辣椒紫色性状遗传分析、相关基因克隆和种间杂种创制
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摘要
紫色辣椒因植株茎、叶、花及未成熟果均富含花青素而呈现紫色,具有耐高温、抗干旱和病毒病等综合抗性特征。花青素及其降解产物在人和动物体内能有效地阻止DNA等大分子物质的氧化损害,具有预防、治疗疾病与身体保健等功能。利用紫色辣椒作为亲本材料,通过与绿色辣椒杂交,实现色素功能互补,提高辣椒营养品质,培育适合夏秋季栽培的专用品种,是解决辣椒生产中“夏淡”的有效途径之一。本文研究了紫色辣椒在夏季环境下坐果及光合特性,分析了紫色性状遗传规律,并利用紫色亲本材料创制了紫色的辣椒种间杂种。主要研究内容与结果如下：
1.紫色辣椒种群归类、夏季坐果率以及光合性能
利用RAPD-PCR技术对来自不同生境类型2份紫色和8份绿色辣椒材料的亲缘关系进行研究,结果表明：从45条RAPD随机引物中筛选出34条引物,共扩增出369清晰条带,其中318条带表现多态性,平均多态性比例为86.2%。聚类分析表明,10份辣椒种质在欧氏距离平方为10.5时,能明显地区分为四大类,其中待测紫色辣椒7033、7036与3份C. annuum代表性材料的遗传相似性最大(0.622-0.732),因此应归为辣椒属的C. annuum种。
为了探索紫色辣椒对高温逆境的耐受性,在夏季塑料日光温室中采用人工蕾期授粉法,对18份紫色和21份绿色辣椒进行自交,并选择其中10份紫色和9份绿色材料做完全随机相互杂交,统计坐果率；选择叶片颜色具有代表性的5个辣椒品种,检测其叶片色素含量、主要光合过程指标和叶绿素荧光参数。结果表明：(1)紫色、绿色辣椒平均自交坐果率分别为72.19%和37.12%；以紫色品种为母本、绿色品种为母本平均杂交坐果率分别为55.99%和21.60%,紫色辣椒自交、杂交坐果率都极显著地高于绿色品种；(2)紫色品种叶片花青素(An)含量、叶绿素a、b比值(Ch1.a/b)都显著高于绿色品种,但Chl.b含量两者无差异；多数紫色品种的Chl.a及类胡萝卜素(Car)含量高于绿色品种的；(3)所有辣椒品种的蒸腾速率(Tr)、气孔导度(Gs)及Pn日变化趋势一致,呈现双峰曲线,中午前后存在着气孔与非气孔双重限制。紫色品种净光合速率(Pn)达极大值时的光合有效辐射(PAR)、光补偿点(LCP)和光饱和点(LSP)、在一天中的高温时段实际Pn都显著地高于绿色品种,而表观量子效率(AQY)低于绿色品种,说明紫色品种耐强光、高温的能力高于绿色品种；(4)绿色辣椒只在光系统Ⅱ(PSⅡ)的“反应中心水平”减轻过剩能量的伤害,紫色辣椒除此之外还在“天线水平”防止太阳能被原初过多吸收；与绿色品种相比,紫色辣椒7033和7034所吸收(ABS/CS).捕获(TRo/CS)和传递(ETo/CS)的能量总量较低,但同时耗散(DIo/CS)也低,而最大荧光(Fm)、最大光化学效率(Fv/Frm)、潜在光化学效率(Fv/Fo)和电子传递效率(ETo/TRo)等都很高,致使能量在光合膜上传递和利用经济高效。有望通过杂交手段,在辣椒分离世代中选择出具有适中的叶绿素与花青素含量比值,高ABS/CS、Fm、ETo/TRo、Pn值和低F0值的色素功能互补型植株,实现辣椒在高温、强光和低湿下的高光效育种。
2.紫色辣椒叶片色素的提取和花青素含量遗传规律
采用三因素完全随机设计方法研究了紫色辣椒叶片色素的提取条件,结果表明：最佳浸提组合为30℃、0.1 mol·L-1盐酸处理叶片1h。该色素具有水溶性,在可见光区540 nm处有吸收峰。对其稳定性研究表明：直射光、pH值、Fe3+及Ag+显著地降低其稳定；过氧化氢、亚硫酸钠和亚硝酸钠等氧化剂和还原剂对其减色作用显著,抗坏血酸和淀粉等食品添加剂也具有一定的减色作用。
通过构建紫色与绿色辣椒3个杂交组合的6世代群体(P1、P2、F1、B1、B2和F2),用分光光度法测定辣椒叶片花青素相对含量,研究其遗传规律,结果表明：3个杂交组合的分离群体叶片花青素含量呈现出多峰或单峰偏态分布,表现为主基因+多基因的数量遗传特征；通过多世代联合分析表明,辣椒叶片花青素含量遗传模式符合2对加性-显性-上位性主基因+加性-显性-上位性多基因模型(E-O模型)；F2群体主基因遗传率较高,为78.89%～91.47%,多基因遗传率较低,为5.19%～17.00%。因此对辣椒紫叶性状的遗传选择可在分离早期世代进行。
3.紫色性状的AFLP分子标记和与花青素苷合成相关的基因克隆
基于BSA-AFLP技术,在特紫DNA池中扩增出2条AFLP特异条带E-AC/M-CTC180, E-AC/M-CTC181。对片段进行了回收和测序,序列分析显示,181 bp序列与辣椒C. frutescens栽培种BAC文库的抗烟草花叶病毒病L基因座(FJ597540,FJ597541)、C. annuum栽培种YAC文库的抗细菌性疮痂病的Bs2基因座(AY702979)有76%-83%的同源性；180 bp序列与辣椒C. annuum栽培种BAC文库的长末端重复序列的返转座子部分序列(GU048886)有87%的同源性。
基于RT-PCR技术及同源克隆策略设计引物并克隆了紫色辣椒7033花色苷合成过程的结构基因PAL、CHS、CHI、5GT及调控转录因子R2R3-MYB的部分片段。测序分析表明：获得的这些基因片段长度分别为664 bp、764 bp、579 bp、864 bp和413 bp；紫色辣椒7033的PAL基因片段序列与GenBank中绿色辣椒C. annuum和C. chinense2个栽培种PAL的同源性分别为98%和97%,推测翻译得到的氨基酸序列与前者氨基酸序列完全相同,与后者有2个氨基酸差异；CHS基因片段序列与绿色辣椒C. annuum栽培种相关序列同源性为99%,翻译得到的氨基酸序列与辣椒C. annuum、马铃薯和番茄分别有3、12和12个氨基酸差异；CHI基因片段序列与番茄的PSY 2序列同源性最高为95%,推测有4个氨基酸差异；5GT基因片段序列与番茄、茄子的相关序列同源性均为90%,氨基酸一致性低；R2R3-MYB基因片段序列与已知紫色辣椒C. annuum有99%的同源性,并获得了完全相同的蛋白质序列。为进一步获得参与紫色辣椒花青素苷合成的基因全长及其表达研究奠定了基础。
4.利用胚胎拯救技术创制紫色的辣椒种间杂种(C. annuum×C. chinense)
