楸子不同类型抗旱耐盐性评价及遗传差异分析
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摘要
楸子（Malus prunifolia）是我国特有的苹果野生种质资源，对不同自然条件有很强的适应性，与苹果嫁接亲和力强，长势旺盛，是我国西北地区重要的苹果砧木。在我国经长期天然杂交，加上基因突变及环境因素的影响，使得楸子产生很多不同的类型（变种）。有些类型在形态上极为相似，以至于很难将其区分，这对楸子不同类型的命名造成一定的困难，对其在分类学上的划分产生较大争议。此外，楸子作为一种野生种质资源，其不同类型之间的抗逆性，还未曾有过系统地分析和评价。本研究通过分析干旱、盐胁迫下各类型的生理响应，对楸子不同类型的抗旱、耐盐性做出综合评价，筛选出不同抗性的苹果砧木资源，这对于苹果抗性育种研究以及经济栽培具有重要的理论和实用价值；同时，利用SSR分子标记技术，结合聚类分析及主成分分析，对楸子不同类型之间的亲缘关系在分子水平上做进一步鉴定，为阐明苹果砧木资源的分类、优化种质资源的收集与管理工作提供理论依据。主要研究结果如下：
1.将‘富士’嫁接于8种苹果砧木（7个楸子类型，1个新疆野苹果）后进行12d干旱处理，其叶片相对含水量（RWC）与对照相比（0d）显著降低，总叶绿素含量与光合参数在胁迫过程中也有不同程度的下降。胁迫第9d，以东北黄海棠、新疆野苹果和富平楸子为砧木的‘富士’仍然保持相对较高的Pn，分别为8.48，7.55和6.34μmol m-2s-1，而以红海棠和白海棠为砧木的‘富士’的Pn则分别下降到2.54和3.82μmol m-2s-1。此时，以富平楸子为砧木的‘富士’的WUEi最高，达到5.71μmol mmol-1，WUEi最小的是以红海棠作为砧木的‘富士’，只有2.42μmol mmol-1。
2.干旱胁迫下，所有植株的APX活性在胁迫前3d急剧上升，在第9d开始下降，不同砧木的‘富士’所达到的最大值不同。以红海棠、五棱海棠和白海棠为砧木的‘富士’的POD活性在胁迫后3d便开始下降，而以富平楸子、东北黄海棠、新疆野苹果以及崂山柰子为砧木的‘富士’的POD活性仍有上升趋势。在胁迫初期，以红海棠和白海棠为砧木的‘富士’的CAT活性上升较快，但并没有持续太长时间；而以富平楸子、东北黄海棠和新疆野苹果为砧木的‘富士’的CAT活性在胁迫初期上升较为缓慢，但第3d以后，活性突然升高，并在第6d达到最大值。
3.干旱胁迫12d后，所有植株叶片的膜透性以及丙二醛（MDA）含量与对照相比均有不同程度的升高。抗旱性较弱的红海棠和白海棠的相对电导率与对照相比分别高出46.37%和45.86%，丙二醛含量与对照相比分别增加了30.04%和31.92%。抗旱性较强的砧木，如富平楸子、东北黄海棠以及新疆野苹果，其相对电导率与对照相比升高了23.33～32.94%；丙二醛含量与对照相比升高了22.06～23.94%。聚类分析将8种砧木的抗旱性分为三类，抗旱性强的砧木有富平楸子、东北黄海棠和新疆野苹果；中等抗旱的砧木为吴起楸子、五棱海棠以及崂山柰子；抗旱性相对较弱的是红海棠和白海棠。
4.用150mM NaCl处理11种苹果砧木（9个楸子类型，另外2种分别为新疆野苹果和平邑甜茶）10d，统计盐害指数（SI）显示：处理3d后，红海棠、白海棠、冬白果和内蒙古海红较早地表现出了盐害症状（SI≥0.10）；在处理后第10d，富平楸子、东北黄海棠、崂山柰子、新疆野苹果和平邑甜茶的盐害指数相对较低，而白海棠、五棱海棠和内蒙古海红，它们的盐害指数相对较高，分别达到0.519、0.509和0.625。
5.通过比较净光合速率（Pn）、相对电导率变化百分比（VP）、相对含水量（RWC），结合盐害指数以及生物量抗逆系数（RAC）的聚类分析结果，11种砧木的耐盐性被分为三大类，耐盐性强的砧木包括富平楸子、东北黄海棠、崂山柰子、新疆野苹果和平邑甜茶；耐盐性中等的砧木有吴起楸子和冬白果；耐盐性弱的砧木分别为红海棠、白海棠、五棱海棠及内蒙古海红。除红海棠之外，聚类分析的结果与统计盐害指数的结果一致。
6.利用降落PCR结合SSR分子标记技术，对15个楸子类型进行遗传差异分析。10对SSR引物所检测出的170条谱带中，有156条为多态性带，多态性谱带比率达到90.74%，平均遗传相似系数（GC）为0.660，GC最高的是富平楸子与吴起楸子，为0.882；其遗传距离（GD）也只有0.125；冬白果与河北热滚子的GC最低，只有0.529，但GD却达到0.636。
7.利用非加权算数平均法（UPGMA）在遗传相似值0.650处，15个楸子类型被分为三大类。这与主成分分析（PCA）的结果一致。聚类分析表明，多数聚在一起的楸子类型与它们的地理分布相一致，但也有部分个体的DNA谱带与其起源地不一致，5个起源于新疆的类型就被分散在不同类群，这在一定程度上反映了楸子不同类型复杂的遗传背景。
