砂梨果皮性状形成机制研究
摘要
梨果皮性状主要由表皮色泽、果点大小与密度、果面蜡质、表皮锈斑等因素构成。早熟砂梨主栽品种‘翠冠’果实表面易发生果锈,外观品质受到严重影响。为揭示砂梨果皮性状形成机制,本研究以‘翠冠’等不同皮色砂梨品种为试材,研究了果实生长发育过程中果皮色素含量的动态变化特点、果实表皮组织显微和超微结构特征；利用BSA分类法筛选与果实皮色相关基因连锁的AFLP标记；探讨了苹果基因序列在砂梨同源基因克隆中的应用,在此基础上开展了砂梨果实木质素形成调控分子基础的初步研究,主要结果如下：
1、以‘西子绿’和‘清香’为试材,对乙醇和丙酮2种有机溶剂浸提法提取果皮和果肉中叶绿素及类胡萝卜素的效果进行了比较分析,结果表明：80%丙酮浸提法和95%乙醇浸提法的效果相似,虽然应用95%乙醇浸提法的各种色素测定结果略偏低,但除了叶绿素a之外,2种提取方法间的差异不显著。并发现4个绿皮品种和4个褐皮品种果实发育过程中果实表皮及果肉中叶绿素、类胡萝卜素的含量与果实皮色表型无关。同时,以‘翠冠’与‘玉冠’为试材,测定了果实发育进程中纵横径、果皮与果肉色素含量的动态变化,以及在成熟前期可滴定酸、可溶性固形物、糖含量的变化特性。结果显示,两品种果实膨大过程均呈单“s”型曲线动态变化。在成熟前14d(翠冠)、28d(玉冠)至采收,果实糖、酸含量急剧变化,且两品种间的变化特性不同,生产上应依据不同品种果实糖、酸形成特性,确定适宜采收时期。
2、以绿皮品种‘翠冠’、‘翠玉’与褐皮品种‘清香’、‘玉冠’为试材,进行了果实表皮组织结构显微和超微水平观察,结果表明：避雨栽培能显著减小‘翠冠’果点的大小,而对‘翠玉’的果点大小没有显著影响。‘翠冠’果实表皮细胞的形状和排列方式与‘翠玉’不同,且角质膜分布厚薄不均,厚度仅为‘翠玉’的一半；两品种果实表皮的角质膜均出现破裂,并形成网状结构,但‘翠玉’龟裂产生的裂纹均匀且浅,而‘翠冠’龟裂产生的裂纹紊乱,且深及表皮层内部。‘玉冠’和‘清香’的果点形成时期要比‘翠冠’和‘翠玉’早50d左右。‘翠冠’果锈是由单层木栓化的表皮细胞构成,‘玉冠’褐皮则由多层木栓化的细胞构成,上层细胞片状分布。表皮细胞超微结构观察结果显示,在角质膜下第1-2层细胞的细胞质稠密,且含丰富的脂质体,脂质体内含圆形小脂滴。这一结构特征表明,梨果实表皮角质膜下第1-2层细胞具有旺盛的分泌蜡脂的能力,但品种间存在差异,‘翠玉’脂质体中的脂滴密集,而‘翠冠’中脂滴稀少。研究还表明,‘翠冠’的果锈形成期主要在果实发育中后期；水分是诱发果锈形成的主要外因。
3、以‘秋荣’ב初夏绿’的杂交后代群体为试材,通过64对AFLP标记引物在亲本和分离群体中的筛选和验证,获得与梨果实皮色相关基因连锁的标记2个,即E-AAG/M-CAG和E-ACT/M-CTT,前者在74%的非纯绿材料中扩增出目标带,在80%的纯绿皮材料中表现为缺失带型；在89%的非纯褐材料中扩增出了目标带,后者在77%的纯褐材料中表现为缺失带型。
4、选取苹果3种类型的8个基因(液泡质子泵焦磷酸水解酶基因、液泡质子泵三磷酸腺苷酶A亚基基因、生长素受体基因、脱落酸受体基因、乙烯受体基因、果重基因、抗病相关基因),分别在起始密码子上游区、终止密码子下游区、外显子区和内含子区设计引物,分别对苹果和砂梨的基因组DNA进行PCR扩增。结果显示,除了抗病相关基因,其余的7个基因均有引物在砂梨基因组DNA中扩增出特异条带。与基因其它区域扩增产物相比,起始密码子上游区扩增产物在不同砂梨材料间多态性较丰富,外显子和内含子区扩增产物在不同砂梨材料间的保守性更高。外显子区扩增产物测序结果表明：砂梨和苹果在该区对应基因片段具有89.6%-97.5%的核苷酸序列同源性。应用同源克隆方法从砂梨(Pyrus pyrifolia)中获得4个长度分别为821bp,874bp,972bp和914bp木质素合成调控因子MYB基因的启动子序列,GenBank登录号分别为JF900710、JF900709、JF900707和JF900708。生物信息学预测发现,这些序列具有典型的启动子序列特征。各启动子序列在果皮褐与绿的砂梨品种间的等位变异分析发现,PpMYBx2启动子区的一个SNP (A/G)位点与砂梨果皮褐、绿色性状间存在相关性。本研究为进一步开展砂梨MYB基因的表达分析、遗传定位,以及探明砂梨果实木质素形成的分子机制奠定了基础。
Appearance quality of pear fruits is mainly composed of skin color, size and density of fruit dot, the fruit surface waxiness, epidermal rust and other factors. The sand pear 'Cuiguan' is a cultivar with early maturity. It usually generates surface rust, which largely damages the fruit appearance quality. In order to reveal the mechanism of the appearance quality formation, we used 'Cuiguan' and other cultivars with different fruit peel color as materials and studied fruit peel pigment content dynamic changes and microscopic and ultrastructural features of epidermal tissue during different developmental stages of fruits; screened the AFLP markers linkage with fruit color genes using BSA classification; analyzed the application of apple fruit peel color related genes in pear homologues cloning, and based on which the preliminary study of molecular regulation of sand pear lignin biosynthesis was carried out. The major results are as follows:
1. Using 'Xizilv' and 'Qingxiang' as materials, we compared the efficiency of ethanol and acetone in chlorophyll and carotenoid extracting in peel and pulp. The results showed that the extracting efficiency of80%acetone is nearly similar with that of95%ethanol. Compared with80%acetone,95%ethanol resulted slightly lower pigment extraction, but there was no significant difference in the statistic analysis (except for chlorophyll a). The comparison of chlorophyll and carotenoid content in peel and pulp between the four green and four russet varieties at different development stages showed that the content of the pigments has no relation with the fruit skin color phenotype.'Cuiguan' and 'Yuguan' were used as materials to measure the dynamics changes of fruit vertical and horizontal diameter, pigments content in peel and pulp during the fruit development, and to measure titratable acid, soluble solids, and sugar contents during fruit maturing and harvest stage. The results showed that the development of fruit size of two varieties presented "S" curve; there was no significant differences of total chlorophyll and carotenoid content in peel and pulp between russet and green pear. During maturation (14days before harvest of'Cuiguan',28days before harvest of'Yuguan') and havest stage, fruit growth rate decreased, and the contents of titratable acid and sugars dramatically increased. The properties of these were difference between the two cultivars. Different harvest date should be determined according to the character of fruit inner quality formation for certain cultivar.
2. The epidermis structure observation was carried out at microscopic and ultra-microscopic levels using green-skin fruit sand pear cultivars'Cuiguan' and 'Cuiyu' and russet-skin fruit sand pear cultivars 'Qingxiang' and 'Yuguan'. The results showed that rain shelter cultivation could significantly reduce'Cuiguan'fruit dot size, while the'Cuiyu 'the fruit dot size does not significantly affect.the epidermal cell shape and arrangement of 'Cuiguan' is different with that of'Cuiyu'. The distribution of cuticle thickness in 'Cuiguan' is uneven, and its cuticle thickness is half of that of'Cuiyu'; the cuticle of both two cultivars ruptured and formed netted texture. But 'Cuiyu' generated uniform and shallow cracks,'Cuiguan' generated disorder cracks and deep into the inner epidermis.'Yu Guan' and 'Qingxiang' fruit formation period than 'Cuiguan' and 'Cuiyu' early about50d.'Cuiguan' fruit rust is composed of single-layer corky epidermal cells that constitute the 'Yu Guan' brown leather by multilayer corky consists of cells, the cell patchy distribution. The ultra-structure of epidermal cells showed that the1-2layer cells under the cuticle had dense cytoplasm, and rich of liposomes. The liposomes contain small lipid droplets. The structural feature indicates that the1-2layer cells under the cuticle had strong ability to secrete waxes, but the ability differed with cultivars. The fatty granule was intensive in 'Cuiyu' liposomes, but'Cuiguan'lacked of liposomes. The research also showed that the fruit rust of 'Cuiguan' mainly formed in the latter period of fruit development; moisture is a main external factor resulted in the rust formation.
