胞发育介于日本原产完全甜柿和非完全甜柿之间,且存在可溶性单宁的‘凝固’现象。
2.通过同源克隆和快速扩增cDNA末端(RACE)技术从‘罗田甜柿’(Diospyros kaki Thunb.'Luotian-tianshi',中国原产完全甜柿)幼果中扩增出长1379 bp包含ORF的无色花色素还原酶(leucoanthocyanidin reductase, LAR)基因cDNA全长序列,其编码349个氨基酸残基,命名为DkLAR。该序列有LAR蛋白所特有的THD、RFLP和ICCN保守序列和还原酶—异构酶—脱氢酶家族所特有的Rossmann二核苷酸结合位点。DkLAR表达水平变化趋势与我国原产完全甜柿单宁积累趋势相一致。
3.通过RACE和热不对称交错RT-PCR (TAIL-PCR)等技术从‘磨盘柿’(Diospyros kaki Thunb.'Mopanshi',中国原产完全涩柿)幼果中分离到4个丙酮酸脱羧酶(pyruvate decarboxylase, PDC)基因家族成员,分别命名为DkPDC1、DkPDC2、DkPDC3和DkPDC4;得到2个乙醇脱氢酶(alcohol dehydrogenase, ADH)基因家族成员,分别命名为DkADHl和DkADH2。通过qRT-PCR分析结果表明,在供试我国原产完全甜柿可溶性单宁向不溶性单宁主要转化时期,果肉中DkPDC2基因表达活性升高,DkADH1基因表达活性降低,维持在较低表达水平,可能与我国原产完全甜柿自然脱涩相关。
4乙醇处理‘罗田甜柿’、‘小果甜柿’(D. kaki Thunb.'Xiaoguo-tianshi',中国原产完全甜柿)、‘骏河’(D. kaki Thunb.'Suruga',日本原产完全甜柿)和‘禅寺丸’(D. kaki Thunb.'Zenjimaru',日本原产非完全甜柿)果实,除‘骏河’不受乙醇处理影响外其它材料果实6天内完成脱涩,且其可溶性单宁含量下降量和不溶性单宁含量上升量基本相等。DkADH和DkPDC基因在‘骏河’果皮、果肉和果心部位转录活性均极低,乙醇处理后其表达水平无明显变化,与乙醇处理对单宁含量无影响相一致。DkADH和DkPDC基因在‘罗田甜柿’果皮、果肉、果心和种子中表达活性均较低,但高于在‘骏河’中的表达水平,经乙醇处理后DkADH1和DkPDC2基因表达水平在种子部位明显升高而在果皮、果肉和果心部位的变化不明显,与乙醇处理后其可溶性单宁含量迅速下降和不溶性单宁含量迅速上升相一致。DkADH1和DkPDC2基因在‘小果甜柿’和‘禅寺丸’种子部位高效表达,乙醇处理后DkADH1基因在这两个品种的种子部位表达水平均明显下调,DkPDC2基因在‘小果甜柿’种子中表达活性升高,而在‘禅寺丸’种子中的转录活性影响较小,DkADH2和DkPDC1、DkPDC3和DkPDC4在‘小果甜柿’和‘禅寺丸’中总体表达水平较低,且无明显组织特异性,乙醇处理反应变化较小。可能说明与单宁‘凝固’有关的关键基因为DkADH1和DkPDC2,转录活性主要在种子部位,且自然脱涩过程中单宁‘凝固’能力大小顺序：‘禅寺丸’>‘小果甜柿’>‘罗田甜柿’。
综上所述,中国原产完全甜柿单宁细胞发育特点介于日本原产完全甜柿和非完全甜柿之间,其单宁细胞在果实发育早期停止膨大,单宁积累持续到果实发育中后期,且存在可溶性单宁向不溶性单宁转化过程。单宁生物合成关键基因DkLAR表达水平变化趋势与单宁细胞内单宁积累趋势相一致,单宁凝固相关基因DkPDC和DkADH两基因家族中DkPDC2和DkADH1基因表达模式与单宁凝固相关。因此,中国原产完全甜柿自然脱涩既有单宁细胞‘稀释效应’又有可溶性单宁的‘凝固作用’,且在凝固过程中DkPDC2和DkADH1基因起关键作用。
Pollination-constant and non-astringent (PCNA) fruit can be eaten like an apple without any postharvest treatment and is the most desirable for fresh consumption. So, it has been the main breeding objective. However, there are only a few cultivars of PCNA type, and they have very narrow genetic variability. The project of persimmon breeding is cornerd. Recentaly, a series of new PCNA types including'Luotian-tianshi' were found in Luotian country Hubei province. But it was not reported about genetic background and the value as parents for the breeding projects in persimmon. In this study, PCNA type origined in China was the main material, and non-PCNA and PCNA typesorigined in Japan were the materials as compared with Chinese PCNA types. The development pattern of tannin cell and the expreesion pattern of the genes related with tannin biosynthesis and tannin cogulation during the development of the fruit were studied, and its relationship with nature loss of astringency was analyzed. These studies would be provided scientific insights on the mechanism of the PCNA nature loss of astringency and genetic improvement in Japanese persimmon. The main results of this study are as follows:
1. Tannin cell size in Chinese PCNA type was not distinctly increased after July and the biggest area was smaller than 30×103μm2; the accumulation of tannin continued until September, and some tannin were coagulated during the stage of accumulation. These results suggested development pattern of tannin in Chinese PCNA type between Japanese PCNA and non-PCNA type.