通过绿色和紫色辣椒正反组合的杂交试验,研究了不同授粉后天数(DAP)及培养基激素浓度对胚胎拯救成功率以及不同配方生根培养基对增殖苗生根的影响,结果表明：在32-34 DAP时能获得胚胎拯救苗,其中MS+8%蔗糖+0.1 mg·L-1GA3+0.05NAA拯救成苗率最高(4.5%～4.84%),其次组合MS+8%蔗糖+0.15 mg·L-1 GA3+0.1NAA也能获得拯救苗(1.43%)。综合分析生根茎段数(率)、生根总数、平均生根数等指标的差异,筛选出实验范围内茎段最适合的生根培养基为1/2MS+0.5 mg·L-1 IBA。由此建立了克服种间杂交受精后种子败育类型的遗传障碍的胚胎拯救适宜方法。
对采用胚胎拯救技术获得的以紫色材料(C. annuum)为母本、绿色辣椒(C. chinense)为父本的F1杂种植株,进行了杂种真实性鉴定。形态学观察发现,杂种一代植株多数性状特别是器官的紫色介于父母本之间；F1代杂种的细胞学研究表明,花粉母细胞减数分裂期染色体的行为发生异常,可育性花粉的比例显著地低于相应的父母本；过氧化物同工酶谱带及RAPD标记分析显示,F1杂种具有部分特异的父本条带；直接测序方法得到的核糖体DNA转录间隔区(ITS)序列表明,与其父母本序列相比较,F1杂种存在186个变异位点,其中139个位点(74.7%)继承了父本序列,23个位点(12.4%)继承了母本序列,其余为碱基颠换/转换。从而由形态学、细胞学、同工酶学及分子生物学等方面的一致性证明了所获得F1杂种植株为真实的种间杂种。
Purple pepper plants have a high resistance to high temperature, drought and virus because of the purple color in the stem, branches, leaves, flowers and immature fruits rich in anthocyanins. The anthocyanins and degraded metabolic products in body of human and animal can effectively prevent the macromolecules such as DNA from damaging by oxidation, thus with functions of the preventing and curing diseases, protecting health and so on. Hybridizing purple pepper, as one of parent materials, with green pepper to achieve complementary function of different pigments, improve the nutritional quality of capsicum fruites and cultivate cultivars suitable for seasons of summer and autumn is one of the efficacious ways to solve the problem of the "short summer". In this paper, fruit setting rate of purple capsicum and the photosynthesis characteristics during summer conditions, inheritance rule of purple trait was investigated, and the accession of purple pepper was utilized as a parent to create purple interspecific hybrids successfully. The main contents and results are as follows:
1. Classifying, fruit setting rate of purple capsicum and the photosynthesis characteristics during summer conditions
Phylogeny of 10 Capsicum accessions including purple varieties 7033,7036 from different habitats was researched using random amplified polymorphic DNA-polymerase chain reaction (RAPD-PCR) technique, the results showed that 369 scored bands were amplified in total with 318 bands having polymorphism amplified with 34 primers screened from 45 RAPD primers in the genome DNA of 10 Capsicum accessions, and the percentage of polymorphic loci was 86.2%. Analysis with SPSS statistical software demonstrated that when the threshold of Squared Euclidean distance was at 10.5, the 10 Capsicum resources could be clustered into 4 groups, C. annuum, C. frutescens, C. chinense and C. baccatum, in which purple peppers 7033,7036 should be classified as C. annuum species of the genus Capsicum for there were similarity maximum, from 0.622 to 0.732, between them and other 3 representative C. annuum accessions.
