番茄种质遗传多样性及种子质量调控技术研究
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摘要
茄科的番茄(Solanum lycopersicum L.)是全世界广泛种植的重要经济作物之一在人们的日常饮食和营养中具有重要作用。针对目前我国番茄育种和种子生产过程中存在的主要问题,如育种中经常使用少数骨干亲本,使番茄品种的遗传基础趋于狭窄；番茄种子质量差,杂交种种子纯度检测技术较为落后、种子活力低下等,本研究利用SSR分子标记对不同类型的番茄自交系进行遗传多样性研究,探索构建番茄杂交种子纯度快速鉴定的分子技术体系和利用种子引发技术提高番茄种子活力的方法,为番茄育种和生产服务。取得的主要研究结果如下：
1、以105核心种质为试验材料,在正常、盐、干旱胁迫下对番茄种子活力进行鉴定评价。结果表明,在正常条件下发芽势(GP)、发芽率(GR)变幅分别为0～98%、0～98%；在10%PEG干旱胁迫下GP、GR变幅分别为0～98%、0～98%；在100mM NaCl胁迫下GP、GR变幅分别为0%～52%、0～66%。在105份种质资源中,GP、GR在品种间存在较大的差异。在正常、干旱、盐胁迫条件下,绝大部分品种GP分别为80～100%、80～100%、0%,分别占42.86%、47.6%、69.5%；而绝大部分品种GR分别为80～100%、80～100%、<10%,分别占55.23%、60.95%、63.80%。本研究还筛选出在正常条件下、干旱、盐胁迫下高种子活力品种,在正常条件下,高活力种子(GR、 GP>90%)品种有20个,占19%；在10%PEG胁迫下,高活力种子(GR、GP>90%)品种分别有22个,占20.9%；在100mM NaCl胁迫下,高活力种子(GR、GP>50%)品种分别有4个,占3.8%。这些番茄种质资源的利用有利于高活力番茄品种的培育,及为今后基因定位克隆研究奠定基础。
2、利用35对SSR分子标记对来自中国、日本、韩国和美国的39份无限生长型和有限生长型番茄种质的遗传多样性进行研究。结果表明,35对引物共扩增出150个条带,平均每对引物扩增出4.3个条带,平均多态性信息含量(PIC)为0.31。聚类分析研究表明在遗传相似系数0.80处将39份材料分成4组。来自美国的有限生长型番茄自交系之间最具有遗传相似性,而来自美国的有限生长型自交系Us-16和来自日本的无限生长型自交系Ja-2遗传相似性最低。中国的番茄自交系具有较高的遗传相似性(平均遗传相似系数为0.909)、其次是日本(平均遗传相似系数为0.76)和韩国(平均遗传相似系数为0.80)的番茄自交系。聚类结果与番茄生态型和生长类型分类相一致,研究结果为今后番茄育种提供重要信息。
3、利用35对SSR分子标记对生产上广泛种植的番茄杂交种“ZZ1”进行种子纯度的分子鉴定研究,结果表明,有16对SSR引物在双亲中表现多态性,其中7对引物能表现明显的共显性。进一步选择扩增条带清晰、结果稳定的3对引物(AI778183、 SSR136、LEat014)对人工配置的不同批次和不同纯度的番茄杂交种“ZZ1”的种子样品进行分子水平上的检测,同时在田间进行种植鉴定,结果发现,两种方法鉴定的结果基本一致,误差在1%以下。该分子检测技术已经申请国家专利,可以作为番茄杂交种种子纯度鉴定的一种快速、简单和可靠性好的方法在种子生产和销售中应用,并可探索其在更多番茄品种上的应用。
4、引发作为种子播前处理技术,可以提高逆境条件下种子活力。本研究探讨了渗调引发对番茄杂交种子在老化和盐胁迫下种子活力的影响。番茄杂交种“ZZ1”种子分别在自然条件(老化)和-2℃条件下(未老化)贮藏4年,利用10%(w/v)聚乙二醇(PEG)在20±1℃黑暗条件下引发处理2d。老化种子在正常条件下(蒸馏水)25±1℃发芽7d,分析引发对老化种子活力的影响。结果表明,与未引发老化种子相比,引发能显著提高老化种子发芽势(GP)、发芽率(GR)、发芽指数(GI)、,提高幼苗根长(RL)、芽长(SL)、幼苗总鲜重(FW),缩短平均发芽时间(MGT)；其种子活力与未老化种子的活力相近。同时,未老化种子在100mM NaCl胁迫条件下25±+1℃发芽7d,分析引发对盐胁迫下种子活力的影响,结果表明与未引发种子相比,引发能显著提高GP、GI、FW,显著降低MGT,但GR、RL、SL两者无显著差异；在田间条件下,引发能显著提高直播番茄成苗率、幼苗RL、SL、FW。进一步研究表明,引发处理能显著降低种子吸胀期脂质过氧化。与未引发处理种子相比,引发能显著降低老化和盐胁迫下种子吸胀期细胞膜相对电导率(REL)、丙二醛含量(MDA)。相关分析表明MDA、REL与种子活力呈显著负相关,表明引发提高胁迫条件下种子活力可能与降低种子脂质过氧化相关。
Tomato(Solanum lycopersicum L.) as one of the important economic crops is widely cultivated all over the world, which plays an important role in people's daily diet and nutrition intake. Several problems are exsited in the current process of tomato breeding and seed production. For example, the genetic basis of tomato varieties are narrow because only a small number of key breeding parents used in breeding, and the level of hybrid seed purity testing and the seed vigor of varieties is low. Therefore, the SSR technology was used to analysis the genetic diversity in inbred lines of different tomato types in this study, the SSR technology was used to build a rapid tomato seed purity testing system, and the seed priming technology was used to improve the seed vigor. The main results obtained are as follows:
