马铃薯野生种Solanum berthaultii抗低温糖化基因的分离及表达特征分析
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摘要
为实现马铃薯全年供应,同时减少采后病虫害、块茎萌发和失水等带来的损失,用于加工的马铃薯通常需要储藏在4℃左右的低温环境中,但这样又带来另一个严重的负面影响—低温糖化(Cold-induced sweetening, CIS),即块茎低温储藏过程中,易发生糖化,出现淀粉含量下降、还原糖积累的现象。发生糖化的块茎在油炸加工过程中,还原糖和游离氨基酸发生Maillard反应,以致出现褐色变化、口感变差,并产生具有神经毒性的致癌物质丙烯酰胺,严重降低了加工产品的品质,给马铃薯种植业带来巨大经济损失。马铃薯品种改良是控制这一危害的重要途径。
不同马铃薯基因型发生低温糖化的程度具有差异,特别是一些野生种,具有较强的抗低温糖化的能力。因此,了解马铃薯低温糖化的调节机制和发掘低温糖化的关键基因,对马铃薯加工品质的改良具有重要意义。
本研究选用抗低温糖化的野生种Solanum berthaultii (accession CW2-1, ber)和不抗低温糖化的栽培品种(S. tuberosum)鄂马铃薯3号(E3)为材料,筛选块茎低温应答基因,进而通过在不同抗性材料中的基因表达谱分析,获得与低温糖化相关的基因。研究取得的主要结果如下：
1.不同抗低温糖化马铃薯的炸片色泽指数和可溶性糖含量对低温的响应
为评价储藏温度对不同块茎炸片色泽、还原糖以及蔗糖含量的影响,本研究分别分析了马铃薯ber和E3块茎在4℃和20℃储藏0、3、5、10、15、20、30、45、60、75和90d时相关指标的变化情况。结果表明,在20℃储藏时块茎炸片色泽和糖含量变化较小。总体来说,低温储藏时,块茎还原糖和蔗糖含量随着储藏时间延长而不断增加,尽管在储藏后期有小幅下降,但ber块茎的还原糖含量明显低于E3块茎,蔗糖含量则高于E3块茎,而且还原糖含量和炸片色泽之间呈显著线性正相关。表明野生种ber相对于栽培种E3而言具有较高的抗低温糖化能力。
2.反向差减杂交文库的构建
为进一步分析ber低温反应及抗低温糖化的机理,本研究在杨建文(2005)构建的ber正向低温差减杂交文库的基础上,分别以4℃和20℃储藏5d的ber块茎为Driver和Tester,构建反向差减杂交文库。在经过两轮差减杂交和选择性PCR扩增之后,将获得的差减杂交产物连入pBlueScript SK(-),通过Amp/X-gal/IPTG筛选获得1920个白色单斑,经过进一步PCR扩增筛选,最终获得由1584个有效克隆构成的反向SSH文库,克隆中的插入片段长度约为500bp。
3.ber块茎低温反应基因的确定
通过PCR扩增获得ber正向文库中2112条插入序列和反向文库中1584条插入序列,纯化后定量,作为点制芯片的探针。提取ber块茎4℃和20℃储藏5d后的总RNA,逆转录时分别用Cy5和Cy3进行标记,并与cDNA芯片进行杂交,共获得736个差异表达基因,其中663个上调,73个下调。差异表达基因序列分析时共有719个克隆获得良好测序结果。经序列比对、合并重复后,共获得188个代表不同基因的非重复序列片段,其中上调表达序列138个,下调表达序列50个。经分析,149个已知功能的差异表达基因可被归为14种功能类型,主要与细胞自救与防御、代谢、能量、蛋白命运等细胞过程相关。本研究首次对马铃薯低温响应的差异表达基因进行了系统分析,为进一步研究低温糖化机制奠定了基础。
4.块茎低温糖化相关基因的筛选
以抗低温糖化的ber块茎和易低温糖化的E3块茎为材料,通过qRT-PCR方法全面分析上述188个差异表达基因在4℃和20℃储藏时动态表达变化。结果表明,这些基因中的绝大部分在两种块茎中表现出相同的调节模式,反映了不同基因型块茎低温应答机制具有同一性；但是仍有一些基因的表达模式或转录水平在两种基因型块茎中存在较大差异,初步推测这些“特异表达基因”的表达强弱,决定了不同基因型块茎间低温糖化水平的差异。例如,低温储藏时,ber C20-3-O14(GWD)和C20-6-K02(木糖异构酶)基因的转录被抑制,而E3块茎则为上调表达,并且表达量均高于ber块茎；ber和E3块茎C20-2-D03(BMY7)和C20-4-C15(InvInh)基因具有明显的低温诱导特性,但her BMY7转录水平低于E3,而ber InvInh则高于E3；另外,低温下ber和E3块茎C20-6-F06(GAPDH)和C20-3-A14(丙酮酸激酶)基因都呈先上调后下调的表达模式,但这两个基因在ber块茎中的累积量始终高于E3块茎。因此,与E3块茎相比,ber低温储藏时很可能具有较低的淀粉分解率和较高的糖酵解速率,且ber蔗糖向还原糖转化受到限制。因此,根据本实验结果和已有资料,我们推测导致ber块茎抗低温糖化特征的分子机制,主要与淀粉分解、蔗糖分解和糖酵解三种代谢途径相关。进一步深入研究这些基因的功能,将对马铃薯低温糖化机制提供新的依据,并为马铃薯品质的基因工程改良提供理论指导。
In order to prolonging the storage for year-round processing of potato tubers, and reduce the loss caused by microorganism infection, sprout growth, and water loss after harvest, low temperature (around4℃) storage is commonly used. However, the cold-induced sweetening (CIS) characterized by accumulation of reducing sugars in the tubers often occurs. As potato tubers are processed into chips and fries, reducing sugars react with amino acids to generate unacceptable dark-colored, bitter-tasting products in the non-enzymatic Maillard reaction, as well as accumulating acrylamide-a known neurotoxin and suspected carcinogen. CIS has posed a significant challenge to potato industry and is of commercial interest.