Malus prunifolia is an endemic wild apple germplasm resource in our country withstrong adaptability to different natural conditions. It is also an important apple rootstock forgood grafting affinity ability and widely used in northern and northwestern China. Thisdiverse species has formed many biotypes (variant) as long-term natural hybridization,genetic mutation and environmental factors influence. Some biotypes are so similar inphenotypic that it is difficult to distinguish them from each other. All these lead to itsnomenclature is confusing and the taxonomy is fraught with problems. On the other hand, nocomprehensive and systematic evaluation has been made of its drought and salt tolerance.Therefore, in our study, through explored the physiological response and evaluated thetolerance in different biotypes of M. prunifolia under drought and salt stress, it provide animportant theoretical and practical value for apple stress breeding and cultivation. Meanwhile,we used SSR markers in combination with cluster and principal coordinate analysis toevaluate the genetic relationship among biotypes of M. prunifolia; the findings are helpful tooptimize the collection and management of apple germplasm resources, and also provide atheoretical reference for traditional morphological classifications of Malus prunifolia at themolecular level.
The main results were as follows:
1. Eight apple rootstocks (seven biotypes of M. prunifolia and M. sieversii) grafted with‘Fuji’ subjected to drought stress for12d. Compared with the control (0d), the leaf relativewater content (RWC) values decreased significantly in different rootstocks, and the amount ofleaves total chlorophyll as well as photosynthesis parameters also declined to varying degrees.M. prunifolia ‘dongbeihuanghaitang’, M. sieversii, and M. prunifolia ‘fupingqiuzi’ stillmaintained relatively high Pn (8.48,7.55, and6.34μmol m-2s-1, respectively) after9d stress.Two more-sensitive biotypes of M. prunifolia—‘honghaitang’ and ‘baihaitang’—which hadthe lowest Pn values, at2.54and3.82μmol m-2s-1, respectively. Moreover, WUEiwas thehighest,5.71, in the more-resistant M. prunifolia ‘fupingqiuzi’, compared with the lowestvalue,2.42, for M. prunifolia ‘honghaitang’.