3. Two markers linked with fruit skin color were obtained through screening64pairs of AFLP primers using 'Akibae'×'Chuxialu' hybrid offspring. One of the markers (E-AAG/M-CAG) amplified missing band in80%green-skin-fruit offspring, and amplified the target band in74%of non-green materials; the other maker (E-ACT/M-CTT) amplified missing band in77%russet-skin-fruit offspring, and amplified the target band in89%non-pure-russett-skin-fruit materials.
4. Eight of three types genes including vacuolar proton pump PPase, subunit A of ATPase, auxin receptor gene, ABA receptor gene, ethylene receptor gene and the genes involved in fruit weight and disease resistance were selected for PCR analysis of apple (Malus domestica Borkh) and pear (Pyrus pyrifolia). For each gene, the primers were designed in exon and intron regions as well as in the regions upstream of start codon and downstream of stop codon, respectively. These primers were used to amplify apple and sand pear genomic DNA. It was found that all the genes have at least one pair of primers amplifying specific bands in pear genome DNA except the disease-resistant gene. Compared to the products amplified at gene other regions, the PCR products at upstream of start codon were more polymorphics, and the PCR products at exon and intron regions showed higher conservation, among different sand pear genotypes. Results of exon PCR product sequencing showed that the fragments of sand pear and apple corresponding genes in the area shared89.6%to97.5%homology. Four putative promoter sequences of lignin biosynthetic regulatory factor gene MYB were cloned from the sand pear (P. pyrifolia) with homology method. The four clones have the length of821bp,874bp,972bp and914bp, and their GenBank accession number was JF900710, JF900709, JF900707and JF900708, respectively. These sequences have the typical characteristics of gene promoters. The allelic analysis of the promoter sequences between green-and russet-skin fruit sand pear genotypes revealed that a SNP (A/G) site in PpMYBx2promoter region may have correlation with the fruit skin character.
1、以‘西子绿’和‘清香’为试材,对乙醇和丙酮2种有机溶剂浸提法提取果皮和果肉中叶绿素及类胡萝卜素的效果进行了比较分析,结果表明：80%丙酮浸提法和95%乙醇浸提法的效果相似,虽然应用95%乙醇浸提法的各种色素测定结果略偏低,但除了叶绿素a之外,2种提取方法间的差异不显著。并发现4个绿皮品种和4个褐皮品种果实发育过程中果实表皮及果肉中叶绿素、类胡萝卜素的含量与果实皮色表型无关。同时,以‘翠冠’与‘玉冠’为试材,测定了果实发育进程中纵横径、果皮与果肉色素含量的动态变化,以及在成熟前期可滴定酸、可溶性固形物、糖含量的变化特性。结果显示,两品种果实膨大过程均呈单“s”型曲线动态变化。在成熟前14d(翠冠)、28d(玉冠)至采收,果实糖、酸含量急剧变化,且两品种间的变化特性不同,生产上应依据不同品种果实糖、酸形成特性,确定适宜采收时期。
2、以绿皮品种‘翠冠’、‘翠玉’与褐皮品种‘清香’、‘玉冠’为试材,进行了果实表皮组织结构显微和超微水平观察,结果表明：避雨栽培能显著减小‘翠冠’果点的大小,而对‘翠玉’的果点大小没有显著影响。‘翠冠’果实表皮细胞的形状和排列方式与‘翠玉’不同,且角质膜分布厚薄不均,厚度仅为‘翠玉’的一半；两品种果实表皮的角质膜均出现破裂,并形成网状结构,但‘翠玉’龟裂产生的裂纹均匀且浅,而‘翠冠’龟裂产生的裂纹紊乱,且深及表皮层内部。‘玉冠’和‘清香’的果点形成时期要比‘翠冠’和‘翠玉’早50d左右。‘翠冠’果锈是由单层木栓化的表皮细胞构成,‘玉冠’褐皮则由多层木栓化的细胞构成,上层细胞片状分布。表皮细胞超微结构观察结果显示,在角质膜下第1-2层细胞的细胞质稠密,且含丰富的脂质体,脂质体内含圆形小脂滴。这一结构特征表明,梨果实表皮角质膜下第1-2层细胞具有旺盛的分泌蜡脂的能力,但品种间存在差异,‘翠玉’脂质体中的脂滴密集,而‘翠冠’中脂滴稀少。研究还表明,‘翠冠’的果锈形成期主要在果实发育中后期；水分是诱发果锈形成的主要外因。
3、以‘秋荣’ב初夏绿’的杂交后代群体为试材,通过64对AFLP标记引物在亲本和分离群体中的筛选和验证,获得与梨果实皮色相关基因连锁的标记2个,即E-AAG/M-CAG和E-ACT/M-CTT,前者在74%的非纯绿材料中扩增出目标带,在80%的纯绿皮材料中表现为缺失带型；在89%的非纯褐材料中扩增出了目标带,后者在77%的纯褐材料中表现为缺失带型。