Studies suggest that the level of CIS varies between potato cultivars, some wild species exhibit high level resistance to CIS. Understanding the mechanism of CIS through identifying the regulative pathways and associated genes should be critical for improving potato processing quality.
In present study, two potato species, a CIS-resistant wild type species Solanum berthaultii (accession CW2-1, ber) and a CIS-sensitive cultivator (S. tuberosum) E-potato-3(E3), were employed. Differentially expressed genes responsible to low temperature were classified and the genes involved in CIS were elucidated. The main results are as following.
1. Response of potato genotypes with distinct resistance to CIS to low temperature in terms of variation in chip color index and soluble sugar contents
To determine the effects of storage temperature on chip color and contests of reducing sugars and sucrose, ber and E3tubers were stored at4℃and20℃, and sampled at0,3,5,10,15,20,30,45,60,75and90d, respectively. The results indicated that both ber and E3tubers displayed little variation when stored at20℃. However, dramatic variations were observed when tubers stored at4℃. Generally, the reducing sugars and sucrose contents increased along with the time course although there were some declines at the end of the storage. Comparing between the two potato genotypes, ber had a lower reducing sugar and a higher sucrose contents than E3. Importantly, there was a positive linear relationship between the reducing sugar content and the chip color index, suggesting that the wild potato species ber was more resistant to CIS than E3.
2. Construction of the reverse suppression subtractive hybridization (SSH) library
In order to identify the genes associated with CIS, a forward SSH libraries of ber tubers subjected to cold stimulation was constructed previously by Yang (2005). A reverse SSH library was constructed in the present study. ber tubers stored at4℃and20℃for5d were sampled and used as Driver and Tester, respectively. The special expressed sequences were obtained by subtraction and selective PCR amplification, and ligated into the pBlueScript SK (-), then transformed into Escherichia coli DH5a competent cells. The transformed E. coli cells were selected by Amp/X-gal/IPTG and PCR. A total of1920white-single colonies were selected. After PCR screening, a total of1584valid clones (a single amplicon around500bp in length) were contained in the reverse SSH library.