To delve into resistance to high temperature of purple pepper, an experiment was conducted in a plastic greenhouse to assess fruit setting rate and major photosynthetic and chlorophyll characteristics of purple and green pepper varieties in summer. For self fruit setting studies,18 and 21 purple and green pepper varieties, respectively were hand pollinated at blossom bud stage. Similarly,10 purple and 9 green peppers were randomly selected for reciprocal hybridization to evaluate cross fruit setting rate. Main parameters of photosynthesis and chlorophyll fluorescence were measured from 5 pepper varieties with diverse leaf colours to assess the pigment content. The following results were obtained:(1) significantly higher fruit self-setting rate of purple (72.19%) than the green peppers (37.12%), significantly higher fruit cross-setting rate was recorded. The fruit cross setting rate was 55.99 and 21.60% when purple and green pepper varieties were used as female parents respectively. This indicates stronger tolerance ability of purple pepper variety to summer conditions; (2) the amount anthocyanin (An) and ratio of chlorophyll a and b (Chl. a/b) were significantly higher in leaves of purple peppers than the green ones, amount Chl. a and carotenoid (Car) were also higher in most purple peppers, but insignificantly in amount Chl. b; (3) transpiration rate (Tr), stomatal conductance (Gs) and net photosynthetic rate (Pn) of purple and green pepper showed a similar trend of double peak curve and double stomatal and non stomatal limitation at around noon. Light compensation point (LCP), light saturation point (LSP) and actual Pn during the day time of high temperatures were significantly higher while apparent quantum efficiency (AQY) were lower in all purple varieties than the green ones. Thus the capacity of purple varieties resistance to high light and temperature is higher than the green ones; (4) purple pepper plants could prevent excessive absorption of solar energy by pigment antennas besides by partly reversible inactivation of the reaction center (RC) of photosystemⅡ(PSⅡ) to reduce the harm of excess energy absorbed, the only strategy found in green pepper. The energy of dissipation (DIo/CS) as well as absorption (ABS/CS), capture (TRo/CS) and delivery (ETo/CS) in purple varieties 7033 and 7034 was lower, while the maximum of fluorescence (Fm), maximum of photochemical efficiency (Fv/Fm), potential photochemical efficiency (Fv/Fo) and electron transport efficiency (ETo/TRo) were higher than the green pepper. It can be concluded that energy in photosynthetic membranes of purple peppers was transferred more economically. Therefore, high photosynthetic efficiency varieties that will get high resistance to high temperature, strong light and low humidity may be bred up by hybridizing, selecting the individual plants with the appropriate ratio of chlorophyll to anthocyanin content, higher values of ABS/CS, Fm, ETo/TRo, Pn and lower Fo, and pigment functional complementary in segregating generation.
2. Extraction method of pigment in purple capsicum leaves and inheritance of anthocyanin contents
Completely randomized experiment design was employed to study the effects of temperature, acid concentration and time on extraction of pigment in purple pepper leaves, results showed that optimized combination was that of leaf material soaking in 0.1 mol·L-1 hydrochloric acid for 1 h at temperature 30℃. The pigment was a kind of water soluble pigments and its absorbency peak in visible spectra was observed at wavelength 540nm. Research on pigment stability in purple pepper presented the stability was significantly affected by sunlight, pH and metallic ions of Fe3+ and Ag+, as well as oxidants of hydrogen peroxide and reducers of sodium sulfite and sodium nitrite. To some extent, the stability was also impaired by other food additives such as ascorbic acid and starch.
