本文主要研究内容
作者钟亚琴（2019）在《嫁接西瓜响应低钾胁迫的生理和分子机制研究》一文中研究指出：钾是植物生长所需元素之一,在植物生长发育过程中有着重要作用。我国土壤缺钾的状况仍未得到有效解决,已经成为影响作物产量和品质的限制性因子之一。西瓜[Citrullus lanatus(Thunb.)Matsum.and Nakai]是我国重要的园艺作物,在全国各地均有广泛种植。西瓜在其生长发育中需要大量的钾元素,土壤缺钾问题己经成为我国西瓜生产中重要的限制因子。本研究通过对嫁接西瓜进行短期和长期低钾胁迫处理,从嫁接西瓜生长生理、果实产量及品质、叶片和果实的转录组及果实代谢组等方面来阐明嫁接提高西瓜耐低钾性的生理和分子机理。本研究的主要结果如下:1.砧木嫁接维持叶片较高的光合特性、抗氧化活性和根系钾吸收效率,是砧木嫁接缓解低钾胁迫对西瓜嫁接苗生长抑制的重要原因。以栽培西瓜品种“早佳8424”(Z)、野生西瓜砧木“勇士”(Y)、葫芦砧木“京欣砧一号”(J)和南瓜砧木“青研砧一号”(Q)为材料构建四个嫁接组合:Z/Z(西瓜自嫁)、Z/Y(勇士嫁接西瓜)、Z/J(京欣砧一号嫁接西瓜)和Z/Q(青研砧一号嫁接西瓜)。分别用正常供钾(6 mM K~+)和低钾胁迫(0.1 mM K~+)处理16 d,营养液栽培,研究嫁接西瓜对低钾胁迫响应。结果表明,低钾胁迫导致叶面积减少,干重降低,根系表面积、根系平均直径和根体积显著下降,西瓜嫁接苗的生长受到抑制。在0.1 mM K~+时,异源砧木嫁接能缓解低钾胁迫对西瓜嫁接苗生长抑制。低钾胁迫导致嫁接西瓜光合速率下降,活性氧积累,引起细胞膜脂过氧化,影响细胞正常生理生化功能。异源砧木嫁接一方面提高了西瓜嫁接苗在低钾胁迫下的抗氧化酶(SOD、CAT和APX)活性,有效地清除活性氧,降低膜脂过氧化程度,使其在低钾胁迫下能够维持正常的生长发育。另一方面,异源砧木利用其强大的根系,提高了低钾环境下的钾吸收量,增强了钾效率。2.叶片中光合和氧化还原相关基因在转录水平差异表达是嫁接西瓜耐低钾性的主要分子机制。比较了嫁接西瓜Z/Q和Z/Z叶片转录组对低钾胁迫(0.1 mM K~+)16 d的响应,正常钾浓度为6 mM K~+,营养液栽培。在Z/Z叶片中检测到3223个差异表达基因(699个下调表达,2524个上调表达)。Z/Q中检测到2887个差异表达基因(715个下调表达,2172个上调表达)。GO(Gene Ontology)功能富集分析表明,差异基因功能涉及氧化还原、碳水化合物代谢、磷酸化、细胞识别、光合作用的光反应和光合电子传递等生物学过程。KEGG(Kyoto Encyclopedia of Genes and Genomes)代谢通路分析表明,差异表达基因显著富集在光合作用、次生代谢产物的生物合成、氨基酸的生物合成、淀粉和蔗糖代谢等代谢通路。低钾胁迫导致光系统I、光系统II、电子传递链及碳同化产物代谢相关基因的下调表达,是低钾诱导光合下降的重要机制。低钾胁迫下,异源砧木嫁接西瓜叶片转录组中抗氧化酶和抗氧化物质的相关基因,如过氧化物酶基因、谷胱甘肽转移酶基因、谷胱甘肽过氧化物酶基因等上调表达,这可能是异源砧木嫁接西瓜抗氧化活性比自嫁西瓜高的原因。3.低钾胁迫下,砧木嫁接提高了西瓜单果重和果实品质。以Z/Z、Z/Y、Z/J和Z/Q为材料,基质栽培,用正常供钾(6 mM K~+)和低钾胁迫(0.1 mM K~+)处理60 d,研究低钾胁迫对嫁接西瓜果实产量和品质的影响。结果表明,低钾胁迫降低了嫁接西瓜单果重和果实品质(授粉后30 d成熟果实)。在2种钾浓度下,Z/Y、Z/J和Z/Q的钾吸收量和单果重均显著高于Z/Z。在低钾胁迫下,异源砧木嫁接西瓜的果实品质优于自嫁西瓜,可溶性固形物、蔗糖、Vc、β-胡萝卜素的含量高于自嫁西瓜。砧木嫁接西瓜叶片ZR(玉米素核苷)含量和叶绿素含量(SPAD)高于自嫁接西瓜。异源砧木嫁接西瓜通过低钾胁迫下根系对K~+的高效吸收和地上部维持较高的ZR含量,缓解低钾胁迫对西瓜生长发育的抑制作用。4.自嫁西瓜果实转录组对低钾胁迫更敏感。比较了嫁接西瓜Z/Z、Z/Y、Z/J和Z/Q在2种钾浓度(6 mM K~+,0.1 mM K~+)下果肉转录组,基质栽培。和正常供钾(6 mM K~+)相比,低钾(0.1 mM K~+)胁迫下,Z/Z嫁接西瓜果实有670个差异表达基因(451个上调表达,219个下调表达);Z/Y嫁接西瓜果实有27个差异表达基因(7个上调表达,20个下调表达);Z/J嫁接西瓜果实有16个差异表达基因(6个上调表达,10个下调表达);Z/Q嫁接西瓜果实有15个差异表达基因(11个上调表达,4个下调表达)。Z/Z果实转录组对低钾胁迫更敏感。GO功能富集分析表明,Z/Z中差异基因主要富集在甘油磷酸二酯酶活性、多元醇代谢、单生物体糖代谢、细胞糖代谢、甘油代谢等过程。KEGG代谢通路分析表明,差异基因主要富集在牛磺酸和亚牛磺酸代谢、DNA复制、植物激素信号转导、光合生物中的碳固定、氮代谢等代谢途径。5.异源砧木嫁接能提高西瓜果实一些功能性物质的含量。以Z/Z、Z/Y、Z/J和Z/Q为材料,正常供钾(6 mM K~+)和低钾胁迫(0.1 mM K~+)处理,基质栽培,研究嫁接西瓜果肉代谢组。