1. Seed vigor of105core collection was identified under normal, salt and drought stress conditions in tomato. The results showed that the amplitude of germination potential (GP) and germination rate (GR) were0to98%and0to98%under normal conditions; the amplitude of GP and GR were0to98%and0to98%under10%PEG drought stress conditions; the amplitude of GP and GR were0to52%and0to66%under100mM NaC1stress conditions. There were significant differences in GP and GR between varieties. The GP of most varieties were80to100%,80to100%and0%, respectively, the percentages were42.86%,47.6%and69.5%of total germplasm under normal, drought and salt stress conditions. However, the GR of most varieties were80to100%,80to100%and<10%, respectively, the percentages were55.23%,60.95%and63.80%of total germplasm under normal, drought and salt stress conditions. The varieties with high seed vigor were also screened in this study. For example,20varieties with high vigor seeds (GR, GP>90%) were screened and the percentage was19%of total germplasm under normal conditions;22varieties with high vigor seeds (GR, GP>90%) were screened and the percentage was20.9%of total germplasm under10%PEG drought stress conditions;4varieties with high vigor seeds (GR, GP>50%) were screened and the percentage was3.8%of total germplasm under100mM NaC1stress conditions. These tomato germplasm will benefit to the cultivation of tomato varieties with high seed vigor and to gene mapping and cloning in future study.
2. A study was conducted to determine the genetic diversity of39determinate and indeterminate tomato inbred lines collected from China, Japan, S. Korea, and USA. Using35SSR polymorphic markers, a total of150alleles were found with moderate levels of diversity, and a high number of unique alleles existing in these tomato lines. The mean number of alleles per locus was4.3and the average polymorphism information content (PIC) was0.31. Unweighted Pair Group Method with Arithmetic Mean (UPGMA) clustering at genetic similarity value of0.85grouped the inbred lines into four groups, where one USA cultivar formed a separate and more distant cluster. The most similar inbred lines are from USA, both with determinate type, whereas the most different lines are from USA (Us-16) and Japan (Ja-2) with determinate and indeterminate growth habit, respectively. Clustering was consistent with the known information regarding geographical location and growth habit. The genetic distance information reported in this study might be used by breeders when planning future crosses among these inbred lines.
