生长调节剂对‘砀山酥梨’果实萼片脱落与宿存的影响机制研究
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摘要
‘砀山酥梨’(Pyrus bretschneideri cv. Dangshansuli)果实有脱萼果与宿萼果之分,脱萼果品质优于宿萼果,如何通过栽培技术措施来提高脱萼果率,对于‘砀山酥梨’优质高效生产具有重要意义。本研究以‘砀山酥梨’为试材,研究了生长调节剂对梨果实萼片发育、品质和新梢生长的影响；探讨了脱萼果与宿萼果内源激素和矿质元素含量的差异及外源PP333、GA3对幼果Fe吸收、IAA含量及其代谢途径中相关酶活性的影响；并开展了‘砀山酥梨’脱萼果与宿萼果中相关基因的克隆与表达特性研究,主要结果如下：
一、高浓度PBO、PP333虽能提高‘砀山酥梨’脱萼果率、果实固形物与可溶性糖含量,降低有机酸含量,但使果实石细胞明显增多、硬度增加、果点增大。2500mg·L-1PP333.2000mg· L-1PBO处理果实石细胞含量分别为1.52%和1.47%；果肉硬度分别为6.34kg· cm-2和6.36kg· cm-2;果点直径分别为1.97mm和1.87mm,均显著高于对照。PBO、Flu对果实品质和新梢生长影响较PP333小,花期喷施其适宜浓度为200mg·L-1和100μL·L-1.2500mg·L-1PP333处理果点明显增大,超微结构观察表明,PP333处理使细胞排列紧凑、细胞壁变薄、木栓化程度提高,且表皮裂痕增大。
二、授粉品种与花粉萌发状况对‘砀山酥梨’脱萼果率没有影响；宿萼果中IAA含量明显高于脱萼果,而GA3、ABA和ZR并无显著差异；PBO、PP333显著提高了
‘砀山酥梨’脱萼果率,且明显降低了幼果中内源IAA含量；GA3能显著降低脱萼果率,且明显提高了幼果中IAA含量。生长调节剂影响果实萼片的发育,很可能与调节幼果中内源IAA水平有关。
‘砀山酥梨’和‘丰水’梨花柱中激素与矿质元素含量呈梯度分布,其中内源激素ABA、IAA和ZR含量从花柱形态学上端到下端呈逐渐升高的趋势,而GA3呈逐渐下降的趋势；矿质元素K、Ca、Fe、Zn和B含量从花柱形态学上端到下端均呈逐渐下降的趋势。用2.5mg·L-1IAA、5mg·L-16-BA处理花柱对花粉管伸长有显著促进作用；100mg·L-1H3BO4、300mg·L-1Ca(NO3)2处理对花粉管伸长的促进效果不如外源激素处理。进一步分析发现,梨花柱中ABA、IAA含量高于花粉,说明花粉萌发过程中所需的主要激素可能源自花柱,当花柱中激素含量增加时,可促进花粉管的伸长。梨花粉中矿质元素K、Ca、Fe、Zn、B高于花柱与子房,推测花粉萌发过程中所需的矿质元素可能来自花粉自身,当花柱中矿质元素含量增加时,对花粉管的伸长并无明显的促进作用。
三、‘砀山酥梨’脱萼果中Fe含量极显著低于宿萼果。本试验克隆了3个铁蛋白基因,即PbFer2、PbFer3和PbFer4,其中,脱萼果中PbFer2相对表达量要低于宿萼果,GA3能提高幼果中PbFer2的表达量,说明PbFer2与幼果中铁的贮存与吸收有关；脱萼果中PbFer3的表达量却显著高于宿萼果,推断环境胁迫可能诱导了PbFer3的表达。‘砀山酥梨’宿萼幼果与萼片中Fe含量差异最大,萼片中Fe含量是幼果的6.13倍,其它元素差异较小；萼片发育过程中对Fe的需求量明显高于幼果,而对其他矿质元素的需求量则少于幼果。400mg·L-1GA3处理增加了宿萼果率,且增加了幼果与萼片Fe的含量；2500mg·L-1PP333处理降低了宿萼果率,且减少了幼果与萼片中Fe的含量,喷施0.3%FeS04又提高了宿萼果率。因此,Fe是萼片发育重要的矿质元素之一,Fe充足时果实萼片易宿存。同时,萼片中叶绿素a含量的动态变化与Fe含量的动态变化呈显著性正相关,表明萼片中大量的Fe可能用于叶绿素a的合成。
四、通过cDNA-AFLP分析,在‘砀山酥梨’脱萼果与宿萼果中发现了10条TDFs (Transcript-derived fragments)与已登录的相关基因同源性较高。通过RACE (Rapid-amplification of cDNA ends)技术,克隆了ARF和MT基因的cDNA全长。其中,ARF基因全长1280bp,其中5'非翻译区44bp,3'非翻译区315bp,编码区为921bp,编码306个氨基酸,推测蛋白分子量为32.2243kDa,理论等电点为7.73。该基因具有Aux/IAA蛋白结构域和Aux/IAA-ARF-dimerisation2种结构域。IAA-ARF-dimerisation是Aux/IAA.ARF二聚体化的作用位点。推断该蛋白具有ARF类蛋白功能,可能是生长素信号转导途径中的转录因子。系统进化分析表明,与苹果MdAFR3基因编码的氨基酸序列同源性最高。因此,将其命名为PbARF3;MT基因全长552bp,其中5'非翻译区129bp,3’非翻译区222bp,编码区为201bp,编码66个氨基酸,推测蛋白分子量为6.4498kDa,理论等电点为4.79。并具有金属硫蛋白基因的典型结构域特征,富含半胱氨酸残基。系统进化分析表明,与苹果(Malus domestica)和樱桃(Prunus avium)MT2基因编码的氨基酸序列同源性高。因此,将其命名为PbMT2基因。GA3能显著提高幼果中PbARF3的表达量,且能提高幼果中PbMT2基因的表达量。其中PbARF3可能调控果实萼片的发育。PbMT2能提高了植物对金属离子的吸收和利用,而前期试验结果表明,Fe有利于果实萼片的宿存,据此推测,PbMT2调控幼果萼片的发育可能与促进其对Fe的吸收和利用有关。