2. The full length cDNA of LAR gene named DkLAR was cloned from the young fruit of'Luotian-tianshi'(Diospyros kaki Thunb., Chinese PCNA type) using degenerate PCR and Rapid amplication cDNA End (RACE) method, and registered in NCBI (GenBank accession number:EU747876). The 1356 bp long DkLAR cDNA contained a 1047 bp open reading frame (ORF) and encoded 349 amino acids. The DkLAR protein contained the THD, RFLP and ICCN motifs conserved in LAR protein, and include the Rossmann dinucleotide-binding domain. Expression pattern of DkLAR was coincident with the tannin cell development.
3. Four PDC cDNA fragments (named DkPDCl, DkPDC2, DkPDC3 and DkPDC4) and two full length cDNA of ADH gene (named DkADH1 and DkADH2) were cloned from the young fruit of 'Mopanshi' (Diospyros kaki Thunb., Chinese PCA type) using degenerate PCR, RACE and TAIL-PCR method. The expression patterns of DkPDC and DkADH were performed in the pulp of the fruit during the throughout of the fruit development. DkPDC2 expression level increased and DkADH1 expression level decrease at the late development stage of the fruit in which soluble tannin was converted into insoluble tannin. It suggested that the transcritive activity of DkPDC2 and DkADH1 was relatived with the PCNA nature loss astrigency.
4. The fruits of the'Luotian-tianshi' (Diospyros kaki Thunb., Chinese PCNA type), 'xiaoguo-tianshi' (Diospyros kaki Thunb., Chinese PCNA type),'Suruga' (Diospyros kaki Thunb., Japanese PCNA type) and'Zenjimaru' (Diospyros kaki Thunb., Japanese non-PCNA type) were enclosed in polyethylene bags containing small amount ethanol on the tree to remove astringency. The astringency of the fruits from all materials except 'Suruga' disappered completely in 6 days, and the decrease of soluble tannin were correlated with a increase of insoluble tannin in treated fruits. The expressions of DkPDC and DkADH in peel, pulp, core and seed of the'Suruga' were low and not related with treatment of ethanol. The expression pattern of DkPDC and DkADH gene in 'Luotian-tianshi' was similar to in'Suruga', but after treatment, DkPDC2 and DkADHl expression level in seed was increased significantly. In 'Xiaoguo-tianshi' and 'Zenjimaru', the expreesion level of DkADH1 was very high, but decreased after treatment using ethanol; the DkPDC2 expression was high in seeds of the'Xiaoguo-tianshi' and 'Zenjimaru', after treatment using ethanol, increased in'Xiaoguo-tianshi' and changed little in'Zenjimaru'. DkADH2, DkPDCl, DkPDC3 and DkPDC4 expression level was low and no specific in pulp, peel, core and seed in 'Xiaoguo-tianshi' and 'Zenjimaru'. These results suggested that DkPDC2 and DkADHl gene in seeds played an important role in tannin congulation, and the order of ability of removing astringency during nature loss of astringency was'Zenjimaru'>'Xiaoguo-tianshi'>'Luotian-tianshi'.