Three hybridization combinations and their respective six-generations, F1, F2, backcrosses B1 and B2 and their parents P1 and P2, were made with purple and green leaf peppers. Inheritance of anthocyanin contents measured with spectrophotometer was studied through joint analysis of multiple generations. The results showed that relative anthocyanin contents in leaves of segregating populations of three combinations displayed one asymmetry peak or multimodal distribution, indicating that the trait was quantitative in nature and controlled by major gene plus polygene. The heritance of anthocyanin contents in leave of chili pepper fitted the pattern of additive-dominance-epitasis major gene plus additive-dominance-epitasis polygene (E-0 model). The major gene heritability is 78.89%～91.47% and polygene heritability is 5.19%～17.00% in F2 generations in three crosses, which suggested artificial selection to purple leaves trait of pepper could be carried out in the separation of the early generations.
3. AFLP molecular markers related to purple trait and genes cloning of anthocyanin biosynthesis
An extreme purple pepper varieties C. annuum7033 and green C. chinense7020 were used as parents to make F1 and then selfed to build F2 segregated population of leaf colour. Extreme purple and extreme green DNA pools were developed separately using bulked segregant analysis (BSA). AFLP technique with 64 primer combinations were employed to find the polymorphisms between these DNA pools, and in extreme purple DNA pool two specific fragments,181 and 180 bp, were amplified in the primer combination of E-AC/M-CTC. Sequence analyses showed E-AC/M-CTC181 had 76% and 83% homology with the species C. frutescens sequences named FJ597540 and FJ597541, two BAC clones of L gene that confer resistance to tobacco mosaic virus, and 83% homology with the species C. annuum sequence named AY702979, a YAC clone of Bs2 gene that confers resistance to bacterial spot of pepper, respectively. E-AC/M-CTC180 had 87% homology with the species C. annuum sequence named GU048886, a BAC clone of retrotransposon of long terminal repeat (LTR) sequence.
Based on RT-PCR technique and homologous cloning strategy, primers were designed to clone the fragments relevant to anthocyanin biosynthesis in the purple pepper 7033, the structure genes PAL, CHS, CHI,5GT and transcription regulator gene R2R3-MYB, and the fragments length of 664,764,579,864 and 413 bp was obtained by sequencing, respectively. The comparison of sequence homology showed that the gene PAL fragment in purple pepper 7033 had an identity 98% to that of green species C. annuum and 97% to C. chinense in GenBank; the deduced amino acid sequence was identical with that of the former, but two amino acid differences to the latter. The gene CHS sequence had 99% homology to the green pepper C. annuum, and 3,12 and 12 deduced amino acid differences to the pepper C. annuum, potato and tomato, respectively. The gene CHI sequence had a highest homology of 95% to that of tomato PSY 2, but 4 amino acid differences. The gene 5GT sequence had an identity 90% to that of both tomato and eggplant, and low deduced amino acid consistency. The gene R2R3-MYB had 99% homology nucleotide sequence and the identical protein sequence to purple pepper C. annuum reported. The present research laid the foundation of cloning and expression of full-length genes involved in the path of anthocyanin biosynthesis in the purple pepper.
4. Using embryo rescue techniques to create purple Capsicum interspecific hybrids (C. annuum×C. chinense)
Interspecific hybridization experiments between green of C. chinense and purple variety of C. annuum were performed to research the effects of days after pollination (DAP) and hormone concentration on percentage of plants survived by embryo rescuing, and different culture mediums on stem segment rooting. The results showed that in treatments of 32～34 DAP hybrid embryos were rescued successfully. Number/percentage, 4.5%～4.84% of plants survived was highest using MS+8% sucrose+0.1 mg-L"1 GA3+ 0.05 NAA as rescuing medium, followed by the combination of MS+8% sucrose+0.15 mg·L-1 GA3+0.1 NAA,1.43%. Comprehensively analyzing indices of number/percentage of stem segment rooting, total number of roots, average of roots each stem segment, the most suitable medium for rooting,1/2 MS+0.5 mg·L-1 IBA, was selected. Thus, a suitable method of embryo rescue to overcome seed abortion after fertilization, one type of genetic barriers, was made.
Hybridity identifying to the putative F1 plant obtained through embryo rescue technique, the purple accession (C. annuum) as female and the green (C. chinense) as male parent. Morphological observations revealed that most traits of F1 hybrids were intermediate to both parents especially the organic purple colour. Cytological investigation of F1 hybrid exhibited abnormal chromosome behaviour at meiotic stage of pollen mother cells (PMCs). Pollen grains stainability percentage was significantly lower than the corresponding parents. Both peroxidase isozyme and random amplified polymorphic DNA profiles of F1 hybrid possessed specific bands of the male parent. The nuclear ribosomal DNA internal transcribed spacer (ITS) region sequences from putative F1 and the possible parents obtained by direct sequencing method showed that 186 variable loci.139 loci (74.7%) and 23 loci (12.4%) of F1 hybrid were inherited from male and female parents, respectively. The consistency in morphological characteristics, cytological features, isozyme banding patterns and ITS region sequences thus confirmed hybridity of F1 plant.
