茉莉（Jasminum sambac L.）对低温胁迫的生理响应及离体快繁研究
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摘要
本研究选用单、双瓣茉莉2年生扦插苗为材料进行低温处理,探讨茉莉在模拟降温和自然降温过程中多个生理生化指标变化规律,旨在探讨茉莉对低温胁迫的抗性机制；应用外源物质对茉莉叶片进行预处理,研究水杨酸(SA)和脱落酸(ABA)对茉莉耐寒性的调控效果及机制；克隆与脯氨酸合成相关的△-1-吡咯琳-5-羧酸合成酶基因(P5CS),并通过半定量RT-PCR方法研究在低温胁迫条件下该基因的表达、脯氨酸含量及其相互之间的关系；并通过以带芽茎段为材料,优化茉莉的离体快繁体系。主要研究结果如下：
1.以2年生单、双瓣茉莉扦插苗为材料,采用人工模拟低温的方法,初步研究不同温度梯度和低温处理时间对茉莉植株多个生理指标的影响。结果表明：随着温度的降低和胁迫时间的延长,低温对单、双瓣茉莉的影响逐渐加剧；3℃低温胁迫8d后,茉莉各项生理指标的变化最大。随着温度的降低,叶片相对电导率、MDA含量、脯氨酸含量、可溶性蛋白含量、POD活性逐渐升高；根系活力和叶绿素含量则逐渐下降；其他指标则在不同处理时间后的变化趋势较为复杂。单瓣茉莉叶片SOD活性在各温度处理2d和8d的情况下均呈现先升后降趋势,而双瓣SOD活性则持续增加,且比单瓣的SOD活性强。双瓣茉莉叶片可溶性糖含量在各温度处理2d和4d时随温度降低并无显著变化,而在各温度处理8d的情况下则呈显著增加的趋势。综合分析发现,双瓣茉莉的耐寒能力强于单瓣茉莉。
2.以单、双瓣茉莉2年生扦插苗为试验材料,分析其在2009年南京地区秋冬自然降温过程中多个抗寒生理指标的特性,并比较单、双瓣茉莉间抗寒能力的差异。研究结果表明,在自然降温条件下,单、双瓣茉莉的相对电导率、MDA含量、脯氨酸含量均显著上升,可溶性蛋白含量先上升后下降,不同抗氧化酶活性表现出的趋势并不一致,根系活力呈下降趋势。综合多个生理指标分析得出：在本实验多次降温过程中双瓣茉莉的耐寒能力略强于单瓣茉莉,其中双瓣茉莉抗氧化酶活性表现最为突出。
3.用50、100、175、250mg·L-1的SA水溶液预处理单、双瓣茉莉幼苗,并经6℃低温胁迫处理3d后,综合比较叶片相对电导率、MDA含量、脯氨酸含量、可溶性糖含量、SOD酶活性等多项指标可以得出,喷施50mg·L-1的SA对单、双瓣茉莉缓解低温胁迫均起到了较好的效果；用0.5、5、10、20mg·L-1的ABA水溶液预处理单、双瓣茉莉叶片,并经6℃低温胁迫处理3d后,综合比较叶片相对电导率、MDA含量、脯氨酸含量、可溶性糖含量、可溶性蛋白含量、POD酶活性等多项指标可以得出,喷施5mg·L-1的ABA对单、双瓣茉莉缓解低温胁迫均起到较好的效果。
4.利用同源克隆的方法,从双瓣茉莉叶片中克隆得了脯氨酸合成路径中的关键酶基因(P5CS)片段。该基因片段大小为1205bp,编码401个氨基酸,序列分析表明该基因片段与蜡烛果、猕猴桃、烟草、番茄P5CS基因片段同源性达到75%以上。运用半定量RT-PCR方法研究该基因在低温胁迫不同时间的表达情况。研究结果表明,在3℃低温胁迫下,P5CS基因的表达量随着低温胁迫时间延长先升高后降低,高峰期是胁迫第4d,而脯氨酸含量的高峰期则为低温胁迫第6d,存在着一定的滞后性。同时,P5CS基因的转录水平到达高峰期后随着低温胁迫时间的延长而减弱,这与脯氨酸含量变化规律基本一致,这表明脯氨酸含量的增加可能与P5CS基因表达的增加存在正相关,但是这需要进一步的研究加以明确。
5.以8年生双瓣茉莉为材料,系统研究茉莉茎段快繁体系中多个影响因素,包括消毒处理和基本培养基对外植体启动培养的影响、腋芽增殖培养的最佳配方、最佳丛生芽诱导配方以及生根诱导配方等。主要结果如下：
(1)外植体采摘最佳时间为4-9月份(避开梅雨季节)。以当年生半木质化带腋芽的茎段为外植体,最佳消毒处理方法是先75%酒精60s,再用0.1%HgCl2消毒52min,无菌水冲洗3次。腋芽萌发的最佳基本培养基为WPM基本培养基。
(2)在WPM基本培养基中,通过添加不同浓度生长素、细胞分裂素、赤霉素进行单因素或多因素综合试验,发现腋芽最佳伸长生长配方：WPM+2mg·L-1IBA+0.5mg·L-16-BA+0.7mg·L-1GA3。在WPM培养基中添加1mg·L-1ZT和0.5mg·L-1IBA能有效诱导丛生芽,最多可达每茎段诱导11个丛生芽。
(3)在组培苗生根方面,本试验发现在无菌条件下,通过将芽苗浸泡在已过滤灭菌的450mg·L-1NAA溶液里10min,然后接种到1/2WPM培养基中进行生根诱导效果最好,生根率可达86.4%,且根系健壮,通过炼苗移栽组培苗成活率78%。
In this study, the physiological and biochemical characteristics of chilling stress under artificial low temperature and natural low temperature were investigated in2-year-old cuttings of two cultivars of Jasminum sambac L., single petal jasmine (SPJ) and double petal jasmine (DPJ). In order to unravel mechanism underlying the chilling tolerance of jasmine and to facilitate chilling tolerance breeding, the effect of exogenous salicylic acid (SA) and abscisic acid (ABA) on physiological and biochemical response