3.35SSR markers were used to screen the polymorphic and co-dominant markers among tomato hybrids and their parents. The results indicated that16SSR markers showed polymorphic in the parents in which7SSR markers were co-dominant performance. The co-dominant SSR markers (AI778183、SSR136、LEat014) that could amplify clear and stable bands were selected to identify the accuracy of SSR testing in different artificial configuration purity of tomato samples, and the testing of field also was conducted. The similar results were found between the SSR and field testing with error about1%. The SSR testing system of seed purity will be broadly applied in future with the advantages of rapid and accurate identification in tomato.
4. Priming is a pre-sowing treatment that improves seed vigor under stress conditions. This study investigates the effect of osmopriming on seed vigor under aging and salinity stress of a tomato hybrid. Tomato seeds of the ZZ1hybrid variety, stored for four years under natural (aged) or-20℃(unaged) conditions, were primed in10%(w/v) polyethylene glycol (PEG) solution for2d at20±1℃in the dark. The vigor of primed, aged seeds under normal condition (water) at25±1℃for7d was evaluated. The results showed that the germination potential (GP), germination rate (GR) and germination index (GI) of primed, aged seeds were significantly enhanced, with a substantial increase in the radicle length (RL), shoot length (SL) and total fresh weight (FW) and a reduction of the mean germination time (MGT) compared with unprimed, aged seeds. The vigor of aged seeds was comparable to that of unaged seeds due to the priming treatment. Furthermore, the vigor of primed, unaged seeds was evaluated under the100mM NaCl condition at25±1℃for7d. Similarly, the GP, GI and FW significantly increased in primed seeds compared with unprimed seeds under salinity stress. In contrast, MGT significantly decreased, while GR, RL and SL did not show significant differences. Under the field condition, the priming of tomato seeds significantly improved the stand seedling establishment and the seedlings from primed seeds had greater RL, SL, FW than the un-primed seeds. Among priming treatments, further results showed that osmopriming could reduce seed lipid peroxidation. A decline in the relative electrolyte leakage (REL) and in malondialdehyde (MDA) was detected in primed seeds during the imbibition stage compared with unprimed seeds under aging and salinity stress. Similarly, there were negative correlations between seed vigor and REL and MDA, which suggests that seed priming improves seed vigor under stress conditions associated with a decrease in lipid peroxidation.
1、以105核心种质为试验材料,在正常、盐、干旱胁迫下对番茄种子活力进行鉴定评价。结果表明,在正常条件下发芽势(GP)、发芽率(GR)变幅分别为0～98%、0～98%；在10%PEG干旱胁迫下GP、GR变幅分别为0～98%、0～98%；在100mM NaCl胁迫下GP、GR变幅分别为0%～52%、0～66%。在105份种质资源中,GP、GR在品种间存在较大的差异。在正常、干旱、盐胁迫条件下,绝大部分品种GP分别为80～100%、80～100%、0%,分别占42.86%、47.6%、69.5%；而绝大部分品种GR分别为80～100%、80～100%、<10%,分别占55.23%、60.95%、63.80%。本研究还筛选出在正常条件下、干旱、盐胁迫下高种子活力品种,在正常条件下,高活力种子(GR、 GP>90%)品种有20个,占19%；在10%PEG胁迫下,高活力种子(GR、GP>90%)品种分别有22个,占20.9%；在100mM NaCl胁迫下,高活力种子(GR、GP>50%)品种分别有4个,占3.8%。这些番茄种质资源的利用有利于高活力番茄品种的培育,及为今后基因定位克隆研究奠定基础。