1.紫色辣椒种群归类、夏季坐果率以及光合性能
利用RAPD-PCR技术对来自不同生境类型2份紫色和8份绿色辣椒材料的亲缘关系进行研究,结果表明：从45条RAPD随机引物中筛选出34条引物,共扩增出369清晰条带,其中318条带表现多态性,平均多态性比例为86.2%。聚类分析表明,10份辣椒种质在欧氏距离平方为10.5时,能明显地区分为四大类,其中待测紫色辣椒7033、7036与3份C. annuum代表性材料的遗传相似性最大(0.622-0.732),因此应归为辣椒属的C. annuum种。
为了探索紫色辣椒对高温逆境的耐受性,在夏季塑料日光温室中采用人工蕾期授粉法,对18份紫色和21份绿色辣椒进行自交,并选择其中10份紫色和9份绿色材料做完全随机相互杂交,统计坐果率；选择叶片颜色具有代表性的5个辣椒品种,检测其叶片色素含量、主要光合过程指标和叶绿素荧光参数。结果表明：(1)紫色、绿色辣椒平均自交坐果率分别为72.19%和37.12%；以紫色品种为母本、绿色品种为母本平均杂交坐果率分别为55.99%和21.60%,紫色辣椒自交、杂交坐果率都极显著地高于绿色品种；(2)紫色品种叶片花青素(An)含量、叶绿素a、b比值(Ch1.a/b)都显著高于绿色品种,但Chl.b含量两者无差异；多数紫色品种的Chl.a及类胡萝卜素(Car)含量高于绿色品种的；(3)所有辣椒品种的蒸腾速率(Tr)、气孔导度(Gs)及Pn日变化趋势一致,呈现双峰曲线,中午前后存在着气孔与非气孔双重限制。紫色品种净光合速率(Pn)达极大值时的光合有效辐射(PAR)、光补偿点(LCP)和光饱和点(LSP)、在一天中的高温时段实际Pn都显著地高于绿色品种,而表观量子效率(AQY)低于绿色品种,说明紫色品种耐强光、高温的能力高于绿色品种；(4)绿色辣椒只在光系统Ⅱ(PSⅡ)的“反应中心水平”减轻过剩能量的伤害,紫色辣椒除此之外还在“天线水平”防止太阳能被原初过多吸收；与绿色品种相比,紫色辣椒7033和7034所吸收(ABS/CS).捕获(TRo/CS)和传递(ETo/CS)的能量总量较低,但同时耗散(DIo/CS)也低,而最大荧光(Fm)、最大光化学效率(Fv/Frm)、潜在光化学效率(Fv/Fo)和电子传递效率(ETo/TRo)等都很高,致使能量在光合膜上传递和利用经济高效。有望通过杂交手段,在辣椒分离世代中选择出具有适中的叶绿素与花青素含量比值,高ABS/CS、Fm、ETo/TRo、Pn值和低F0值的色素功能互补型植株,实现辣椒在高温、强光和低湿下的高光效育种。
2.紫色辣椒叶片色素的提取和花青素含量遗传规律
采用三因素完全随机设计方法研究了紫色辣椒叶片色素的提取条件,结果表明：最佳浸提组合为30℃、0.1 mol·L-1盐酸处理叶片1h。该色素具有水溶性,在可见光区540 nm处有吸收峰。对其稳定性研究表明：直射光、pH值、Fe3+及Ag+显著地降低其稳定；过氧化氢、亚硫酸钠和亚硝酸钠等氧化剂和还原剂对其减色作用显著,抗坏血酸和淀粉等食品添加剂也具有一定的减色作用。
通过构建紫色与绿色辣椒3个杂交组合的6世代群体(P1、P2、F1、B1、B2和F2),用分光光度法测定辣椒叶片花青素相对含量,研究其遗传规律,结果表明：3个杂交组合的分离群体叶片花青素含量呈现出多峰或单峰偏态分布,表现为主基因+多基因的数量遗传特征；通过多世代联合分析表明,辣椒叶片花青素含量遗传模式符合2对加性-显性-上位性主基因+加性-显性-上位性多基因模型(E-O模型)；F2群体主基因遗传率较高,为78.89%～91.47%,多基因遗传率较低,为5.19%～17.00%。因此对辣椒紫叶性状的遗传选择可在分离早期世代进行。
3.紫色性状的AFLP分子标记和与花青素苷合成相关的基因克隆
基于BSA-AFLP技术,在特紫DNA池中扩增出2条AFLP特异条带E-AC/M-CTC180, E-AC/M-CTC181。对片段进行了回收和测序,序列分析显示,181 bp序列与辣椒C. frutescens栽培种BAC文库的抗烟草花叶病毒病L基因座(FJ597540,FJ597541)、C. annuum栽培种YAC文库的抗细菌性疮痂病的Bs2基因座(AY702979)有76%-83%的同源性；180 bp序列与辣椒C. annuum栽培种BAC文库的长末端重复序列的返转座子部分序列(GU048886)有87%的同源性。
基于RT-PCR技术及同源克隆策略设计引物并克隆了紫色辣椒7033花色苷合成过程的结构基因PAL、CHS、CHI、5GT及调控转录因子R2R3-MYB的部分片段。测序分析表明：获得的这些基因片段长度分别为664 bp、764 bp、579 bp、864 bp和413 bp；紫色辣椒7033的PAL基因片段序列与GenBank中绿色辣椒C. annuum和C. chinense2个栽培种PAL的同源性分别为98%和97%,推测翻译得到的氨基酸序列与前者氨基酸序列完全相同,与后者有2个氨基酸差异；CHS基因片段序列与绿色辣椒C. annuum栽培种相关序列同源性为99%,翻译得到的氨基酸序列与辣椒C. annuum、马铃薯和番茄分别有3、12和12个氨基酸差异；CHI基因片段序列与番茄的PSY 2序列同源性最高为95%,推测有4个氨基酸差异；5GT基因片段序列与番茄、茄子的相关序列同源性均为90%,氨基酸一致性低；R2R3-MYB基因片段序列与已知紫色辣椒C. annuum有99%的同源性,并获得了完全相同的蛋白质序列。为进一步获得参与紫色辣椒花青素苷合成的基因全长及其表达研究奠定了基础。
4.利用胚胎拯救技术创制紫色的辣椒种间杂种(C. annuum×C. chinense)
通过绿色和紫色辣椒正反组合的杂交试验,研究了不同授粉后天数(DAP)及培养基激素浓度对胚胎拯救成功率以及不同配方生根培养基对增殖苗生根的影响,结果表明：在32-34 DAP时能获得胚胎拯救苗,其中MS+8%蔗糖+0.1 mg·L-1GA3+0.05NAA拯救成苗率最高(4.5%～4.84%),其次组合MS+8%蔗糖+0.15 mg·L-1 GA3+0.1NAA也能获得拯救苗(1.43%)。综合分析生根茎段数(率)、生根总数、平均生根数等指标的差异,筛选出实验范围内茎段最适合的生根培养基为1/2MS+0.5 mg·L-1 IBA。由此建立了克服种间杂交受精后种子败育类型的遗传障碍的胚胎拯救适宜方法。