2. The activities of ascorbate peroxidase (APX), peroxidase (POD), and catalase (CAT)changed significantly during drought treatment. For APX activity, all biotypes had a sharpincrease during the first3d and began to decline after9d stress. However, the maximumactivity varied by biotypes. The POD activity began to drop in M. prunifolia ‘honghaitang’, M.prunifolia ‘wulenghaitang’, and M. prunifolia ‘baihaitang’ after3d, but M. prunifolia‘fupingqiuzi’, M. prunifolia ‘dongbeihuanghaitang’, M. prunifolia ‘laoshannaizi’ and M.sieversii still maintanted an upward trend. CAT activity raised quickly in M. prunifolia—‘honghaitang’ and ‘baihaitang’ at the stress started, but it did not maintant very long;however, M. prunifolia ‘fupingqiuzi’, M. prunifolia ‘dongbeihuanghaitang’, and M. sieversiiincreased slowly during the first3d treatment, then raised at a faster rate and reached thepeak values on6d.
3. All treated plants had elevated electrolyte leakage (EL) and malondialdehyde (MDA)contents after12d of stress. Compared with their corresponding control, values for EL inless-tolerant M. prunifolia ‘honghaitang’ and ‘baihaitang’ increased by46.37%and45.86%,MDA contents increased by30.04%and31.92%; Such damage was not as severe amonghighly drought-tolerant rootstocks, e.g., M. prunifolia ‘fupingqiuzi’, M. prunifolia‘dongbeihuanghaitang’, and M. sieversii, which EL values increased by23.33～32.94%andMDA contents increased by22.06～23.94%compared with the control. Cluster analysis cangroup these Malus biotypes into three categories. The high tolerance included two M.prunifolia—‘fupingqiuzi’ and ‘dongbeihuanghaitang’—plus M. sieversii Roem; moderatetolerance were three M. prunifolia—‘wuqiqiuzi’,‘wulenghaitang’, and ‘laoshannaizi’; lowtolerance were two M. prunifolia—‘honghaitang’ and ‘baihaitang’.
4. Eleven apple rootstocks (nine biotypes of Malus prunifolia plus M. sieversii and M.hupehensis) were exposed to150mM NaCl for10d. Salt injury index (SI) showed that, fourbiotypes of M. prunifolia—‘honghaitang’,‘baihaitang’,‘dongbaiguo’, and‘neimengguhaihong’ were more susceptible than other biotypes after3days of exposure (SI≥0.10); after10d stress, M. prunifolia ‘fupingqiuzi’, M. prunifolia ‘dongbeihuanghaitang’, andM. prunifolia ‘laoshannaizi’, as well as plants of M. sieversii and M. hupehensis, hadrelatively low SI. However, three more sensitive biotypes of M. prunifolia—‘baihaitang’,‘wulenghaitang’, and ‘neimengguhaihong’—that had higher SI values of0.519,0.509, and0.625, respectively.
5. By comparison with net photosynthetic rate (Pn), variation percentage (VP) ofelectrolyte leakage, relative water content (RWC), SI and biomass adverse resistancecoefficients (ARC), we grouped these biotypes into three classes. The high tolerance included:three for M. prunifolia—‘fupingqiuzi’,‘dongbeihuanghaitang’ and ‘laoshannaizi’—plus M. sieversii and M. hupehensis; the moderate tolerance were: M. prunifolia ‘wuqiqiuzi’ and‘dongbaiguo’; the low tolerance were four for M. prunifolia—‘honghaitang’,‘baihaitang’,‘wulenghaitang’, and ‘neimengguhaihong’. Except for M. prunifolia ‘honghaitang’, all ofthese groupings followed a pattern similar to that found with SI values.
6. We used10pairs of simple sequence repeat (SSR) primers in combination withtouchdown PCR to evaluate the genetic relationships among15biotypes of Malus prunifolia.Out of170detected fragments,156(90.74%) were polymorphic. The average geneticsimilarity coefficient (GC) between pairs of biotypes was0.66. The greatest GC (0.88) wasfound between ‘fupingqiuzi’ and ‘wuqiqiuzi’, while the biotype pairing of ‘dongbaiguo’ and‘hebeiregunzi’ showed the least GC (0.53).