五、‘砀山酥梨’宿萼果中色氨酸(Trp)含量和吲哚乙醛氧化酶(IAAldO)活性均高于脱萼果,而色氨酸转氨酶(TRT)活性无明显差异。GA3提高了幼果中TRT和IAAldO的活性。宿萼果中过氧化物酶(POD)低于脱萼果,且PP333处理提高了幼果中吲哚乙酸氧化酶(IAAO)和POD的活性。因此,GA3提高‘砀山酥梨’幼果内IAA的含量主要与提高IAA合成途径中相关酶的活性有关,而PP333降低幼果内IAA的含量主要与提高IAA降解途径中相关酶的活性有关,研究结果进一步揭示了PP333和GA3调控梨果实萼片发育的生理机制。
There are two types of Pyrus bretschneideri cv.'Dangshansuli' fruit:CSF (calyx-shed fruit) and CPF (calyx-persistency fruit). The fruit quality of CSF is better than that of CPF. It is therefore very important to reduce the occurrence of CPF through optimum culture technology, which can improve the fruit quality and planting benefit of 'Dangshansuli' pear.'Dangshansuli' was used as experiment materials in this paper, the effects of growth regulators on the fruit calyx development, fruit quality and shoot growth were researched. Differences of the endogenous hormones and the mineral element content between in CSF and CPF, effects of PP333(Paclobutrazol) and GA3(Giberellic acid) on the Fe absorption and IAA content in young fruit, and the effects of PP333and GA3on the key enzyme activities in IAA metabolic pathways were all discussed. The relatived genes clone and its expression characters in CSF and CPF were studied. The main results as follows:
Ⅰ. The percentage of the CSF, the soluble solids and soluble suger contents in fruit were increased, and the organic acid contents in fruit, was decreased, but the stone cell content, flesh firmness and fruit spot diameter were respectively increased by application of the high concentrations of PBO (Mixture of paclobutrazol and6-benzyladenine), PP333. The stone cell content in fruit were1.52%and1.47%, the flesh firmness were6.34kg· cm-2and6.36kg· cm-2, and fruit spot diameter were1.97mm and1.87mm treated with the2500mg· L-1PP333and2000mg· L-1PBO respectively, they were all significant higher than the contrast. Flu (Flusilazole) and PBO have a smaller effect on the fruit quality and shoot growth comparing to PP333treatment, and the appropriate concentrations of spraying with PBO and Flu were respectively200mg· L-1and100μL·L-1at the full-bloom stage. The fruit spots were enlarged, the cells arrangement of fruit dot was compact, the cell wall was thinning, the skin suberization and cracks were increased by the treatment of2500mg· L-1 PP333.
