外源硅对不同砧木嫁接黄瓜果面蜡粉和硅吸收及相关基因表达的影响
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摘要
为探明砧木影响嫁接黄瓜果面蜡粉形成的机制以及与外源硅的关系,以‘山农5号’黄瓜为接穗,‘云南黑籽南瓜’(去蜡粉能力弱)和‘黄诚根2号’南瓜(去蜡粉能力强)为砧木进行嫁接,采用水培法,研究营养液中添加硅与否对黄瓜硅吸收、硅转运蛋白相关基因表达和果实表面蜡粉形成的影响。结果表明:加硅显著提高了‘云南黑籽南瓜’嫁接黄瓜和自根黄瓜果面蜡粉量及各器官硅含量,但加硅后‘黄诚根2号’嫁接黄瓜的果面蜡粉量和各器官硅含量均较低,与不加硅条件下的嫁接和自根黄瓜相近;加硅处理后‘云南黑籽南瓜’嫁接黄瓜苗根系中硅转运蛋白基因Cm Lsi1、Cm Lsi2-1、Cm Lsi2-2和Cm Lsi3的表达量均高于‘黄诚根2号’嫁接黄瓜苗;随着硅处理时间延长,‘云南黑籽南瓜’嫁接黄瓜苗根系中硅转运蛋白基因表达下调,而‘黄诚根2号’嫁接黄瓜苗根系中硅转运蛋白基因的表达同样下调。
In order to investigate the mechanism underlying the effects of rootstocks on fruit bloom formation of grafted cucumbers under different silicon nutrition conditions,the rootstocks‘Yunnan figleaf gourd'and‘Huangchenggen 2'were used to study the absorption and distribution of silicon,expression of silicon transporter genes and bloom formation in the grafted plants of cucumber‘Shannong 5',which were grown by hydroponics with the nutrient solution with silicon added or without. The results showed that the fruit bloom and silicon contents in organ of the cucumbers grafted on‘Yunnan figleaf gourd'and self-rooted cucumbers were significantly increased by added silicon,and much higher than those in the cucumbers grafted on‘Huangchenggen 2'. The fruit bloom and silicon contents in organs of the cucumbers grafted on‘Huangchenggen 2'by added silicon were similar with those of the cucumbers grafted or self-rooted cucumbers in the condition without silicon added. The expression of Cm Lsi1,Cm Lsi2-1,Cm Lsi2-2 and Cm Lsi3 in roots of cucumbers grafted on‘Yunnan figleaf gourd'was higher than that in roots of cucumbers grafted on‘Huangchenggen 2'. With the prolongation of silicon treatment,the expression of silicon transporter genes in roots of cucumbers grafted onto‘Yunnan figleaf gourd'was down-regulated,the expression of those genes in roots of cucumbers grafted on‘Huangchenggen 2'also decreased.
In order to investigate the mechanism underlying the effects of rootstocks on fruit bloom formation of grafted cucumbers under different silicon nutrition conditions,the rootstocks‘Yunnan figleaf gourd'and‘Huangchenggen 2'were used to study the absorption and distribution of silicon,expression of silicon transporter genes and bloom formation in the grafted plants of cucumber‘Shannong 5',which were grown by hydroponics with the nutrient solution with silicon added or without. The results showed that the fruit bloom and silicon contents in organ of the cucumbers grafted on‘Yunnan figleaf gourd'and self-rooted cucumbers were significantly increased by added silicon,and much higher than those in the cucumbers grafted on‘Huangchenggen 2'. The fruit bloom and silicon contents in organs of the cucumbers grafted on‘Huangchenggen 2'by added silicon were similar with those of the cucumbers grafted or self-rooted cucumbers in the condition without silicon added. The expression of Cm Lsi1,Cm Lsi2-1,Cm Lsi2-2 and Cm Lsi3 in roots of cucumbers grafted on‘Yunnan figleaf gourd'was higher than that in roots of cucumbers grafted on‘Huangchenggen 2'. With the prolongation of silicon treatment,the expression of silicon transporter genes in roots of cucumbers grafted onto‘Yunnan figleaf gourd'was down-regulated,the expression of those genes in roots of cucumbers grafted on‘Huangchenggen 2'also decreased.