of these two cultivars were examined. Also, the relationship between delta-1-Pyrroline-5-Carboxylate Synthetase(P5CS) gene and the proline content was studied through semi quantitative reverse transcription and polymerase chain reaction (Sq RT-PCR) under low temperature stress. Besides, the efficient in vitro micropropagation system of jasmine was developed by nodal segments with axillary buds.
From this research, the main results were as follows:
1. During the artificial temperature falling experiment, the nine kinds of physiological and biochemical indexes of chilling tolerance during the low temperature stress were investigated in2-year-old cuttings of two cultivars of J. sambac, SPJ and DPJ. The results showed that the effect of low temperature stress on two cultivars intensified gradually with temperature falling and elongation of treating time. The several physiological and biochemical indexes of jasmines changed severely while they were treated under3℃for8days. The relative electrolyte leakage (REL), peroxidase (POD) activity, the content of malondialdehyde (MDA), proline and soluble protein increased gradually; while the root activity and chlorophyll content decreased by degrees; and the other physiological and biochemical indexes changed complicatedly under different low temperature and treating time. The superoxide dismutase (SOD) activity of SPJ increased firstly and then decreased when it was treated under different temperature for2days and8days. The soluble protein content of DPJ was not different remarkably under different temperature treating for2days and4days but increased significantly under different temperature treating for8days. Through comprehensive analysis of these indexes, the chilling tolerance of DPJ was betterr than that of SPJ.
2. The chilling injury parameter and eight physiological and biochemical indexes of chilling-tolerance were investigated in2-year-old cuttings of SPJ and DPJ under the condition of natural temperature falling in2009year, Nanjing City. The results showed that the REL and the content of MDA and proline of these two cultivars increased significantly, the content of soluble protein increased at first and decreased afterwards, while the tendencies of antioxidant enzymes activity of the two cultivars were different and their root activity decreased gradually. Comprehensive assessment of those physiological indexes showed that the DPJ was a little bit more chilling-resistant than SPJ, especially the change of antioxidant enzymes activity of DPJ.