共鉴定出38种代谢物,其中包括15种氨基酸、8种有机酸盐、6种糖、4种核酸、5种其他化合物。最丰富的五种代谢物是葡萄糖、果糖、蔗糖、瓜氨酸和谷氨酰胺。在正常供钾下,野生西瓜嫁接提高了腺苷、蛋氨酸的含量;葫芦嫁接提高了腺苷、瓜氨酸、葡萄糖、蛋氨酸的含量;南瓜嫁接提高了天冬酰胺,瓜氨酸、富马酸、谷氨酰胺、异亮氨酸、蛋氨酸、氧化型辅酶Ⅰ、苯丙氨酸、酪氨酸、缬氨酸的含量。低钾胁迫下,野生西瓜嫁接提高了天冬酰胺、氧化型辅酶Ⅰ、肌醇和丙氨酸的含量;葫芦嫁接提高了天冬酰胺、半乳糖和丙氨酸的含量;南瓜嫁接提高了肌醇、天冬酰胺的含量。综上所述,砧木嫁接是提高耐低钾性的重要措施。砧木嫁接提高了西瓜在低钾胁迫下根系钾吸收效率和的抗氧化能力;维持叶片较高的光合能力玉米素核苷的含量,是嫁接西瓜耐低钾的重要原因。砧木嫁接还是提高西瓜果实营养品质的主要手段。本研究发掘的低钾响应基因为未来开展西瓜的钾营养分子机制研究工作奠定了基础。
Abstract
jia shi zhi wu sheng chang suo xu yuan su zhi yi ,zai zhi wu sheng chang fa yo guo cheng zhong you zhao chong yao zuo yong 。wo guo tu rang que jia de zhuang kuang reng wei de dao you xiao jie jue ,yi jing cheng wei ying xiang zuo wu chan liang he pin zhi de xian zhi xing yin zi zhi yi 。xi gua [Citrullus lanatus(Thunb.)Matsum.and Nakai]shi wo guo chong yao de yuan yi zuo wu ,zai quan guo ge de jun you an fan chong zhi 。xi gua zai ji sheng chang fa yo zhong xu yao da liang de jia yuan su ,tu rang que jia wen ti ji jing cheng wei wo guo xi gua sheng chan zhong chong yao de xian zhi yin zi 。ben yan jiu tong guo dui jia jie xi gua jin hang duan ji he chang ji di jia xie pai chu li ,cong jia jie xi gua sheng chang sheng li 、guo shi chan liang ji pin zhi 、xie pian he guo shi de zhuai lu zu ji guo shi dai xie zu deng fang mian lai chan ming jia jie di gao xi gua nai di jia xing de sheng li he fen zi ji li 。ben yan jiu de zhu yao jie guo ru xia :1.zhen mu jia jie wei chi xie pian jiao gao de guang ge te xing 、kang yang hua huo xing he gen ji jia xi shou xiao lv ,shi zhen mu jia jie huan jie di jia xie pai dui xi gua jia jie miao sheng chang yi zhi de chong yao yuan yin 。yi zai pei xi gua pin chong “zao jia 8424”(Z)、ye sheng xi gua zhen mu “yong shi ”(Y)、hu lu zhen mu “jing xin zhen yi hao ”(J)he na gua zhen mu “qing yan zhen yi hao ”(Q)wei cai liao gou jian si ge jia jie zu ge :Z/Z(xi gua zi jia )、Z/Y(yong shi jia jie xi gua )、Z/J(jing xin zhen yi hao jia jie xi gua )he Z/Q(qing yan zhen yi hao jia jie xi gua )。fen bie yong zheng chang gong jia (6 mM K~+)he di jia xie pai (0.1 mM K~+)chu li 16 d,ying yang ye zai pei ,yan jiu jia jie xi gua dui di jia xie pai xiang ying 。