Ⅱ. No positive differences were discovered between the calyx shedding behavior of young fruit and either the different pollinators, or the different pollen germination conditions. The IAA content of fruit without calyx was lower than that of fruit with calyx, while the levels of GA3, ABA and BR did not differ significantly between the two types of fruit. The percentage of calyx-shed fruit could be significantly increased by spaying PBO and PP333in blossom, but the IAA content in young fruit decreased correspondingly. On the contrary, calyx-shedding was reduced by spraying with GA3, but the IAA content in young fruit increased as a result. In summary, we showed that fruit calyx development in 'Dangshansuli' pear were unaffected by pollinators and pollen germination conditions, and that calyx shedding is probably due to reduced levels of endogenous IAA in young fruit, which can be brought about by the application of PBO and PP333.
The hormone levels in style of 'Dangshansuli' and 'Housi' exhibit gradient distribution, from the top to the bottom in style morphology, the endogenous hormones ABA, IAA, ZR contents were gradually increased, while GA3was gradually decreased; The mineral elements K, Ca, Fe, Zn, B showed a gradually decreased. The elongation of pollen tube could be promoted treated with2.5mg·L-1IAA and5mg·L-16-BA; Further analysis showed that the ABA, IAA contents in style is higher than that in pollen, so the major hormones for pollen germination requirement probably come from style, because the elongation of pollen tubes could be promoted by increased hormone levels in style. The mineral elements K, Ca, Fe, Zn and B in pollen higher than the style and ovary, therefore, the required mineral elements in the process of pollen germination may come from the pollen itself, because the elongation of pollen tubes could not be significantly promoted by increased mineral levels in style.
Ⅲ. The Fe content in CPF was higher than that in CSF among the tested nine mineral elements. There are3Ferritin genes were cloned, which are PbFer2、PbFer3and PbFer4, the PbFer2relative expression in CPF was higher than in CSF, and GA3could enhance PbFer2expression level in young fruits too, so the PbFer2could promote iron storage and absorption for young fruit, The PbFer3expression level in CSF were remarkably higher than in CPF, we can peculated that the environmental stress may induce the expression of PbFer3. The Fe content difference between in fruitlet and in its calyx whose Fe content was6.13times higher than that in fruitlet was the biggest among the tested nine mineral elements. During the sepal developing process, the Iron demands were obviously higher than the fruitlets but at the same time of less demands for other mineral elements. The percentage of calyx persistency fruit, Fe content in fruitlet and its calyx were all increased under the spraying of400mg· L-1GA3. On the contrary, the percentage of calyx persistency fruit and the Fe content were decreased under th treatment of2500mg· L-1PP333, otherwisely, the calyx persistency fruit percentage was increased after spraying0.3%FeSO4solution too. So it was postulated that the sufficient Fe content was the guarantees to both fruit calyx development and the fruit calyx persistency. There was an positive correlation between Fe and Chla content dynamic changes in fruitlet calyx, it indicated that the abundant of Fe was probably used for Chla synthesis.