3. Identification of differentially expressed (DE) genes responsive to low temperature
A total of3696inserts, including2112clones from the forward library and1584from the reverse library, were amplified by PCR and purified, then qualified to ensure adequate and equal PCR products. The amplicons were printed onto glass slides to produce the microarrays. Total RNAs from ber tubers stored at4℃and20℃for5d were prepared and reverse transcribed separately in presence of Cy5-and Cy3-labeled dUTP. The hybridization was preformed using the labeled cDNAs. Finally,736cold-related clones were identified, of which663were putatively up-regulated and73down-regulated. Then the inserted fragments were sequenced and719clones produced good results. A total of188non-redundant sequences were identified by comparative analyses against the GenBank database, including138down-regulated and50up-regulated genes. One hundred and forty-nine genes of known function were classified into14functional categories. These functional genes were mostly related to cell rescue, defense and virulence, metabolism, energy and protein fate, representing various processes of plant defense against abiotic stresses. These results present an extensive investigation of the DE genes of tubers in response to cold stress, providing a basis in understanding possible mechanisms of CIS.
4. Screening of the genes associated with CIS of potato tubers
Profiling of the188DE genes as mentioned above in CIS-resistant ber tubers and in CIS-sensitive E3tubers were further investigated by qRT-PCR, tubers were stored at4℃and sampled at20℃for0,5,15and30d, respectively. The results indicated that most of the genes had nearly the same expression patterns in ber and E3tubers, indicating that tubers with different genotypes shared similar cold responses mechanisms. However, some "special genes", which might determine the degree of CIS, were found differentially regulated in ber and E3tubers. During cold storage, transcripts of C20-3-O14(GWD) and C20-6-K02(xylose isomerase) were suppressed in ber tubers, but they were induced in E3tubers, the transcripts accumulation of the two genes were less in ber tubers than in E3. C20-2-D03(BMY7) and C20-4-C15(InvInh) were cold induced genes in ber and E3tubers, but ber tubers had more transcripts accumulation of InvInh and less BMY7than E3. Transcriptional patterns of C20-6-F06(GAPDH) and C20-3-A14(pyruvate kinase) were first up-regulated and then down-regulated in the cold stored ber and E3tubers. However, both the two genes had higher transcriptional levels in ber tubers than in E3. Comparing to E3tubers, ber might have lower amylolysis level but higer glycolysis level under cold conditions, so the sucrose decomposition might be limited in ber tubers. Together, it was hypothesized that amylolysis, sucrose decomposition and glycolysis pathways might be three key-determinants of CIS. Further investigation of these cold-regulated genes will deepen our understanding of the regulatory mechanisms of potato CIS and direct approaches for the genetic improvement of potato processing quality.