7. Cluster analysis sorted all biotypes into three groups, which was then confirmed byprincipal component analysis (PCA). A dendrogram showed that some of the clusteredbiotypes were largely congruent for geographical distribution. However, the DNA patterns forsome biotype groups did not demonstrate relative agreement in their origin. Five biotypesarising from the Xinjiang Autonomous Region were dispersed within the dendrogram,reflecting their complex genetic backgrounds.
1.将‘富士’嫁接于8种苹果砧木（7个楸子类型，1个新疆野苹果）后进行12d干旱处理，其叶片相对含水量（RWC）与对照相比（0d）显著降低，总叶绿素含量与光合参数在胁迫过程中也有不同程度的下降。胁迫第9d，以东北黄海棠、新疆野苹果和富平楸子为砧木的‘富士’仍然保持相对较高的Pn，分别为8.48，7.55和6.34μmol m-2s-1，而以红海棠和白海棠为砧木的‘富士’的Pn则分别下降到2.54和3.82μmol m-2s-1。此时，以富平楸子为砧木的‘富士’的WUEi最高，达到5.71μmol mmol-1，WUEi最小的是以红海棠作为砧木的‘富士’，只有2.42μmol mmol-1。
2.干旱胁迫下，所有植株的APX活性在胁迫前3d急剧上升，在第9d开始下降，不同砧木的‘富士’所达到的最大值不同。以红海棠、五棱海棠和白海棠为砧木的‘富士’的POD活性在胁迫后3d便开始下降，而以富平楸子、东北黄海棠、新疆野苹果以及崂山柰子为砧木的‘富士’的POD活性仍有上升趋势。在胁迫初期，以红海棠和白海棠为砧木的‘富士’的CAT活性上升较快，但并没有持续太长时间；而以富平楸子、东北黄海棠和新疆野苹果为砧木的‘富士’的CAT活性在胁迫初期上升较为缓慢，但第3d以后，活性突然升高，并在第6d达到最大值。
3.干旱胁迫12d后，所有植株叶片的膜透性以及丙二醛（MDA）含量与对照相比均有不同程度的升高。抗旱性较弱的红海棠和白海棠的相对电导率与对照相比分别高出46.37%和45.86%，丙二醛含量与对照相比分别增加了30.04%和31.92%。抗旱性较强的砧木，如富平楸子、东北黄海棠以及新疆野苹果，其相对电导率与对照相比升高了23.33～32.94%；丙二醛含量与对照相比升高了22.06～23.94%。聚类分析将8种砧木的抗旱性分为三类，抗旱性强的砧木有富平楸子、东北黄海棠和新疆野苹果；中等抗旱的砧木为吴起楸子、五棱海棠以及崂山柰子；抗旱性相对较弱的是红海棠和白海棠。
4.用150mM NaCl处理11种苹果砧木（9个楸子类型，另外2种分别为新疆野苹果和平邑甜茶）10d，统计盐害指数（SI）显示：处理3d后，红海棠、白海棠、冬白果和内蒙古海红较早地表现出了盐害症状（SI≥0.10）；在处理后第10d，富平楸子、东北黄海棠、崂山柰子、新疆野苹果和平邑甜茶的盐害指数相对较低，而白海棠、五棱海棠和内蒙古海红，它们的盐害指数相对较高，分别达到0.519、0.509和0.625。
5.通过比较净光合速率（Pn）、相对电导率变化百分比（VP）、相对含水量（RWC），结合盐害指数以及生物量抗逆系数（RAC）的聚类分析结果，11种砧木的耐盐性被分为三大类，耐盐性强的砧木包括富平楸子、东北黄海棠、崂山柰子、新疆野苹果和平邑甜茶；耐盐性中等的砧木有吴起楸子和冬白果；耐盐性弱的砧木分别为红海棠、白海棠、五棱海棠及内蒙古海红。除红海棠之外，聚类分析的结果与统计盐害指数的结果一致。
6.利用降落PCR结合SSR分子标记技术，对15个楸子类型进行遗传差异分析。10对SSR引物所检测出的170条谱带中，有156条为多态性带，多态性谱带比率达到90.74%，平均遗传相似系数（GC）为0.660，GC最高的是富平楸子与吴起楸子，为0.882；其遗传距离（GD）也只有0.125；冬白果与河北热滚子的GC最低，只有0.529，但GD却达到0.636。
7.利用非加权算数平均法（UPGMA）在遗传相似值0.650处，15个楸子类型被分为三大类。这与主成分分析（PCA）的结果一致。聚类分析表明，多数聚在一起的楸子类型与它们的地理分布相一致，但也有部分个体的DNA谱带与其起源地不一致，5个起源于新疆的类型就被分散在不同类群，这在一定程度上反映了楸子不同类型复杂的遗传背景。
Malus prunifolia is an endemic wild apple germplasm resource in our country withstrong adaptability to different natural conditions. It is also an important apple rootstock forgood grafting affinity ability and widely used in northern and northwestern China. Thisdiverse species has formed many biotypes (variant) as long-term natural hybridization,genetic mutation and environmental factors influence. Some biotypes are so similar inphenotypic that it is difficult to distinguish them from each other. All these lead to itsnomenclature is