Ⅳ. The results of the cDNA-AFLP analysis showed that there were10TDFs (Transcript-derived fragments) between CSF and CPF, their homology were high with related genes by NCBI alignments. The full length of PbARF and PbMT genes were cloned by RACE (Rapid-amplification of cDNA ends) techniques. The full length cDNA of ARF gene was1280bp, including44bp of5' untranslated region and315bp of3'untranslated region. It had a coding sequence of921bp, which encoding306amino acid residues. The ARF protein molecular weight was32.2243kD and the theoretical pI was7.73. It has two kinds of structure domain about Aux/IAA protein structure domain and the Aux/IAA-ARF-dimerisation. IAA-ARF-dimerisation is the target site of Aux/IAA-ARF dimers, which related to the function of ARF kind protein, and involved in auxin signal transduction pathways. The phylogenetic tree analysis revealed that the gene have highest similarity with Malus domestic AFR3gene amino acid sequence, therefore it was named PbARF3. The full length cDNA of MT gene was552bp, including129bp of5'untranslated region and222bp of3'untranslated region. It had a coding sequence of201bp, which encoded66amino acid residues. The protein molecular weight was6.4498kD and the theoretical pI was4.79. It contained the conserved domains of all known MTs, rich in homocysteine residual base. The phylogenetic tree analysis revealed that that the amino acids sequence of MT shared higher similarity with Malus domestica and Prunus avium, therefore, it was named PbMT2. The PbARF3expression quantity in young fruit was significantly improved by GA3, and the PbMT2gene expression quantity was significantly improved by GA3in young fruit at the beginning of calyx abscission. It concluded that PbARF3may control the fruit calyx development. The above research showed that the sufficient Fe was beneficial to fruit calyx persistency, so the regulation of PbMT2on calyx development probably was attributed to improve iron absorbing and utilizing of young fruit.
V. Tryptophane (Trp) and indole aldehyde oxidase (IAAldO) in CPF were higher than that of CSF, and the Trp transaminase (TRT) activity have no obvious difference between in CPF and CSF, the TRT and IAAldO activity in young fruit were improved by GA3. The peroxidase (POD) in CPF was lower than that in CSF, and IAA oxidase (IAAO) and POD activity were increased by PP333. Therefore, the increase of IAA content treated with GA3was attributed to the related enzymes activities in IAA synthesis pathway, and the decrease of IAA content treated with PP333was attributed to the related enzymes activities in IAA degradation pathway. The main results laid a foundation to reveal the physiological mechanism of fruit calyx development of 'Dangshansuli' pear regulated by exogenous growth regulators further more.
一、高浓度PBO、PP333虽能提高‘砀山酥梨’脱萼果率、果实固形物与可溶性糖含量,降低有机酸含量,但使果实石细胞明显增多、硬度增加、果点增大。2500mg·L-1PP333.2000mg· L-1PBO处理果实石细胞含量分别为1.52%和1.47%；果肉硬度分别为6.34kg· cm-2和6.36kg· cm-2;果点直径分别为1.97mm和1.87mm,均显著高于对照。PBO、Flu对果实品质和新梢生长影响较PP333小,花期喷施其适宜浓度为200mg·L-1和100μL·L-1.2500mg·L-1PP333处理果点明显增大,超微结构观察表明,PP333处理使细胞排列紧凑、细胞壁变薄、木栓化程度提高,且表皮裂痕增大。