不同马铃薯基因型发生低温糖化的程度具有差异,特别是一些野生种,具有较强的抗低温糖化的能力。因此,了解马铃薯低温糖化的调节机制和发掘低温糖化的关键基因,对马铃薯加工品质的改良具有重要意义。
本研究选用抗低温糖化的野生种Solanum berthaultii (accession CW2-1, ber)和不抗低温糖化的栽培品种(S. tuberosum)鄂马铃薯3号(E3)为材料,筛选块茎低温应答基因,进而通过在不同抗性材料中的基因表达谱分析,获得与低温糖化相关的基因。研究取得的主要结果如下：
1.不同抗低温糖化马铃薯的炸片色泽指数和可溶性糖含量对低温的响应
为评价储藏温度对不同块茎炸片色泽、还原糖以及蔗糖含量的影响,本研究分别分析了马铃薯ber和E3块茎在4℃和20℃储藏0、3、5、10、15、20、30、45、60、75和90d时相关指标的变化情况。结果表明,在20℃储藏时块茎炸片色泽和糖含量变化较小。总体来说,低温储藏时,块茎还原糖和蔗糖含量随着储藏时间延长而不断增加,尽管在储藏后期有小幅下降,但ber块茎的还原糖含量明显低于E3块茎,蔗糖含量则高于E3块茎,而且还原糖含量和炸片色泽之间呈显著线性正相关。表明野生种ber相对于栽培种E3而言具有较高的抗低温糖化能力。
2.反向差减杂交文库的构建
为进一步分析ber低温反应及抗低温糖化的机理,本研究在杨建文(2005)构建的ber正向低温差减杂交文库的基础上,分别以4℃和20℃储藏5d的ber块茎为Driver和Tester,构建反向差减杂交文库。在经过两轮差减杂交和选择性PCR扩增之后,将获得的差减杂交产物连入pBlueScript SK(-),通过Amp/X-gal/IPTG筛选获得1920个白色单斑,经过进一步PCR扩增筛选,最终获得由1584个有效克隆构成的反向SSH文库,克隆中的插入片段长度约为500bp。
3.ber块茎低温反应基因的确定
通过PCR扩增获得ber正向文库中2112条插入序列和反向文库中1584条插入序列,纯化后定量,作为点制芯片的探针。提取ber块茎4℃和20℃储藏5d后的总RNA,逆转录时分别用Cy5和Cy3进行标记,并与cDNA芯片进行杂交,共获得736个差异表达基因,其中663个上调,73个下调。差异表达基因序列分析时共有719个克隆获得良好测序结果。经序列比对、合并重复后,共获得188个代表不同基因的非重复序列片段,其中上调表达序列138个,下调表达序列50个。经分析,149个已知功能的差异表达基因可被归为14种功能类型,主要与细胞自救与防御、代谢、能量、蛋白命运等细胞过程相关。本研究首次对马铃薯低温响应的差异表达基因进行了系统分析,为进一步研究低温糖化机制奠定了基础。
4.块茎低温糖化相关基因的筛选
以抗低温糖化的ber块茎和易低温糖化的E3块茎为材料,通过qRT-PCR方法全面分析上述188个差异表达基因在4℃和20℃储藏时动态表达变化。结果表明,这些基因中的绝大部分在两种块茎中表现出相同的调节模式,反映了不同基因型块茎低温应答机制具有同一性；但是仍有一些基因的表达模式或转录水平在两种基因型块茎中存在较大差异,初步推测这些“特异表达基因”的表达强弱,决定了不同基因型块茎间低温糖化水平的差异。例如,低温储藏时,ber C20-3-O14(GWD)和C20-6-K02(木糖异构酶)基因的转录被抑制,而E3块茎则为上调表达,并且表达量均高于ber块茎；ber和E3块茎C20-2-D03(BMY7)和C20-4-C15(InvInh)基因具有明显的低温诱导特性,但her BMY7转录水平低于E3,而ber InvInh则高于E3；另外,低温下ber和E3块茎C20-6-F06(GAPDH)和C20-3-A14(丙酮酸激酶)基因都呈先上调后下调的表达模式,但这两个基因在ber块茎中的累积量始终高于E3块茎。因此,与E3块茎相比,ber低温储藏时很可能具有较低的淀粉分解率和较高的糖酵解速率,且ber蔗糖向还原糖转化受到限制。因此,根据本实验结果和已有资料,我们推测导致ber块茎抗低温糖化特征的分子机制,主要与淀粉分解、蔗糖分解和糖酵解三种代谢途径相关。进一步深入研究这些基因的功能,将对马铃薯低温糖化机制提供新的依据,并为马铃薯品质的基因工程改良提供理论指导。
In order to prolonging the storage for year-round processing of potato tubers, and reduce the loss caused by microorganism infection, sprout growth, and water loss after harvest, low temperature (around4℃) storage is commonly used. However, the cold-induced sweetening (CIS) characterized by accumulation of reducing sugars in the tubers often occurs. As potato tubers are processed into chips and fries, reducing sugars react with amino acids to generate unacceptable dark-colored, bitter-tasting products in the non-enzymatic Maillard reaction, as well as accumulating acrylamide-a known neurotoxin and suspected carcinogen. CIS has posed a significant challenge to potato industry and is of commercial interest.
Studies suggest that the level of CIS varies between potato cultivars, some wild species exhibit high level resistance to CIS. Understanding the mechanism of CIS through identifying the regulative pathways and associated genes should be critical for improving potato processing quality.
In present study, two potato species, a CIS-resistant wild type species Solanum berthaultii (accession CW2-1, ber) and a CIS-sensitive cultivator (S. tuberosum) E-potato-3(E3), were employed. Differentially expressed genes responsible to low temperature were classified and the genes involved in CIS were elucidated. The main results are as following.