4、选取苹果3种类型的8个基因(液泡质子泵焦磷酸水解酶基因、液泡质子泵三磷酸腺苷酶A亚基基因、生长素受体基因、脱落酸受体基因、乙烯受体基因、果重基因、抗病相关基因),分别在起始密码子上游区、终止密码子下游区、外显子区和内含子区设计引物,分别对苹果和砂梨的基因组DNA进行PCR扩增。结果显示,除了抗病相关基因,其余的7个基因均有引物在砂梨基因组DNA中扩增出特异条带。与基因其它区域扩增产物相比,起始密码子上游区扩增产物在不同砂梨材料间多态性较丰富,外显子和内含子区扩增产物在不同砂梨材料间的保守性更高。外显子区扩增产物测序结果表明：砂梨和苹果在该区对应基因片段具有89.6%-97.5%的核苷酸序列同源性。应用同源克隆方法从砂梨(Pyrus pyrifolia)中获得4个长度分别为821bp,874bp,972bp和914bp木质素合成调控因子MYB基因的启动子序列,GenBank登录号分别为JF900710、JF900709、JF900707和JF900708。生物信息学预测发现,这些序列具有典型的启动子序列特征。各启动子序列在果皮褐与绿的砂梨品种间的等位变异分析发现,PpMYBx2启动子区的一个SNP (A/G)位点与砂梨果皮褐、绿色性状间存在相关性。本研究为进一步开展砂梨MYB基因的表达分析、遗传定位,以及探明砂梨果实木质素形成的分子机制奠定了基础。
Appearance quality of pear fruits is mainly composed of skin color, size and density of fruit dot, the fruit surface waxiness, epidermal rust and other factors. The sand pear 'Cuiguan' is a cultivar with early maturity. It usually generates surface rust, which largely damages the fruit appearance quality. In order to reveal the mechanism of the appearance quality formation, we used 'Cuiguan' and other cultivars with different fruit peel color as materials and studied fruit peel pigment content dynamic changes and microscopic and ultrastructural features of epidermal tissue during different developmental stages of fruits; screened the AFLP markers linkage with fruit color genes using BSA classification; analyzed the application of apple fruit peel color related genes in pear homologues cloning, and based on which the preliminary study of molecular regulation of sand pear lignin biosynthesis was carried out. The major results are as follows:
1. Using 'Xizilv' and 'Qingxiang' as materials, we compared the efficiency of ethanol and acetone in chlorophyll and carotenoid extracting in peel and pulp. The results showed that the extracting efficiency of80%acetone is nearly similar with that of95%ethanol. Compared with80%acetone,95%ethanol resulted slightly lower pigment extraction, but there was no significant difference in the statistic analysis (except for chlorophyll a). The comparison of chlorophyll and carotenoid content in peel and pulp between the four green and four russet varieties at different development stages showed that the content of the pigments has no relation with the fruit skin color phenotype.'Cuiguan' and 'Yuguan' were used as materials to measure the dynamics changes of fruit vertical and horizontal diameter, pigments content in peel and pulp during the fruit development, and to measure titratable acid, soluble solids, and sugar contents during fruit maturing and harvest stage. The results showed that the development of fruit size of two varieties presented "S" curve; there was no significant differences of total chlorophyll and carotenoid content in peel and pulp between russet and green pear. During maturation (14days before harvest of'Cuiguan',28days before harvest of'Yuguan') and havest stage, fruit growth rate decreased, and the contents of titratable acid and sugars dramatically increased. The properties of these were difference between the two cultivars. Different harvest date should be determined according to the character of fruit inner quality formation for certain cultivar.