二、授粉品种与花粉萌发状况对‘砀山酥梨’脱萼果率没有影响；宿萼果中IAA含量明显高于脱萼果,而GA3、ABA和ZR并无显著差异；PBO、PP333显著提高了
‘砀山酥梨’脱萼果率,且明显降低了幼果中内源IAA含量；GA3能显著降低脱萼果率,且明显提高了幼果中IAA含量。生长调节剂影响果实萼片的发育,很可能与调节幼果中内源IAA水平有关。
‘砀山酥梨’和‘丰水’梨花柱中激素与矿质元素含量呈梯度分布,其中内源激素ABA、IAA和ZR含量从花柱形态学上端到下端呈逐渐升高的趋势,而GA3呈逐渐下降的趋势；矿质元素K、Ca、Fe、Zn和B含量从花柱形态学上端到下端均呈逐渐下降的趋势。用2.5mg·L-1IAA、5mg·L-16-BA处理花柱对花粉管伸长有显著促进作用；100mg·L-1H3BO4、300mg·L-1Ca(NO3)2处理对花粉管伸长的促进效果不如外源激素处理。进一步分析发现,梨花柱中ABA、IAA含量高于花粉,说明花粉萌发过程中所需的主要激素可能源自花柱,当花柱中激素含量增加时,可促进花粉管的伸长。梨花粉中矿质元素K、Ca、Fe、Zn、B高于花柱与子房,推测花粉萌发过程中所需的矿质元素可能来自花粉自身,当花柱中矿质元素含量增加时,对花粉管的伸长并无明显的促进作用。
三、‘砀山酥梨’脱萼果中Fe含量极显著低于宿萼果。本试验克隆了3个铁蛋白基因,即PbFer2、PbFer3和PbFer4,其中,脱萼果中PbFer2相对表达量要低于宿萼果,GA3能提高幼果中PbFer2的表达量,说明PbFer2与幼果中铁的贮存与吸收有关；脱萼果中PbFer3的表达量却显著高于宿萼果,推断环境胁迫可能诱导了PbFer3的表达。‘砀山酥梨’宿萼幼果与萼片中Fe含量差异最大,萼片中Fe含量是幼果的6.13倍,其它元素差异较小；萼片发育过程中对Fe的需求量明显高于幼果,而对其他矿质元素的需求量则少于幼果。400mg·L-1GA3处理增加了宿萼果率,且增加了幼果与萼片Fe的含量；2500mg·L-1PP333处理降低了宿萼果率,且减少了幼果与萼片中Fe的含量,喷施0.3%FeS04又提高了宿萼果率。因此,Fe是萼片发育重要的矿质元素之一,Fe充足时果实萼片易宿存。同时,萼片中叶绿素a含量的动态变化与Fe含量的动态变化呈显著性正相关,表明萼片中大量的Fe可能用于叶绿素a的合成。
四、通过cDNA-AFLP分析,在‘砀山酥梨’脱萼果与宿萼果中发现了10条TDFs (Transcript-derived fragments)与已登录的相关基因同源性较高。通过RACE (Rapid-amplification of cDNA ends)技术,克隆了ARF和MT基因的cDNA全长。其中,ARF基因全长1280bp,其中5'非翻译区44bp,3'非翻译区315bp,编码区为921bp,编码306个氨基酸,推测蛋白分子量为32.2243kDa,理论等电点为7.73。该基因具有Aux/IAA蛋白结构域和Aux/IAA-ARF-dimerisation2种结构域。IAA-ARF-dimerisation是Aux/IAA.ARF二聚体化的作用位点。推断该蛋白具有ARF类蛋白功能,可能是生长素信号转导途径中的转录因子。系统进化分析表明,与苹果MdAFR3基因编码的氨基酸序列同源性最高。因此,将其命名为PbARF3;MT基因全长552bp,其中5'非翻译区129bp,3’非翻译区222bp,编码区为201bp,编码66个氨基酸,推测蛋白分子量为6.4498kDa,理论等电点为4.79。并具有金属硫蛋白基因的典型结构域特征,富含半胱氨酸残基。系统进化分析表明,与苹果(Malus domestica)和樱桃(Prunus avium)MT2基因编码的氨基酸序列同源性高。因此,将其命名为PbMT2基因。GA3能显著提高幼果中PbARF3的表达量,且能提高幼果中PbMT2基因的表达量。其中PbARF3可能调控果实萼片的发育。PbMT2能提高了植物对金属离子的吸收和利用,而前期试验结果表明,Fe有利于果实萼片的宿存,据此推测,PbMT2调控幼果萼片的发育可能与促进其对Fe的吸收和利用有关。
五、‘砀山酥梨’宿萼果中色氨酸(Trp)含量和吲哚乙醛氧化酶(IAAldO)活性均高于脱萼果,而色氨酸转氨酶(TRT)活性无明显差异。GA3提高了幼果中TRT和IAAldO的活性。宿萼果中过氧化物酶(POD)低于脱萼果,且PP333处理提高了幼果中吲哚乙酸氧化酶(IAAO)和POD的活性。因此,GA3提高‘砀山酥梨’幼果内IAA的含量主要与提高IAA合成途径中相关酶的活性有关,而PP333降低幼果内IAA的含量主要与提高IAA降解途径中相关酶的活性有关,研究结果进一步揭示了PP333和GA3调控梨果实萼片发育的生理机制。
There are two types of Pyrus bretschneideri cv.'Dangshansuli' fruit:CSF (calyx-shed fruit) and CPF (calyx-persistency fruit). The fruit quality of CSF is better than that of CPF. It is therefore very important to reduce the occurrence of CPF through optimum culture technology, which can improve the fruit quality and planting benefit of 'Dangshansuli' pear.'Dangshansuli' was used as experiment materials in this paper, the effects of growth regulators on the fruit calyx development, fruit quality and shoot growth were researched. Differences of the endogenous hormones and the mineral element content between in CSF and CPF, effects of PP333(Paclobutrazol) and GA3(Giberellic acid) on the Fe absorption and IAA content in young fruit, and the effects of PP333and GA3on the key enzyme activities in IAA metabolic pathways were all discussed. The relatived genes clone and its expression characters in CSF and CPF were studied. The main results as follows:
Ⅰ. The percentage of the CSF, the soluble solids and soluble suger contents in fruit were increased, and the organic acid contents in fruit, was decreased, but the stone cell content, flesh firmness and fruit spot diameter were respectively increased by application of the high concentrations of PBO (Mixture of paclobutrazol and6-benzyladenine), PP333. The stone cell content in fruit were1.52%and1.47%, the flesh firmness were6.34kg· cm-2and6.36kg· cm-2, and fruit spot diameter were1.97mm and1.87mm treated with the2500mg· L-1PP333and2000mg· L-1PBO respectively, they were all significant higher than the contrast. Flu (Flusilazole) and PBO have a smaller effect on the fruit quality and shoot growth comparing to PP333treatment, and the appropriate concentrations of spraying with PBO and Flu were respectively200mg· L-1and100μL·L-1at the full-bloom stage. The fruit spots were enlarged, the cells arrangement of fruit dot was compact, the cell wall was thinning, the skin suberization and cracks were increased by the treatment of2500mg· L-1 PP333.
Ⅱ. No positive differences were discovered between the calyx shedding behavior of young fruit and either the different pollinators, or the different pollen germination conditions. The IAA content of fruit without calyx was lower than that of fruit with calyx, while the levels of GA3, ABA and BR did not differ significantly between the two types of fruit. The percentage of calyx-shed fruit could be significantly increased by spaying PBO and PP333in blossom, but the IAA content in young fruit decreased correspondingly. On the contrary, calyx-shedding was reduced by spraying with GA3, but the IAA content in young fruit increased as a result. In summary, we showed that fruit calyx development in 'Dangshansuli' pear were unaffected by pollinators and pollen germination conditions, and that calyx shedding is probably due to reduced levels of endogenous IAA in young fruit, which can be brought about by the application of PBO and PP333.
The hormone levels in style of 'Dangshansuli' and 'Housi' exhibit gradient distribution, from the top to the bottom in style morphology, the endogenous hormones ABA, IAA, ZR contents were gradually increased, while GA3was gradually decreased; The mineral elements K, Ca, Fe, Zn, B showed a gradually decreased. The elongation of pollen tube could be promoted treated with2.5mg·L-1IAA and5mg·L-16-BA; Further analysis showed that the ABA, IAA contents in style is higher than that in pollen, so the major hormones for pollen germination requirement probably come from style, because the elongation of pollen tubes could be promoted by increased hormone levels in style. The mineral elements K, Ca, Fe, Zn and B in pollen higher than the style and ovary, therefore, the required mineral elements in the process of pollen germination may come from the pollen itself, because the elongation of pollen tubes could not be significantly promoted by increased mineral levels in style.