引文
Chiba Y,Mitani N,Yamaji N,Ma J F.2009.Hv Lsi1 is a silicon influx transporter in barley.The Plant Journal,57:810-818.
Gao Yan-kui,Chen Pu-hong,Li Xin.2008.Effects of different genotype rootstocks on cucumber yield and fruit quality.Changjiang Vegetables,(11b):48-50.(in Chinese)高彦魁,陈普红,李欣.2008.不同基因型砧木对黄瓜产量和果实品质的影响.长江蔬菜,(11b):48-50.
Gao Yan-kui,Li Xin,Zhao Zhi-jun.2011.Impact of different rootstock genotypes on output,bloom,disease resistance and winter hardness of cucumber.Acta Agriculturae Boreali-occidentalis Sinica,20(3):180-183.(in Chinese)高彦魁,李欣,赵志军.2011.不同基因型砧木对黄瓜产量、果霜及抗病性和抗寒性的影响.西北农业学报,20(3):180-183.
Hisao Yamashita,Itakuchi Takumi,Shoji Kita.1989.The effects of rootstocks on cucumber yields and bloom by forcing cultivation.Journal of Tokushima Agriculture&Experiment,26:9-16.(in Japanese)山下久男,板口巧,北冈祥治.1989.促成キュウリ栽培における少ブル-ム台木の种类がブル-ムの发生と收量に及ぼす影响.德岛农试研报,26:9-16.
Honjima Jung.1989.A simple method to determine the occurrence degree of cucumber fruit bloom.Academy of Art and Garden,64(4):541-542.(in Japanese)本岛俊明.1989.キュウリ果面のブル-ム发生程度の简易な测定法.农业および园芸,64(4):541-542.
Liang Y C,Si J,R?mheld V.2005.Silicon uptake and transport is an active process in Cucumis sativus.New Phytologist,167(3):797-804.
Liu Qing,Wei Min,Shen Qiong,Wang Xiu-feng,Yang Feng-juan,Shi Qing-hua.2012.Effects of different rootstocks on formation and silicon absorption and distribution of grafted cucumber.Acta Horticulturae Sinica,39(5):897-904.(in Chinese)刘青,魏珉,沈琼,王秀峰,杨凤娟,史庆华.2012.不同砧木对嫁接黄瓜蜡粉形成及硅吸收分配的影响.园艺学报,39(5):897-904.
Ma J F,Tamai K,Yamaji N,Mitani N,Konishi S,Katsuhara M,Ishiguro M,Murata Y,Yano M.2006.A silicon transporter in rice.Nature,440(30):688-691.
Ma J F,Yamaji N.2015.A cooperative system of silicon transport in plants.Trends in Plant Science,20:435-442.
Ma J F,Yamaji N,Mitani N,Tamai K,Konishi S,Fujiwara T,Katsuhara M,Yano M.2007.An efflux transporter of silicon in rice.Nature,448(12):209-212.
Mitani N,Chiba Y,Yamaji N.2009a.Identification and characterization of maize and barley Lsi2-like silicon efflux transporters reveals a distinct silicon uptake system from that in rice.Plant Cell,21(7):2133-2142.
Mitani N,Ma J F.2005.Uptake system of silicon in different plant species.Journal of Experimental Botany,56(414):1255-1261.
Mitani N,Yamaji N,Ago Y,Iwasaki K,Ma J F.2011a.Isolation and functional characterization of an influx silicon transporter in two pumpkin cultivars contrasting in silicon accumulation.The Plant Journal,66:231-240.
Mitani N,Yamaji N,Ma J F.2009b.Identification of maize silicon influx transporters.Plant and Cell Physiology,50(1):5-12.
Mitani N,Yamaji N,Ma J F.2011b.Silicon efflux transporters isolated from two pumpkin cultivars contrasting in Si uptake.Plant Signaling&Behavior,6(7):991-994.