对采用胚胎拯救技术获得的以紫色材料(C. annuum)为母本、绿色辣椒(C. chinense)为父本的F1杂种植株,进行了杂种真实性鉴定。形态学观察发现,杂种一代植株多数性状特别是器官的紫色介于父母本之间；F1代杂种的细胞学研究表明,花粉母细胞减数分裂期染色体的行为发生异常,可育性花粉的比例显著地低于相应的父母本；过氧化物同工酶谱带及RAPD标记分析显示,F1杂种具有部分特异的父本条带；直接测序方法得到的核糖体DNA转录间隔区(ITS)序列表明,与其父母本序列相比较,F1杂种存在186个变异位点,其中139个位点(74.7%)继承了父本序列,23个位点(12.4%)继承了母本序列,其余为碱基颠换/转换。从而由形态学、细胞学、同工酶学及分子生物学等方面的一致性证明了所获得F1杂种植株为真实的种间杂种。
Purple pepper plants have a high resistance to high temperature, drought and virus because of the purple color in the stem, branches, leaves, flowers and immature fruits rich in anthocyanins. The anthocyanins and degraded metabolic products in body of human and animal can effectively prevent the macromolecules such as DNA from damaging by oxidation, thus with functions of the preventing and curing diseases, protecting health and so on. Hybridizing purple pepper, as one of parent materials, with green pepper to achieve complementary function of different pigments, improve the nutritional quality of capsicum fruites and cultivate cultivars suitable for seasons of summer and autumn is one of the efficacious ways to solve the problem of the "short summer". In this paper, fruit setting rate of purple capsicum and the photosynthesis characteristics during summer conditions, inheritance rule of purple trait was investigated, and the accession of purple pepper was utilized as a parent to create purple interspecific hybrids successfully. The main contents and results are as follows:
1. Classifying, fruit setting rate of purple capsicum and the photosynthesis characteristics during summer conditions
Phylogeny of 10 Capsicum accessions including purple varieties 7033,7036 from different habitats was researched using random amplified polymorphic DNA-polymerase chain reaction (RAPD-PCR) technique, the results showed that 369 scored bands were amplified in total with 318 bands having polymorphism amplified with 34 primers screened from 45 RAPD primers in the genome DNA of 10 Capsicum accessions, and the percentage of polymorphic loci was 86.2%. Analysis with SPSS statistical software demonstrated that when the threshold of Squared Euclidean distance was at 10.5, the 10 Capsicum resources could be clustered into 4 groups, C. annuum, C. frutescens, C. chinense and C. baccatum, in which purple peppers 7033,7036 should be classified as C. annuum species of the genus Capsicum for there were similarity maximum, from 0.622 to 0.732, between them and other 3 representative C. annuum accessions.