Ⅲ. The Fe content in CPF was higher than that in CSF among the tested nine mineral elements. There are3Ferritin genes were cloned, which are PbFer2、PbFer3and PbFer4, the PbFer2relative expression in CPF was higher than in CSF, and GA3could enhance PbFer2expression level in young fruits too, so the PbFer2could promote iron storage and absorption for young fruit, The PbFer3expression level in CSF were remarkably higher than in CPF, we can peculated that the environmental stress may induce the expression of PbFer3. The Fe content difference between in fruitlet and in its calyx whose Fe content was6.13times higher than that in fruitlet was the biggest among the tested nine mineral elements. During the sepal developing process, the Iron demands were obviously higher than the fruitlets but at the same time of less demands for other mineral elements. The percentage of calyx persistency fruit, Fe content in fruitlet and its calyx were all increased under the spraying of400mg· L-1GA3. On the contrary, the percentage of calyx persistency fruit and the Fe content were decreased under th treatment of2500mg· L-1PP333, otherwisely, the calyx persistency fruit percentage was increased after spraying0.3%FeSO4solution too. So it was postulated that the sufficient Fe content was the guarantees to both fruit calyx development and the fruit calyx persistency. There was an positive correlation between Fe and Chla content dynamic changes in fruitlet calyx, it indicated that the abundant of Fe was probably used for Chla synthesis.
Ⅳ. The results of the cDNA-AFLP analysis showed that there were10TDFs (Transcript-derived fragments) between CSF and CPF, their homology were high with related genes by NCBI alignments. The full length of PbARF and PbMT genes were cloned by RACE (Rapid-amplification of cDNA ends) techniques. The full length cDNA of ARF gene was1280bp, including44bp of5' untranslated region and315bp of3'untranslated region. It had a coding sequence of921bp, which encoding306amino acid residues. The ARF protein molecular weight was32.2243kD and the theoretical pI was7.73. It has two kinds of structure domain about Aux/IAA protein structure domain and the Aux/IAA-ARF-dimerisation. IAA-ARF-dimerisation is the target site of Aux/IAA-ARF dimers, which related to the function of ARF kind protein, and involved in auxin signal transduction pathways. The phylogenetic tree analysis revealed that the gene have highest similarity with Malus domestic AFR3gene amino acid sequence, therefore it was named PbARF3. The full length cDNA of MT gene was552bp, including129bp of5'untranslated region and222bp of3'untranslated region. It had a coding sequence of201bp, which encoded66amino acid residues. The protein molecular weight was6.4498kD and the theoretical pI was4.79. It contained the conserved domains of all known MTs, rich in homocysteine residual base. The phylogenetic tree analysis revealed that that the amino acids sequence of MT shared higher similarity with Malus domestica and Prunus avium, therefore, it was named PbMT2. The PbARF3expression quantity in young fruit was significantly improved by GA3, and the PbMT2gene expression quantity was significantly improved by GA3in young fruit at the beginning of calyx abscission. It concluded that PbARF3may control the fruit calyx development. The above research showed that the sufficient Fe was beneficial to fruit calyx persistency, so the regulation of PbMT2on calyx development probably was attributed to improve iron absorbing and utilizing of young fruit.
V. Tryptophane (Trp) and indole aldehyde oxidase (IAAldO) in CPF were higher than that of CSF, and the Trp transaminase (TRT) activity have no obvious difference between in CPF and CSF, the TRT and IAAldO activity in young fruit were improved by GA3. The peroxidase (POD) in CPF was lower than that in CSF, and IAA oxidase (IAAO) and POD activity were increased by PP333. Therefore, the increase of IAA content treated with GA3was attributed to the related enzymes activities in IAA synthesis pathway, and the decrease of IAA content treated with PP333was attributed to the related enzymes activities in IAA degradation pathway. The main results laid a foundation to reveal the physiological mechanism of fruit calyx development of 'Dangshansuli' pear regulated by exogenous growth regulators further more.
引文
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