2、利用35对SSR分子标记对来自中国、日本、韩国和美国的39份无限生长型和有限生长型番茄种质的遗传多样性进行研究。结果表明,35对引物共扩增出150个条带,平均每对引物扩增出4.3个条带,平均多态性信息含量(PIC)为0.31。聚类分析研究表明在遗传相似系数0.80处将39份材料分成4组。来自美国的有限生长型番茄自交系之间最具有遗传相似性,而来自美国的有限生长型自交系Us-16和来自日本的无限生长型自交系Ja-2遗传相似性最低。中国的番茄自交系具有较高的遗传相似性(平均遗传相似系数为0.909)、其次是日本(平均遗传相似系数为0.76)和韩国(平均遗传相似系数为0.80)的番茄自交系。聚类结果与番茄生态型和生长类型分类相一致,研究结果为今后番茄育种提供重要信息。
3、利用35对SSR分子标记对生产上广泛种植的番茄杂交种“ZZ1”进行种子纯度的分子鉴定研究,结果表明,有16对SSR引物在双亲中表现多态性,其中7对引物能表现明显的共显性。进一步选择扩增条带清晰、结果稳定的3对引物(AI778183、 SSR136、LEat014)对人工配置的不同批次和不同纯度的番茄杂交种“ZZ1”的种子样品进行分子水平上的检测,同时在田间进行种植鉴定,结果发现,两种方法鉴定的结果基本一致,误差在1%以下。该分子检测技术已经申请国家专利,可以作为番茄杂交种种子纯度鉴定的一种快速、简单和可靠性好的方法在种子生产和销售中应用,并可探索其在更多番茄品种上的应用。
4、引发作为种子播前处理技术,可以提高逆境条件下种子活力。本研究探讨了渗调引发对番茄杂交种子在老化和盐胁迫下种子活力的影响。番茄杂交种“ZZ1”种子分别在自然条件(老化)和-2℃条件下(未老化)贮藏4年,利用10%(w/v)聚乙二醇(PEG)在20±1℃黑暗条件下引发处理2d。老化种子在正常条件下(蒸馏水)25±1℃发芽7d,分析引发对老化种子活力的影响。结果表明,与未引发老化种子相比,引发能显著提高老化种子发芽势(GP)、发芽率(GR)、发芽指数(GI)、,提高幼苗根长(RL)、芽长(SL)、幼苗总鲜重(FW),缩短平均发芽时间(MGT)；其种子活力与未老化种子的活力相近。同时,未老化种子在100mM NaCl胁迫条件下25±+1℃发芽7d,分析引发对盐胁迫下种子活力的影响,结果表明与未引发种子相比,引发能显著提高GP、GI、FW,显著降低MGT,但GR、RL、SL两者无显著差异；在田间条件下,引发能显著提高直播番茄成苗率、幼苗RL、SL、FW。进一步研究表明,引发处理能显著降低种子吸胀期脂质过氧化。与未引发处理种子相比,引发能显著降低老化和盐胁迫下种子吸胀期细胞膜相对电导率(REL)、丙二醛含量(MDA)。相关分析表明MDA、REL与种子活力呈显著负相关,表明引发提高胁迫条件下种子活力可能与降低种子脂质过氧化相关。
Tomato(Solanum lycopersicum L.) as one of the important economic crops is widely cultivated all over the world, which plays an important role in people's daily diet and nutrition intake. Several problems are exsited in the current process of tomato breeding and seed production. For example, the genetic basis of tomato varieties are narrow because only a small number of key breeding parents used in breeding, and the level of hybrid seed purity testing and the seed vigor of varieties is low. Therefore, the SSR technology was used to analysis the genetic diversity in inbred lines of different tomato types in this study, the SSR technology was used to build a rapid tomato seed purity testing system, and the seed priming technology was used to improve the seed vigor. The main results obtained are as follows:
1. Seed vigor of105core collection was identified under normal, salt and drought stress conditions in tomato. The results showed that the amplitude of germination potential (GP) and germination rate (GR) were0to98%and0to98%under normal conditions; the amplitude of GP and GR were0to98%and0to98%under10%PEG drought stress conditions; the amplitude of GP and GR were0to52%and0to66%under100mM NaC1stress conditions. There were significant differences in GP and GR between varieties. The GP of most varieties were80to100%,80to100%and0%, respectively, the percentages were42.86%,47.6%and69.5%of total germplasm under normal, drought and salt stress conditions. However, the GR of most varieties were80to100%,80to100%and<10%, respectively, the percentages were55.23%,60.95%and63.80%of total germplasm under normal, drought and salt stress conditions. The varieties with high seed vigor were also screened in this study. For example,20varieties with high vigor seeds (GR, GP>90%) were screened and the percentage was19%of total germplasm under normal conditions;22varieties with high vigor seeds (GR, GP>90%) were screened and the percentage was20.9%of total germplasm under10%PEG drought stress conditions;4varieties with high vigor seeds (GR, GP>50%) were screened and the percentage was3.8%of total germplasm under100mM NaC1stress conditions. These tomato germplasm will benefit to the cultivation of tomato varieties with high seed vigor and to gene mapping and cloning in future study.