To delve into resistance to high temperature of purple pepper, an experiment was conducted in a plastic greenhouse to assess fruit setting rate and major photosynthetic and chlorophyll characteristics of purple and green pepper varieties in summer. For self fruit setting studies,18 and 21 purple and green pepper varieties, respectively were hand pollinated at blossom bud stage. Similarly,10 purple and 9 green peppers were randomly selected for reciprocal hybridization to evaluate cross fruit setting rate. Main parameters of photosynthesis and chlorophyll fluorescence were measured from 5 pepper varieties with diverse leaf colours to assess the pigment content. The following results were obtained:(1) significantly higher fruit self-setting rate of purple (72.19%) than the green peppers (37.12%), significantly higher fruit cross-setting rate was recorded. The fruit cross setting rate was 55.99 and 21.60% when purple and green pepper varieties were used as female parents respectively. This indicates stronger tolerance ability of purple pepper variety to summer conditions; (2) the amount anthocyanin (An) and ratio of chlorophyll a and b (Chl. a/b) were significantly higher in leaves of purple peppers than the green ones, amount Chl. a and carotenoid (Car) were also higher in most purple peppers, but insignificantly in amount Chl. b; (3) transpiration rate (Tr), stomatal conductance (Gs) and net photosynthetic rate (Pn) of purple and green pepper showed a similar trend of double peak curve and double stomatal and non stomatal limitation at around noon. Light compensation point (LCP), light saturation point (LSP) and actual Pn during the day time of high temperatures were significantly higher while apparent quantum efficiency (AQY) were lower in all purple varieties than the green ones. Thus the capacity of purple varieties resistance to high light and temperature is higher than the green ones; (4) purple pepper plants could prevent excessive absorption of solar energy by pigment antennas besides by partly reversible inactivation of the reaction center (RC) of photosystemⅡ(PSⅡ) to reduce the harm of excess energy absorbed, the only strategy found in green pepper. The energy of dissipation (DIo/CS) as well as absorption (ABS/CS), capture (TRo/CS) and delivery (ETo/CS) in purple varieties 7033 and 7034 was lower, while the maximum of fluorescence (Fm), maximum of photochemical efficiency (Fv/Fm), potential photochemical efficiency (Fv/Fo) and electron transport efficiency (ETo/TRo) were higher than the green pepper. It can be concluded that energy in photosynthetic membranes of purple peppers was transferred more economically. Therefore, high photosynthetic efficiency varieties that will get high resistance to high temperature, strong light and low humidity may be bred up by hybridizing, selecting the individual plants with the appropriate ratio of chlorophyll to anthocyanin content, higher values of ABS/CS, Fm, ETo/TRo, Pn and lower Fo, and pigment functional complementary in segregating generation.
2. Extraction method of pigment in purple capsicum leaves and inheritance of anthocyanin contents
Completely randomized experiment design was employed to study the effects of temperature, acid concentration and time on extraction of pigment in purple pepper leaves, results showed that optimized combination was that of leaf material soaking in 0.1 mol·L-1 hydrochloric acid for 1 h at temperature 30℃. The pigment was a kind of water soluble pigments and its absorbency peak in visible spectra was observed at wavelength 540nm. Research on pigment stability in purple pepper presented the stability was significantly affected by sunlight, pH and metallic ions of Fe3+ and Ag+, as well as oxidants of hydrogen peroxide and reducers of sodium sulfite and sodium nitrite. To some extent, the stability was also impaired by other food additives such as ascorbic acid and starch.