Sakata Y,Ohara T,Sugiyama M.2008.The history of melon and cucumber grafting in Japan.Acta Horticulturae,767:217-228.
Seki M,Hotta Y.1997.Effect of bloomless stock cultivar on the growth and mineral uptake of cucumber(Cucumis sativus)plants.Research Bulletin of the Aichi-ken Agricultural Research Center,29:127-133.
Shen Qiong.2013.Studies on bloom formation on fruit surface of cucumber and the relationship with absorption and distribution of silicon[M.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)沈琼.2013.黄瓜果实表面蜡粉形成及其硅吸收分配的关系研究[硕士论文].泰安:山东农业大学.
Sun Hong,Su Tong-bing,Yu Shuan-cang,Zhang Feng-lan,Yu Yang-jun.2017.Analysis of the compositions and synthetic genes related to wax tablets in leaves of Chinese cabbage.China Vegetables,(7):42-48.(in Chinese)孙红,苏同兵,于拴仓,张凤兰,余阳俊.2017.白菜叶片表皮蜡粉成分及合成相关基因表达分析.中国蔬菜,(7):42-48.
Tang Jun,Liu Dong-ming,Liu Ze-zhou,Yang Li-mei,Fang Zhi-yuan,Liu Yu-mei,Zhuang Mu,Zhang Yang-yong,Sun Pei-tian.2015.Studies on characteristics of several glossy mutants in cabbage.Acta Horticulturae Sinica,42(6):1093-1102.(in Chinese)唐俊,刘东明,刘泽洲,杨丽梅,方智远,刘玉梅,庄木,张扬勇,孙培田.2015.几份甘蓝蜡粉缺失突变体特征特性的研究.园艺学报,42(6):1093-1102.
Wang Dong-xu.2014.Cloning,expression and transformation of silicon transport gene CSi T-1 and CSi T-2 in cucumber[M.D.Dissertation].Hangzhou:Zhejiang Agriculture and Forestry University.(in Chinese)王东旭.2014.硅转运相关基因Csi T-1和Csi T-2克隆表达与转化黄瓜研究[硕士论文].杭州:浙江农林大学.
Wang Hui,Zhao Sheng,Wei Min,Wang Xiu-feng,Shi Qing-hua,Yang Feng-juan.2015.Cloning and expression analysis of silicon transporter gene Cm Lsi3 in roots of pumpkin.Acta Horticulturae Sinica,42(10):2075-2082.(in Chinese)王慧,赵升,魏珉,王秀峰,史庆华,杨凤娟.2015.砧木南瓜硅转运蛋白基因Cm Lsi3的克隆与表达分析.园艺学报,42(10):2075-2082.
Wang Tie-chen,Wang Hai-rong,Li Hong-ling,Zhao Shi-chun,Wang Shi-kai.2010.Effect of grafting on different rootstocks on wax characteristics of cucumber.China Vegetables,(8):77-79.(in Chinese)王铁臣,王海荣,李红岭,赵士春,王世凯.2010.不同砧木嫁接对黄瓜蜡粉性状的影响.中国蔬菜,(8):77-79.
Wang Tie-chen,Wang Hai-rong,Zhao Shi-chun,Li Hong-ling,Chi Mei-na.2011.Preliminary report about effects of cucumber grafted with different double rootstock on growth,yield and wax powers characters.China Vegetables,(6):54-57.(in Chinese)王铁臣,王海荣,赵士春,李红岭,池美娜.2011.不同双砧木嫁接对黄瓜长势、产量及蜡粉性状的影响初报.中国蔬菜,(6):54-57.
Yamaji N,Mitatni N,Ma J F.2008.A transporter regulating silicon distribution in rice shoots.Plant Cell,20(5):1381-1389.
Zhang Qian,Chen Jia-na,Li Jian-min.2013.Advances of researches on the silicon uptake and transporters in plants.Chinese Agricultural Science Bulletin,29(12):150-156.(in Chinese)张倩,陈佳娜,李建民.2013.植物硅转运蛋白的研究进展.中国农学通报,29(12):150-156.