confusing and the taxonomy is fraught with problems. On the other hand, nocomprehensive and systematic evaluation has been made of its drought and salt tolerance.Therefore, in our study, through explored the physiological response and evaluated thetolerance in different biotypes of M. prunifolia under drought and salt stress, it provide animportant theoretical and practical value for apple stress breeding and cultivation. Meanwhile,we used SSR markers in combination with cluster and principal coordinate analysis toevaluate the genetic relationship among biotypes of M. prunifolia; the findings are helpful tooptimize the collection and management of apple germplasm resources, and also provide atheoretical reference for traditional morphological classifications of Malus prunifolia at themolecular level.
The main results were as follows:
1. Eight apple rootstocks (seven biotypes of M. prunifolia and M. sieversii) grafted with‘Fuji’ subjected to drought stress for12d. Compared with the control (0d), the leaf relativewater content (RWC) values decreased significantly in different rootstocks, and the amount ofleaves total chlorophyll as well as photosynthesis parameters also declined to varying degrees.M. prunifolia ‘dongbeihuanghaitang’, M. sieversii, and M. prunifolia ‘fupingqiuzi’ stillmaintained relatively high Pn (8.48,7.55, and6.34μmol m-2s-1, respectively) after9d stress.Two more-sensitive biotypes of M. prunifolia—‘honghaitang’ and ‘baihaitang’—which hadthe lowest Pn values, at2.54and3.82μmol m-2s-1, respectively. Moreover, WUEiwas thehighest,5.71, in the more-resistant M. prunifolia ‘fupingqiuzi’, compared with the lowestvalue,2.42, for M. prunifolia ‘honghaitang’.
2. The activities of ascorbate peroxidase (APX), peroxidase (POD), and catalase (CAT)changed significantly during drought treatment. For APX activity, all biotypes had a sharpincrease during the first3d and began to decline after9d stress. However, the maximumactivity varied by biotypes. The POD activity began to drop in M. prunifolia ‘honghaitang’, M.prunifolia ‘wulenghaitang’, and M. prunifolia ‘baihaitang’ after3d, but M. prunifolia‘fupingqiuzi’, M. prunifolia ‘dongbeihuanghaitang’, M. prunifolia ‘laoshannaizi’ and M.sieversii still maintanted an upward trend. CAT activity raised quickly in M. prunifolia—‘honghaitang’ and ‘baihaitang’ at the stress started, but it did not maintant very long;however, M. prunifolia ‘fupingqiuzi’, M. prunifolia ‘dongbeihuanghaitang’, and M. sieversiiincreased slowly during the first3d treatment, then raised at a faster rate and reached thepeak values on6d.