jie guo biao ming ,di jia xie pai dao zhi xie mian ji jian shao ,gan chong jiang di ,gen ji biao mian ji 、gen ji ping jun zhi jing he gen ti ji xian zhe xia jiang ,xi gua jia jie miao de sheng chang shou dao yi zhi 。zai 0.1 mM K~+shi ,yi yuan zhen mu jia jie neng huan jie di jia xie pai dui xi gua jia jie miao sheng chang yi zhi 。di jia xie pai dao zhi jia jie xi gua guang ge su lv xia jiang ,huo xing yang ji lei ,yin qi xi bao mo zhi guo yang hua ,ying xiang xi bao zheng chang sheng li sheng hua gong neng 。yi yuan zhen mu jia jie yi fang mian di gao le xi gua jia jie miao zai di jia xie pai xia de kang yang hua mei (SOD、CAThe APX)huo xing ,you xiao de qing chu huo xing yang ,jiang di mo zhi guo yang hua cheng du ,shi ji zai di jia xie pai xia neng gou wei chi zheng chang de sheng chang fa yo 。ling yi fang mian ,yi yuan zhen mu li yong ji jiang da de gen ji ,di gao le di jia huan jing xia de jia xi shou liang ,zeng jiang le jia xiao lv 。2.xie pian zhong guang ge he yang hua hai yuan xiang guan ji yin zai zhuai lu shui ping cha yi biao da shi jia jie xi gua nai di jia xing de zhu yao fen zi ji zhi 。bi jiao le jia jie xi gua Z/Qhe Z/Zxie pian zhuai lu zu dui di jia xie pai (0.1 mM K~+)16 dde xiang ying ,zheng chang jia nong du wei 6 mM K~+,ying yang ye zai pei 。zai Z/Zxie pian zhong jian ce dao 3223ge cha yi biao da ji yin (699ge xia diao biao da ,2524ge shang diao biao da )。Z/Qzhong jian ce dao 2887ge cha yi biao da ji yin (715ge xia diao biao da ,2172ge shang diao biao da )。GO(Gene Ontology)gong neng fu ji fen xi biao ming ,cha yi ji yin gong neng she ji yang hua hai yuan 、tan shui hua ge wu dai xie 、lin suan hua 、xi bao shi bie 、guang ge zuo yong de guang fan ying he guang ge dian zi chuan di deng sheng wu xue guo cheng 。KEGG(Kyoto Encyclopedia of Genes and Genomes)dai xie tong lu fen xi biao ming ,cha yi biao da ji yin xian zhe fu ji zai guang ge zuo yong 、ci sheng dai xie chan wu de sheng wu ge cheng 、an ji suan de sheng wu ge cheng 、dian fen he zhe tang dai xie deng dai xie tong lu 。di jia xie pai dao zhi guang ji tong I、guang ji tong II、dian zi chuan di lian ji tan tong hua chan wu dai xie xiang guan ji yin de xia diao biao da ,shi di jia you dao guang ge xia jiang de chong yao ji zhi 。