Three hybridization combinations and their respective six-generations, F1, F2, backcrosses B1 and B2 and their parents P1 and P2, were made with purple and green leaf peppers. Inheritance of anthocyanin contents measured with spectrophotometer was studied through joint analysis of multiple generations. The results showed that relative anthocyanin contents in leaves of segregating populations of three combinations displayed one asymmetry peak or multimodal distribution, indicating that the trait was quantitative in nature and controlled by major gene plus polygene. The heritance of anthocyanin contents in leave of chili pepper fitted the pattern of additive-dominance-epitasis major gene plus additive-dominance-epitasis polygene (E-0 model). The major gene heritability is 78.89%～91.47% and polygene heritability is 5.19%～17.00% in F2 generations in three crosses, which suggested artificial selection to purple leaves trait of pepper could be carried out in the separation of the early generations.
3. AFLP molecular markers related to purple trait and genes cloning of anthocyanin biosynthesis
An extreme purple pepper varieties C. annuum7033 and green C. chinense7020 were used as parents to make F1 and then selfed to build F2 segregated population of leaf colour. Extreme purple and extreme green DNA pools were developed separately using bulked segregant analysis (BSA). AFLP technique with 64 primer combinations were employed to find the polymorphisms between these DNA pools, and in extreme purple DNA pool two specific fragments,181 and 180 bp, were amplified in the primer combination of E-AC/M-CTC. Sequence analyses showed E-AC/M-CTC181 had 76% and 83% homology with the species C. frutescens sequences named FJ597540 and FJ597541, two BAC clones of L gene that confer resistance to tobacco mosaic virus, and 83% homology with the species C. annuum sequence named AY702979, a YAC clone of Bs2 gene that confers resistance to bacterial spot of pepper, respectively. E-AC/M-CTC180 had 87% homology with the species C. annuum sequence named GU048886, a BAC clone of retrotransposon of long terminal repeat (LTR) sequence.
Based on RT-PCR technique and homologous cloning strategy, primers were designed to clone the fragments relevant to anthocyanin biosynthesis in the purple pepper 7033, the structure genes PAL, CHS, CHI,5GT and transcription regulator gene R2R3-MYB, and the fragments length of 664,764,579,864 and 413 bp was obtained by sequencing, respectively. The comparison of sequence homology showed that the gene PAL fragment in purple pepper 7033 had an identity 98% to that of green species C. annuum and 97% to C. chinense in GenBank; the deduced amino acid sequence was identical with that of the former, but two amino acid differences to the latter. The gene CHS sequence had 99% homology to the green pepper C. annuum, and 3,12 and 12 deduced amino acid differences to the pepper C. annuum, potato and tomato, respectively. The gene CHI sequence had a highest homology of 95% to that of tomato PSY 2, but 4 amino acid differences. The gene 5GT sequence had an identity 90% to that of both tomato and eggplant, and low deduced amino acid consistency. The gene R2R3-MYB had 99% homology nucleotide sequence and the identical protein sequence to purple pepper C. annuum reported. The present research laid the foundation of cloning and expression of full-length genes involved in the path of anthocyanin biosynthesis in the purple pepper.
4. Using embryo rescue techniques to create purple Capsicum interspecific hybrids (C. annuum×C. chinense)
Interspecific hybridization experiments between green of C. chinense and purple variety of C. annuum were performed to research the effects of days after pollination (DAP) and hormone concentration on percentage of plants survived by embryo rescuing, and different culture mediums on stem segment rooting. The results showed that in treatments of 32～34 DAP hybrid embryos were rescued successfully. Number/percentage, 4.5%～4.84% of plants survived was highest using MS+8% sucrose+0.1 mg-L"1 GA3+ 0.05 NAA as rescuing medium, followed by the combination of MS+8% sucrose+0.15 mg·L-1 GA3+0.1 NAA,1.43%. Comprehensively analyzing indices of number/percentage of stem segment rooting, total number of roots, average of roots each stem segment, the most suitable medium for rooting,1/2 MS+0.5 mg·L-1 IBA, was selected. Thus, a suitable method of embryo rescue to overcome seed abortion after fertilization, one type of genetic barriers, was made.
Hybridity identifying to the putative F1 plant obtained through embryo rescue technique, the purple accession (C. annuum) as female and the green (C. chinense) as male parent. Morphological observations revealed that most traits of F1 hybrids were intermediate to both parents especially the organic purple colour. Cytological investigation of F1 hybrid exhibited abnormal chromosome behaviour at meiotic stage of pollen mother cells (PMCs). Pollen grains stainability percentage was significantly lower than the corresponding parents. Both peroxidase isozyme and random amplified polymorphic DNA profiles of F1 hybrid possessed specific bands of the male parent. The nuclear ribosomal DNA internal transcribed spacer (ITS) region sequences from putative F1 and the possible parents obtained by direct sequencing method showed that 186 variable loci.139 loci (74.7%) and 23 loci (12.4%) of F1 hybrid were inherited from male and female parents, respectively. The consistency in morphological characteristics, cytological features, isozyme banding patterns and ITS region sequences thus confirmed hybridity of F1 plant.
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