1.供试我国原产完全甜柿单宁细胞在果实发育早期(武汉地区7月10日左右)停止膨大,最大面积小于30×103μm2,单宁细胞内单宁积累持续到果实发育中后期(武汉地区9月10日左右),在果实发育整个过程中伴随着可溶性单宁向不溶性单宁的转化,且在果实发育后期(武汉地区9月到10月上旬)转化尤为明显。可能说明我国原产完全甜柿单宁细胞发育介于日本原产完全甜柿和非完全甜柿之间,且存在可溶性单宁的‘凝固’现象。
2.通过同源克隆和快速扩增cDNA末端(RACE)技术从‘罗田甜柿’(Diospyros kaki Thunb.'Luotian-tianshi',中国原产完全甜柿)幼果中扩增出长1379 bp包含ORF的无色花色素还原酶(leucoanthocyanidin reductase, LAR)基因cDNA全长序列,其编码349个氨基酸残基,命名为DkLAR。该序列有LAR蛋白所特有的THD、RFLP和ICCN保守序列和还原酶—异构酶—脱氢酶家族所特有的Rossmann二核苷酸结合位点。DkLAR表达水平变化趋势与我国原产完全甜柿单宁积累趋势相一致。
3.通过RACE和热不对称交错RT-PCR (TAIL-PCR)等技术从‘磨盘柿’(Diospyros kaki Thunb.'Mopanshi',中国原产完全涩柿)幼果中分离到4个丙酮酸脱羧酶(pyruvate decarboxylase, PDC)基因家族成员,分别命名为DkPDC1、DkPDC2、DkPDC3和DkPDC4;得到2个乙醇脱氢酶(alcohol dehydrogenase, ADH)基因家族成员,分别命名为DkADHl和DkADH2。通过qRT-PCR分析结果表明,在供试我国原产完全甜柿可溶性单宁向不溶性单宁主要转化时期,果肉中DkPDC2基因表达活性升高,DkADH1基因表达活性降低,维持在较低表达水平,可能与我国原产完全甜柿自然脱涩相关。
4乙醇处理‘罗田甜柿’、‘小果甜柿’(D. kaki Thunb.'Xiaoguo-tianshi',中国原产完全甜柿)、‘骏河’(D. kaki Thunb.'Suruga',日本原产完全甜柿)和‘禅寺丸’(D. kaki Thunb.'Zenjimaru',日本原产非完全甜柿)果实,除‘骏河’不受乙醇处理影响外其它材料果实6天内完成脱涩,且其可溶性单宁含量下降量和不溶性单宁含量上升量基本相等。DkADH和DkPDC基因在‘骏河’果皮、果肉和果心部位转录活性均极低,乙醇处理后其表达水平无明显变化,与乙醇处理对单宁含量无影响相一致。DkADH和DkPDC基因在‘罗田甜柿’果皮、果肉、果心和种子中表达活性均较低,但高于在‘骏河’中的表达水平,经乙醇处理后DkADH1和DkPDC2基因表达水平在种子部位明显升高而在果皮、果肉和果心部位的变化不明显,与乙醇处理后其可溶性单宁含量迅速下降和不溶性单宁含量迅速上升相一致。DkADH1和DkPDC2基因在‘小果甜柿’和‘禅寺丸’种子部位高效表达,乙醇处理后DkADH1基因在这两个品种的种子部位表达水平均明显下调,DkPDC2基因在‘小果甜柿’种子中表达活性升高,而在‘禅寺丸’种子中的转录活性影响较小,DkADH2和DkPDC1、DkPDC3和DkPDC4在‘小果甜柿’和‘禅寺丸’中总体表达水平较低,且无明显组织特异性,乙醇处理反应变化较小。可能说明与单宁‘凝固’有关的关键基因为DkADH1和DkPDC2,转录活性主要在种子部位,且自然脱涩过程中单宁‘凝固’能力大小顺序：‘禅寺丸’>‘小果甜柿’>‘罗田甜柿’。
综上所述,中国原产完全甜柿单宁细胞发育特点介于日本原产完全甜柿和非完全甜柿之间,其单宁细胞在果实发育早期停止膨大,单宁积累持续到果实发育中后期,且存在可溶性单宁向不溶性单宁转化过程。单宁生物合成关键基因DkLAR表达水平变化趋势与单宁细胞内单宁积累趋势相一致,单宁凝固相关基因DkPDC和DkADH两基因家族中DkPDC2和DkADH1基因表达模式与单宁凝固相关。因此,中国原产完全甜柿自然脱涩既有单宁细胞‘稀释效应’又有可溶性单宁的‘凝固作用’,且在凝固过程中DkPDC2和DkADH1基因起关键作用。
Pollination-constant and non-astringent (PCNA) fruit can be eaten like an apple without any postharvest treatment and is the most desirable for fresh consumption. So, it has been the main breeding objective. However, there are only a few cultivars of PCNA type, and they have very narrow genetic variability. The project of persimmon breeding is cornerd. Recentaly, a series of new PCNA types including'Luotian-tianshi' were found in Luotian country Hubei province. But it was not reported about genetic background and the value as parents for the breeding projects in persimmon. In this study, PCNA type origined in China was the main material, and non-PCNA and PCNA typesorigined in Japan were the materials as compared with Chinese PCNA types. The development pattern of tannin cell and the expreesion pattern of the genes related with tannin biosynthesis and tannin cogulation during the development of the fruit were studied, and its relationship with nature loss of astringency was analyzed. These studies would be provided scientific insights on the mechanism of the PCNA nature loss of astringency and genetic improvement in Japanese persimmon. The main results of this study are as follows:
1. Tannin cell size in Chinese PCNA type was not distinctly increased after July and the biggest area was smaller than 30×103μm2; the accumulation of tannin continued until September, and some tannin were coagulated during the stage of accumulation. These results suggested development pattern of tannin in Chinese PCNA type between Japanese PCNA and non-PCNA type.
2. The full length cDNA of LAR gene named DkLAR was cloned from the young fruit of'Luotian-tianshi'(Diospyros kaki Thunb., Chinese PCNA type) using degenerate PCR and Rapid amplication cDNA End (RACE) method, and registered in NCBI (GenBank accession number:EU747876). The 1356 bp long DkLAR cDNA contained a 1047 bp open reading frame (ORF) and encoded 349 amino acids. The DkLAR protein contained the THD, RFLP and ICCN motifs conserved in LAR protein, and include the Rossmann dinucleotide-binding domain. Expression pattern of DkLAR was coincident with the tannin cell development.
3. Four PDC cDNA fragments (named DkPDCl, DkPDC2, DkPDC3 and DkPDC4) and two full length cDNA of ADH gene (named DkADH1 and DkADH2) were cloned from the young fruit of 'Mopanshi' (Diospyros kaki Thunb., Chinese PCA type) using degenerate PCR, RACE and TAIL-PCR method. The expression patterns of DkPDC and DkADH were performed in the pulp of the fruit during the throughout of the fruit development. DkPDC2 expression level increased and DkADH1 expression level decrease at the late development stage of the fruit in which soluble tannin was converted into insoluble tannin. It suggested that the transcritive activity of DkPDC2 and DkADH1 was relatived with the PCNA nature loss astrigency.
4. The fruits of the'Luotian-tianshi' (Diospyros kaki Thunb., Chinese PCNA type), 'xiaoguo-tianshi' (Diospyros kaki Thunb., Chinese PCNA type),'Suruga' (Diospyros kaki Thunb., Japanese PCNA type) and'Zenjimaru' (Diospyros kaki Thunb., Japanese non-PCNA type) were enclosed in polyethylene bags containing small amount ethanol on the tree to remove astringency. The astringency of the fruits from all materials except 'Suruga' disappered completely in 6 days, and the decrease of soluble tannin were correlated with a increase of insoluble tannin in treated fruits. The expressions of DkPDC and DkADH in peel, pulp, core and seed of the'Suruga' were low and not related with treatment of ethanol. The expression pattern of DkPDC and DkADH gene in 'Luotian-tianshi' was similar to in'Suruga', but after treatment, DkPDC2 and DkADHl expression level in seed was increased significantly. In 'Xiaoguo-tianshi' and 'Zenjimaru', the expreesion level of DkADH1 was very high, but decreased after treatment using ethanol; the DkPDC2 expression was high in seeds of the'Xiaoguo-tianshi' and 'Zenjimaru', after treatment using ethanol, increased in'Xiaoguo-tianshi' and changed little in'Zenjimaru'. DkADH2, DkPDCl, DkPDC3 and DkPDC4 expression level was low and no specific in pulp, peel, core and seed in 'Xiaoguo-tianshi' and 'Zenjimaru'. These results suggested that DkPDC2 and DkADHl gene in seeds played an important role in tannin congulation, and the order of ability of removing astringency during nature loss of astringency was'Zenjimaru'>'Xiaoguo-tianshi'>'Luotian-tianshi'.
引文
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