di jia xie pai xia ,yi yuan zhen mu jia jie xi gua xie pian zhuai lu zu zhong kang yang hua mei he kang yang hua wu zhi de xiang guan ji yin ,ru guo yang hua wu mei ji yin 、gu guang gan tai zhuai yi mei ji yin 、gu guang gan tai guo yang hua wu mei ji yin deng shang diao biao da ,zhe ke neng shi yi yuan zhen mu jia jie xi gua kang yang hua huo xing bi zi jia xi gua gao de yuan yin 。3.di jia xie pai xia ,zhen mu jia jie di gao le xi gua chan guo chong he guo shi pin zhi 。yi Z/Z、Z/Y、Z/Jhe Z/Qwei cai liao ,ji zhi zai pei ,yong zheng chang gong jia (6 mM K~+)he di jia xie pai (0.1 mM K~+)chu li 60 d,yan jiu di jia xie pai dui jia jie xi gua guo shi chan liang he pin zhi de ying xiang 。jie guo biao ming ,di jia xie pai jiang di le jia jie xi gua chan guo chong he guo shi pin zhi (shou fen hou 30 dcheng shou guo shi )。zai 2chong jia nong du xia ,Z/Y、Z/Jhe Z/Qde jia xi shou liang he chan guo chong jun xian zhe gao yu Z/Z。zai di jia xie pai xia ,yi yuan zhen mu jia jie xi gua de guo shi pin zhi you yu zi jia xi gua ,ke rong xing gu xing wu 、zhe tang 、Vc、β-hu luo bo su de han liang gao yu zi jia xi gua 。zhen mu jia jie xi gua xie pian ZR(yu mi su he gan )han liang he xie lu su han liang (SPAD)gao yu zi jia jie xi gua 。yi yuan zhen mu jia jie xi gua tong guo di jia xie pai xia gen ji dui K~+de gao xiao xi shou he de shang bu wei chi jiao gao de ZRhan liang ,huan jie di jia xie pai dui xi gua sheng chang fa yo de yi zhi zuo yong 。4.zi jia xi gua guo shi zhuai lu zu dui di jia xie pai geng min gan 。bi jiao le jia jie xi gua Z/Z、Z/Y、Z/Jhe Z/Qzai 2chong jia nong du (6 mM K~+,0.1 mM K~+)xia guo rou zhuai lu zu ,ji zhi zai pei 。he zheng chang gong jia (6 mM K~+)xiang bi ,di jia (0.1 mM K~+)xie pai xia ,Z/Zjia jie xi gua guo shi you 670ge cha yi biao da ji yin (451ge shang diao biao da ,219ge xia diao biao da );Z/Yjia jie xi gua guo shi you 27ge cha yi biao da ji yin (7ge shang diao biao da ,20ge xia diao biao da );Z/Jjia jie xi gua guo shi you 16ge cha yi biao da ji yin (6ge shang diao biao da ,10ge xia diao biao da );Z/Qjia jie xi gua guo shi you 15ge cha yi biao da ji yin (11ge shang diao biao da ,4ge xia diao biao da )。Z/Zguo shi zhuai lu zu dui di jia xie pai geng min gan 。