3. These two cultivars which were pretreated with SA solution (50,100,175and250mg·L-1) or ABA solution (0.5,5,10and20mg·L-1) respectively for4days were used for detecting the physiological and biochemical responses to6℃treating for3days. The REL, the content of MDA, proline and the soluble sugar, the activity of SOD and POD of two jasmines were investigated. Comprehensively comparation of above indexes of these two cultivars, both of50mg·L-1SA solution and5mg·L-1ABA solution were the suitable pretreatment to enhance the chilling resistance of these two jasmines under6℃treating for3days.
4. The P5CS gene segment which was concerned with proline synthesis was obtained from leaves of DPJ by method of homologous cloning. The gene segment had1205nucleotides and coded401amino acids. In comparison with the published sequences of P5CS gene by BLAST program, the P5CS gene segment of DPJ showed more than89%identity to that gene from Aegiceras corniculatum, Actinidia deliciosa, Nicotiana tabacum and Solanum lycopersicum. The P5CS gene expression during3℃was investigated through Sq RT-PCR method with the18S gene as reference gene. The results showed that the expression of P5CS was first up-regulated and then down-regulated during low temperature stress. The expressional fastigium of P5CS gene was at the fourth day of3℃treating, while the fastigium of proline content was at the sixth day of3℃treating. Although there was hysteresis phenomenon between the P5CS gene expression and proline synthesis, the trend of them was consilient. But further studies were needed to elucidate the relationship between them.
5. The micropropagation in vitro was investigated in8-year-old cuttings of DPJ by stem segment. Several experiments were carried out to detecte the disinfection of explant, germination culture of axillary buds, the optimum formula of proliferation and induction of multiple shoots and rooting. The results were as follows:
(1) April to September (avoiding the mould rains season) was the best period to pick the explants. The one-year-old and semi-lignification twigs with axillary buds were most suitable to be explants. The treatment with75%ethanol for1min,0.1%mercuric chloride for52mins then washing3times in aseptic water sequentially was the best disinfection method. The optimum basic medium of germination for axillary buds was WPM medium.
(2) In order to find out the optimum formula for axillary buds elongation, the experiments with different plant growth regulators alone or in combination were carried out. The best formula for axillary buds elongation was WPM+2mg·L-1IBA+0.5mg·L-16-BA+0.7mg·L-1GA3. The optimum combination of plant growth regulators for induction of multiple shoots, with generation of11axillary shoots, was1mg·L-1ZT supplemented with0.5mg·L-1IBA,