3. All treated plants had elevated electrolyte leakage (EL) and malondialdehyde (MDA)contents after12d of stress. Compared with their corresponding control, values for EL inless-tolerant M. prunifolia ‘honghaitang’ and ‘baihaitang’ increased by46.37%and45.86%,MDA contents increased by30.04%and31.92%; Such damage was not as severe amonghighly drought-tolerant rootstocks, e.g., M. prunifolia ‘fupingqiuzi’, M. prunifolia‘dongbeihuanghaitang’, and M. sieversii, which EL values increased by23.33～32.94%andMDA contents increased by22.06～23.94%compared with the control. Cluster analysis cangroup these Malus biotypes into three categories. The high tolerance included two M.prunifolia—‘fupingqiuzi’ and ‘dongbeihuanghaitang’—plus M. sieversii Roem; moderatetolerance were three M. prunifolia—‘wuqiqiuzi’,‘wulenghaitang’, and ‘laoshannaizi’; lowtolerance were two M. prunifolia—‘honghaitang’ and ‘baihaitang’.
4. Eleven apple rootstocks (nine biotypes of Malus prunifolia plus M. sieversii and M.hupehensis) were exposed to150mM NaCl for10d. Salt injury index (SI) showed that, fourbiotypes of M. prunifolia—‘honghaitang’,‘baihaitang’,‘dongbaiguo’, and‘neimengguhaihong’ were more susceptible than other biotypes after3days of exposure (SI≥0.10); after10d stress, M. prunifolia ‘fupingqiuzi’, M. prunifolia ‘dongbeihuanghaitang’, andM. prunifolia ‘laoshannaizi’, as well as plants of M. sieversii and M. hupehensis, hadrelatively low SI. However, three more sensitive biotypes of M. prunifolia—‘baihaitang’,‘wulenghaitang’, and ‘neimengguhaihong’—that had higher SI values of0.519,0.509, and0.625, respectively.
5. By comparison with net photosynthetic rate (Pn), variation percentage (VP) ofelectrolyte leakage, relative water content (RWC), SI and biomass adverse resistancecoefficients (ARC), we grouped these biotypes into three classes. The high tolerance included:three for M. prunifolia—‘fupingqiuzi’,‘dongbeihuanghaitang’ and ‘laoshannaizi’—plus M. sieversii and M. hupehensis; the moderate tolerance were: M. prunifolia ‘wuqiqiuzi’ and‘dongbaiguo’; the low tolerance were four for M. prunifolia—‘honghaitang’,‘baihaitang’,‘wulenghaitang’, and ‘neimengguhaihong’. Except for M. prunifolia ‘honghaitang’, all ofthese groupings followed a pattern similar to that found with SI values.
6. We used10pairs of simple sequence repeat (SSR) primers in combination withtouchdown PCR to evaluate the genetic relationships among15biotypes of Malus prunifolia.Out of170detected fragments,156(90.74%) were polymorphic. The average geneticsimilarity coefficient (GC) between pairs of biotypes was0.66. The greatest GC (0.88) wasfound between ‘fupingqiuzi’ and ‘wuqiqiuzi’, while the biotype pairing of ‘dongbaiguo’ and‘hebeiregunzi’ showed the least GC (0.53).
7. Cluster analysis sorted all biotypes into three groups, which was then confirmed byprincipal component analysis (PCA). A dendrogram showed that some of the clusteredbiotypes were largely congruent for geographical distribution. However, the DNA patterns forsome biotype groups did not demonstrate relative agreement in their origin. Five biotypesarising from the Xinjiang Autonomous Region were dispersed within the dendrogram,reflecting their complex genetic backgrounds.
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