1. Response of potato genotypes with distinct resistance to CIS to low temperature in terms of variation in chip color index and soluble sugar contents
To determine the effects of storage temperature on chip color and contests of reducing sugars and sucrose, ber and E3tubers were stored at4℃and20℃, and sampled at0,3,5,10,15,20,30,45,60,75and90d, respectively. The results indicated that both ber and E3tubers displayed little variation when stored at20℃. However, dramatic variations were observed when tubers stored at4℃. Generally, the reducing sugars and sucrose contents increased along with the time course although there were some declines at the end of the storage. Comparing between the two potato genotypes, ber had a lower reducing sugar and a higher sucrose contents than E3. Importantly, there was a positive linear relationship between the reducing sugar content and the chip color index, suggesting that the wild potato species ber was more resistant to CIS than E3.
2. Construction of the reverse suppression subtractive hybridization (SSH) library
In order to identify the genes associated with CIS, a forward SSH libraries of ber tubers subjected to cold stimulation was constructed previously by Yang (2005). A reverse SSH library was constructed in the present study. ber tubers stored at4℃and20℃for5d were sampled and used as Driver and Tester, respectively. The special expressed sequences were obtained by subtraction and selective PCR amplification, and ligated into the pBlueScript SK (-), then transformed into Escherichia coli DH5a competent cells. The transformed E. coli cells were selected by Amp/X-gal/IPTG and PCR. A total of1920white-single colonies were selected. After PCR screening, a total of1584valid clones (a single amplicon around500bp in length) were contained in the reverse SSH library.
3. Identification of differentially expressed (DE) genes responsive to low temperature
A total of3696inserts, including2112clones from the forward library and1584from the reverse library, were amplified by PCR and purified, then qualified to ensure adequate and equal PCR products. The amplicons were printed onto glass slides to produce the microarrays. Total RNAs from ber tubers stored at4℃and20℃for5d were prepared and reverse transcribed separately in presence of Cy5-and Cy3-labeled dUTP. The hybridization was preformed using the labeled cDNAs. Finally,736cold-related clones were identified, of which663were putatively up-regulated and73down-regulated. Then the inserted fragments were sequenced and719clones produced good results. A total of188non-redundant sequences were identified by comparative analyses against the GenBank database, including138down-regulated and50up-regulated genes. One hundred and forty-nine genes of known function were classified into14functional categories. These functional genes were mostly related to cell rescue, defense and virulence, metabolism, energy and protein fate, representing various processes of plant defense against abiotic stresses. These results present an extensive investigation of the DE genes of tubers in response to cold stress, providing a basis in understanding possible mechanisms of CIS.
4. Screening of the genes associated with CIS of potato tubers
Profiling of the188DE genes as mentioned above in CIS-resistant ber tubers and in CIS-sensitive E3tubers were further investigated by qRT-PCR, tubers were stored at4℃and sampled at20℃for0,5,15and30d, respectively. The results indicated that most of the genes had nearly the same expression patterns in ber and E3tubers, indicating that tubers with different genotypes shared similar cold responses mechanisms. However, some "special genes", which might determine the degree of CIS, were found differentially regulated in ber and E3tubers. During cold storage, transcripts of C20-3-O14(GWD) and C20-6-K02(xylose isomerase) were suppressed in ber tubers, but they were induced in E3tubers, the transcripts accumulation of the two genes were less in ber tubers than in E3. C20-2-D03(BMY7) and C20-4-C15(InvInh) were cold induced genes in ber and E3tubers, but ber tubers had more transcripts accumulation of InvInh and less BMY7than E3. Transcriptional patterns of C20-6-F06(GAPDH) and C20-3-A14(pyruvate kinase) were first up-regulated and then down-regulated in the cold stored ber and E3tubers. However, both the two genes had higher transcriptional levels in ber tubers than in E3. Comparing to E3tubers, ber might have lower amylolysis level but higer glycolysis level under cold conditions, so the sucrose decomposition might be limited in ber tubers. Together, it was hypothesized that amylolysis, sucrose decomposition and glycolysis pathways might be three key-determinants of CIS. Further investigation of these cold-regulated genes will deepen our understanding of the regulatory mechanisms of potato CIS and direct approaches for the genetic improvement of potato processing quality.
引文
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