(3)The best rooting method was that shoots were first dipped in450mg·L-1NAA solution for10mins and cultured in the basal1/2WPM sequentially. The rooting rate was up to86.7%. In this study, the plantlets were acclimatized with78%survival rate under the suitable humidity and illumination during the initial period of acclimation.
1.以2年生单、双瓣茉莉扦插苗为材料,采用人工模拟低温的方法,初步研究不同温度梯度和低温处理时间对茉莉植株多个生理指标的影响。结果表明：随着温度的降低和胁迫时间的延长,低温对单、双瓣茉莉的影响逐渐加剧；3℃低温胁迫8d后,茉莉各项生理指标的变化最大。随着温度的降低,叶片相对电导率、MDA含量、脯氨酸含量、可溶性蛋白含量、POD活性逐渐升高；根系活力和叶绿素含量则逐渐下降；其他指标则在不同处理时间后的变化趋势较为复杂。单瓣茉莉叶片SOD活性在各温度处理2d和8d的情况下均呈现先升后降趋势,而双瓣SOD活性则持续增加,且比单瓣的SOD活性强。双瓣茉莉叶片可溶性糖含量在各温度处理2d和4d时随温度降低并无显著变化,而在各温度处理8d的情况下则呈显著增加的趋势。综合分析发现,双瓣茉莉的耐寒能力强于单瓣茉莉。
2.以单、双瓣茉莉2年生扦插苗为试验材料,分析其在2009年南京地区秋冬自然降温过程中多个抗寒生理指标的特性,并比较单、双瓣茉莉间抗寒能力的差异。研究结果表明,在自然降温条件下,单、双瓣茉莉的相对电导率、MDA含量、脯氨酸含量均显著上升,可溶性蛋白含量先上升后下降,不同抗氧化酶活性表现出的趋势并不一致,根系活力呈下降趋势。综合多个生理指标分析得出：在本实验多次降温过程中双瓣茉莉的耐寒能力略强于单瓣茉莉,其中双瓣茉莉抗氧化酶活性表现最为突出。
3.用50、100、175、250mg·L-1的SA水溶液预处理单、双瓣茉莉幼苗,并经6℃低温胁迫处理3d后,综合比较叶片相对电导率、MDA含量、脯氨酸含量、可溶性糖含量、SOD酶活性等多项指标可以得出,喷施50mg·L-1的SA对单、双瓣茉莉缓解低温胁迫均起到了较好的效果；用0.5、5、10、20mg·L-1的ABA水溶液预处理单、双瓣茉莉叶片,并经6℃低温胁迫处理3d后,综合比较叶片相对电导率、MDA含量、脯氨酸含量、可溶性糖含量、可溶性蛋白含量、POD酶活性等多项指标可以得出,喷施5mg·L-1的ABA对单、双瓣茉莉缓解低温胁迫均起到较好的效果。
4.利用同源克隆的方法,从双瓣茉莉叶片中克隆得了脯氨酸合成路径中的关键酶基因(P5CS)片段。该基因片段大小为1205bp,编码401个氨基酸,序列分析表明该基因片段与蜡烛果、猕猴桃、烟草、番茄P5CS基因片段同源性达到75%以上。运用半定量RT-PCR方法研究该基因在低温胁迫不同时间的表达情况。研究结果表明,在3℃低温胁迫下,P5CS基因的表达量随着低温胁迫时间延长先升高后降低,高峰期是胁迫第4d,而脯氨酸含量的高峰期则为低温胁迫第6d,存在着一定的滞后性。同时,P5CS基因的转录水平到达高峰期后随着低温胁迫时间的延长而减弱,这与脯氨酸含量变化规律基本一致,这表明脯氨酸含量的增加可能与P5CS基因表达的增加存在正相关,但是这需要进一步的研究加以明确。
5.以8年生双瓣茉莉为材料,系统研究茉莉茎段快繁体系中多个影响因素,包括消毒处理和基本培养基对外植体启动培养的影响、腋芽增殖培养的最佳配方、最佳丛生芽诱导配方以及生根诱导配方等。主要结果如下：
(1)外植体采摘最佳时间为4-9月份(避开梅雨季节)。以当年生半木质化带腋芽的茎段为外植体,最佳消毒处理方法是先75%酒精60s,再用0.1%HgCl2消毒52min,无菌水冲洗3次。腋芽萌发的最佳基本培养基为WPM基本培养基。
(2)在WPM基本培养基中,通过添加不同浓度生长素、细胞分裂素、赤霉素进行单因素或多因素综合试验,发现腋芽最佳伸长生长配方：WPM+2mg·L-1IBA+0.5mg·L-16-BA+0.7mg·L-1GA3。在WPM培养基中添加1mg·L-1ZT和0.5mg·L-1IBA能有效诱导丛生芽,最多可达每茎段诱导11个丛生芽。
(3)在组培苗生根方面,本试验发现在无菌条件下,通过将芽苗浸泡在已过滤灭菌的450mg·L-1NAA溶液里10min,然后接种到1/2WPM培养基中进行生根诱导效果最好,生根率可达86.4%,且根系健壮,通过炼苗移栽组培苗成活率78%。
In this study, the physiological and biochemical characteristics of chilling stress under artificial low temperature and natural low temperature were investigated in2-year-old cuttings of two cultivars of Jasminum sambac L., single petal jasmine (SPJ) and double petal jasmine (DPJ). In order to unravel mechanism underlying the chilling tolerance of jasmine and to facilitate chilling tolerance breeding, the effect of exogenous salicylic acid (SA) and abscisic acid (ABA) on physiological and biochemical response of these two cultivars were examined. Also, the relationship between delta-1-Pyrroline-5-Carboxylate Synthetase(P5CS) gene and the proline content was studied through semi quantitative reverse transcription and polymerase chain reaction (Sq RT-PCR) under low temperature stress. Besides, the efficient in vitro micropropagation system of jasmine was developed by nodal segments with axillary buds.