2. A study was conducted to determine the genetic diversity of39determinate and indeterminate tomato inbred lines collected from China, Japan, S. Korea, and USA. Using35SSR polymorphic markers, a total of150alleles were found with moderate levels of diversity, and a high number of unique alleles existing in these tomato lines. The mean number of alleles per locus was4.3and the average polymorphism information content (PIC) was0.31. Unweighted Pair Group Method with Arithmetic Mean (UPGMA) clustering at genetic similarity value of0.85grouped the inbred lines into four groups, where one USA cultivar formed a separate and more distant cluster. The most similar inbred lines are from USA, both with determinate type, whereas the most different lines are from USA (Us-16) and Japan (Ja-2) with determinate and indeterminate growth habit, respectively. Clustering was consistent with the known information regarding geographical location and growth habit. The genetic distance information reported in this study might be used by breeders when planning future crosses among these inbred lines.
3.35SSR markers were used to screen the polymorphic and co-dominant markers among tomato hybrids and their parents. The results indicated that16SSR markers showed polymorphic in the parents in which7SSR markers were co-dominant performance. The co-dominant SSR markers (AI778183、SSR136、LEat014) that could amplify clear and stable bands were selected to identify the accuracy of SSR testing in different artificial configuration purity of tomato samples, and the testing of field also was conducted. The similar results were found between the SSR and field testing with error about1%. The SSR testing system of seed purity will be broadly applied in future with the advantages of rapid and accurate identification in tomato.
4. Priming is a pre-sowing treatment that improves seed vigor under stress conditions. This study investigates the effect of osmopriming on seed vigor under aging and salinity stress of a tomato hybrid. Tomato seeds of the ZZ1hybrid variety, stored for four years under natural (aged) or-20℃(unaged) conditions, were primed in10%(w/v) polyethylene glycol (PEG) solution for2d at20±1℃in the dark. The vigor of primed, aged seeds under normal condition (water) at25±1℃for7d was evaluated. The results showed that the germination potential (GP), germination rate (GR) and germination index (GI) of primed, aged seeds were significantly enhanced, with a substantial increase in the radicle length (RL), shoot length (SL) and total fresh weight (FW) and a reduction of the mean germination time (MGT) compared with unprimed, aged seeds. The vigor of aged seeds was comparable to that of unaged seeds due to the priming treatment. Furthermore, the vigor of primed, unaged seeds was evaluated under the100mM NaCl condition at25±1℃for7d. Similarly, the GP, GI and FW significantly increased in primed seeds compared with unprimed seeds under salinity stress. In contrast, MGT significantly decreased, while GR, RL and SL did not show significant differences. Under the field condition, the priming of tomato seeds significantly improved the stand seedling establishment and the seedlings from primed seeds had greater RL, SL, FW than the un-primed seeds. Among priming treatments, further results showed that osmopriming could reduce seed lipid peroxidation. A decline in the relative electrolyte leakage (REL) and in malondialdehyde (MDA) was detected in primed seeds during the imbibition stage compared with unprimed seeds under aging and salinity stress. Similarly, there were negative correlations between seed vigor and REL and MDA, which suggests that seed priming improves seed vigor under stress conditions associated with a decrease in lipid peroxidation.
引文
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