2. The epidermis structure observation was carried out at microscopic and ultra-microscopic levels using green-skin fruit sand pear cultivars'Cuiguan' and 'Cuiyu' and russet-skin fruit sand pear cultivars 'Qingxiang' and 'Yuguan'. The results showed that rain shelter cultivation could significantly reduce'Cuiguan'fruit dot size, while the'Cuiyu 'the fruit dot size does not significantly affect.the epidermal cell shape and arrangement of 'Cuiguan' is different with that of'Cuiyu'. The distribution of cuticle thickness in 'Cuiguan' is uneven, and its cuticle thickness is half of that of'Cuiyu'; the cuticle of both two cultivars ruptured and formed netted texture. But 'Cuiyu' generated uniform and shallow cracks,'Cuiguan' generated disorder cracks and deep into the inner epidermis.'Yu Guan' and 'Qingxiang' fruit formation period than 'Cuiguan' and 'Cuiyu' early about50d.'Cuiguan' fruit rust is composed of single-layer corky epidermal cells that constitute the 'Yu Guan' brown leather by multilayer corky consists of cells, the cell patchy distribution. The ultra-structure of epidermal cells showed that the1-2layer cells under the cuticle had dense cytoplasm, and rich of liposomes. The liposomes contain small lipid droplets. The structural feature indicates that the1-2layer cells under the cuticle had strong ability to secrete waxes, but the ability differed with cultivars. The fatty granule was intensive in 'Cuiyu' liposomes, but'Cuiguan'lacked of liposomes. The research also showed that the fruit rust of 'Cuiguan' mainly formed in the latter period of fruit development; moisture is a main external factor resulted in the rust formation.
3. Two markers linked with fruit skin color were obtained through screening64pairs of AFLP primers using 'Akibae'×'Chuxialu' hybrid offspring. One of the markers (E-AAG/M-CAG) amplified missing band in80%green-skin-fruit offspring, and amplified the target band in74%of non-green materials; the other maker (E-ACT/M-CTT) amplified missing band in77%russet-skin-fruit offspring, and amplified the target band in89%non-pure-russett-skin-fruit materials.
4. Eight of three types genes including vacuolar proton pump PPase, subunit A of ATPase, auxin receptor gene, ABA receptor gene, ethylene receptor gene and the genes involved in fruit weight and disease resistance were selected for PCR analysis of apple (Malus domestica Borkh) and pear (Pyrus pyrifolia). For each gene, the primers were designed in exon and intron regions as well as in the regions upstream of start codon and downstream of stop codon, respectively. These primers were used to amplify apple and sand pear genomic DNA. It was found that all the genes have at least one pair of primers amplifying specific bands in pear genome DNA except the disease-resistant gene. Compared to the products amplified at gene other regions, the PCR products at upstream of start codon were more polymorphics, and the PCR products at exon and intron regions showed higher conservation, among different sand pear genotypes. Results of exon PCR product sequencing showed that the fragments of sand pear and apple corresponding genes in the area shared89.6%to97.5%homology. Four putative promoter sequences of lignin biosynthetic regulatory factor gene MYB were cloned from the sand pear (P. pyrifolia) with homology method. The four clones have the length of821bp,874bp,972bp and914bp, and their GenBank accession number was JF900710, JF900709, JF900707and JF900708, respectively. These sequences have the typical characteristics of gene promoters. The allelic analysis of the promoter sequences between green-and russet-skin fruit sand pear genotypes revealed that a SNP (A/G) site in PpMYBx2promoter region may have correlation with the fruit skin character.
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