From this research, the main results were as follows:
1. During the artificial temperature falling experiment, the nine kinds of physiological and biochemical indexes of chilling tolerance during the low temperature stress were investigated in2-year-old cuttings of two cultivars of J. sambac, SPJ and DPJ. The results showed that the effect of low temperature stress on two cultivars intensified gradually with temperature falling and elongation of treating time. The several physiological and biochemical indexes of jasmines changed severely while they were treated under3℃for8days. The relative electrolyte leakage (REL), peroxidase (POD) activity, the content of malondialdehyde (MDA), proline and soluble protein increased gradually; while the root activity and chlorophyll content decreased by degrees; and the other physiological and biochemical indexes changed complicatedly under different low temperature and treating time. The superoxide dismutase (SOD) activity of SPJ increased firstly and then decreased when it was treated under different temperature for2days and8days. The soluble protein content of DPJ was not different remarkably under different temperature treating for2days and4days but increased significantly under different temperature treating for8days. Through comprehensive analysis of these indexes, the chilling tolerance of DPJ was betterr than that of SPJ.
2. The chilling injury parameter and eight physiological and biochemical indexes of chilling-tolerance were investigated in2-year-old cuttings of SPJ and DPJ under the condition of natural temperature falling in2009year, Nanjing City. The results showed that the REL and the content of MDA and proline of these two cultivars increased significantly, the content of soluble protein increased at first and decreased afterwards, while the tendencies of antioxidant enzymes activity of the two cultivars were different and their root activity decreased gradually. Comprehensive assessment of those physiological indexes showed that the DPJ was a little bit more chilling-resistant than SPJ, especially the change of antioxidant enzymes activity of DPJ.
3. These two cultivars which were pretreated with SA solution (50,100,175and250mg·L-1) or ABA solution (0.5,5,10and20mg·L-1) respectively for4days were used for detecting the physiological and biochemical responses to6℃treating for3days. The REL, the content of MDA, proline and the soluble sugar, the activity of SOD and POD of two jasmines were investigated. Comprehensively comparation of above indexes of these two cultivars, both of50mg·L-1SA solution and5mg·L-1ABA solution were the suitable pretreatment to enhance the chilling resistance of these two jasmines under6℃treating for3days.
4. The P5CS gene segment which was concerned with proline synthesis was obtained from leaves of DPJ by method of homologous cloning. The gene segment had1205nucleotides and coded401amino acids. In comparison with the published sequences of P5CS gene by BLAST program, the P5CS gene segment of DPJ showed more than89%identity to that gene from Aegiceras corniculatum, Actinidia deliciosa, Nicotiana tabacum and Solanum lycopersicum. The P5CS gene expression during3℃was investigated through Sq RT-PCR method with the18S gene as reference gene. The results showed that the expression of P5CS was first up-regulated and then down-regulated during low temperature stress. The expressional fastigium of P5CS gene was at the fourth day of3℃treating, while the fastigium of proline content was at the sixth day of3℃treating. Although there was hysteresis phenomenon between the P5CS gene expression and proline synthesis, the trend of them was consilient. But further studies were needed to elucidate the relationship between them.
5. The micropropagation in vitro was investigated in8-year-old cuttings of DPJ by stem segment. Several experiments were carried out to detecte the disinfection of explant, germination culture of axillary buds, the optimum formula of proliferation and induction of multiple shoots and rooting. The results were as follows:
(1) April to September (avoiding the mould rains season) was the best period to pick the explants. The one-year-old and semi-lignification twigs with axillary buds were most suitable to be explants. The treatment with75%ethanol for1min,0.1%mercuric chloride for52mins then washing3times in aseptic water sequentially was the best disinfection method. The optimum basic medium of germination for axillary buds was WPM medium.
(2) In order to find out the optimum formula for axillary buds elongation, the experiments with different plant growth regulators alone or in combination were carried out. The best formula for axillary buds elongation was WPM+2mg·L-1IBA+0.5mg·L-16-BA+0.7mg·L-1GA3. The optimum combination of plant growth regulators for induction of multiple shoots, with generation of11axillary shoots, was1mg·L-1ZT supplemented with0.5mg·L-1IBA,
(3)The best rooting method was that shoots were first dipped in450mg·L-1NAA solution for10mins and cultured in the basal1/2WPM sequentially. The rooting rate was up to86.7%. In this study, the plantlets were acclimatized with78%survival rate under the suitable humidity and illumination during the initial period of acclimation.
引文
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