GOgong neng fu ji fen xi biao ming ,Z/Zzhong cha yi ji yin zhu yao fu ji zai gan you lin suan er zhi mei huo xing 、duo yuan chun dai xie 、chan sheng wu ti tang dai xie 、xi bao tang dai xie 、gan you dai xie deng guo cheng 。KEGGdai xie tong lu fen xi biao ming ,cha yi ji yin zhu yao fu ji zai niu huang suan he ya niu huang suan dai xie 、DNAfu zhi 、zhi wu ji su xin hao zhuai dao 、guang ge sheng wu zhong de tan gu ding 、dan dai xie deng dai xie tu jing 。5.yi yuan zhen mu jia jie neng di gao xi gua guo shi yi xie gong neng xing wu zhi de han liang 。yi Z/Z、Z/Y、Z/Jhe Z/Qwei cai liao ,zheng chang gong jia (6 mM K~+)he di jia xie pai (0.1 mM K~+)chu li ,ji zhi zai pei ,yan jiu jia jie xi gua guo rou dai xie zu 。gong jian ding chu 38chong dai xie wu ,ji zhong bao gua 15chong an ji suan 、8chong you ji suan yan 、6chong tang 、4chong he suan 、5chong ji ta hua ge wu 。zui feng fu de wu chong dai xie wu shi pu tao tang 、guo tang 、zhe tang 、gua an suan he gu an xian an 。zai zheng chang gong jia xia ,ye sheng xi gua jia jie di gao le xian gan 、dan an suan de han liang ;hu lu jia jie di gao le xian gan 、gua an suan 、pu tao tang 、dan an suan de han liang ;na gua jia jie di gao le tian dong xian an ,gua an suan 、fu ma suan 、gu an xian an 、yi liang an suan 、dan an suan 、yang hua xing fu mei Ⅰ、ben bing an suan 、lao an suan 、xie an suan de han liang 。di jia xie pai xia ,ye sheng xi gua jia jie di gao le tian dong xian an 、yang hua xing fu mei Ⅰ、ji chun he bing an suan de han liang ;hu lu jia jie di gao le tian dong xian an 、ban ru tang he bing an suan de han liang ;na gua jia jie di gao le ji chun 、tian dong xian an de han liang 。zeng shang suo shu ,zhen mu jia jie shi di gao nai di jia xing de chong yao cuo shi 。zhen mu jia jie di gao le xi gua zai di jia xie pai xia gen ji jia xi shou xiao lv he de kang yang hua neng li ;wei chi xie pian jiao gao de guang ge neng li yu mi su he gan de han liang ,shi jia jie xi gua nai di jia de chong yao yuan yin 。zhen mu jia jie hai shi di gao xi gua guo shi ying yang pin zhi de zhu yao shou duan 。ben yan jiu fa jue de di jia xiang ying ji yin wei wei lai kai zhan xi gua de jia ying yang fen zi ji zhi yan jiu gong zuo dian ding le ji chu 。
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