Gao Yan-kui,Chen Pu-hong,Li Xin.2008.Effects of different genotype rootstocks on cucumber yield and fruit quality.Changjiang Vegetables,(11b):48-50.(in Chinese)高彦魁,陈普红,李欣.2008.不同基因型砧木对黄瓜产量和果实品质的影响.长江蔬菜,(11b):48-50.
Gao Yan-kui,Li Xin,Zhao Zhi-jun.2011.Impact of different rootstock genotypes on output,bloom,disease resistance and winter hardness of cucumber.Acta Agriculturae Boreali-occidentalis Sinica,20(3):180-183.(in Chinese)高彦魁,李欣,赵志军.2011.不同基因型砧木对黄瓜产量、果霜及抗病性和抗寒性的影响.西北农业学报,20(3):180-183.
Hisao Yamashita,Itakuchi Takumi,Shoji Kita.1989.The effects of rootstocks on cucumber yields and bloom by forcing cultivation.Journal of Tokushima Agriculture&Experiment,26:9-16.(in Japanese)山下久男,板口巧,北冈祥治.1989.促成キュウリ栽培における少ブル-ム台木の种类がブル-ムの发生と收量に及ぼす影响.德岛农试研报,26:9-16.
Honjima Jung.1989.A simple method to determine the occurrence degree of cucumber fruit bloom.Academy of Art and Garden,64(4):541-542.(in Japanese)本岛俊明.1989.キュウリ果面のブル-ム发生程度の简易な测定法.农业および园芸,64(4):541-542.
Liang Y C,Si J,R?mheld V.2005.Silicon uptake and transport is an active process in Cucumis sativus.New Phytologist,167(3):797-804.
Liu Qing,Wei Min,Shen Qiong,Wang Xiu-feng,Yang Feng-juan,Shi Qing-hua.2012.Effects of different rootstocks on formation and silicon absorption and distribution of grafted cucumber.Acta Horticulturae Sinica,39(5):897-904.(in Chinese)刘青,魏珉,沈琼,王秀峰,杨凤娟,史庆华.2012.不同砧木对嫁接黄瓜蜡粉形成及硅吸收分配的影响.园艺学报,39(5):897-904.
Ma J F,Tamai K,Yamaji N,Mitani N,Konishi S,Katsuhara M,Ishiguro M,Murata Y,Yano M.2006.A silicon transporter in rice.Nature,440(30):688-691.
Ma J F,Yamaji N.2015.A cooperative system of silicon transport in plants.Trends in Plant Science,20:435-442.
Ma J F,Yamaji N,Mitani N,Tamai K,Konishi S,Fujiwara T,Katsuhara M,Yano M.2007.An efflux transporter of silicon in rice.Nature,448(12):209-212.
Mitani N,Chiba Y,Yamaji N.2009a.Identification and characterization of maize and barley Lsi2-like silicon efflux transporters reveals a distinct silicon uptake system from that in rice.Plant Cell,21(7):2133-2142.
Mitani N,Ma J F.2005.Uptake system of silicon in different plant species.Journal of Experimental Botany,56(414):1255-1261.
Mitani N,Yamaji N,Ago Y,Iwasaki K,Ma J F.2011a.Isolation and functional characterization of an influx silicon transporter in two pumpkin cultivars contrasting in silicon accumulation.The Plant Journal,66:231-240.
Mitani N,Yamaji N,Ma J F.2009b.Identification of maize silicon influx transporters.Plant and Cell Physiology,50(1):5-12.
Mitani N,Yamaji N,Ma J F.2011b.Silicon efflux transporters isolated from two pumpkin cultivars contrasting in Si uptake.Plant Signaling&Behavior,6(7):991-994.
Sakata Y,Ohara T,Sugiyama M.2008.The history of melon and cucumber grafting in Japan.Acta Horticulturae,767:217-228.
Seki M,Hotta Y.1997.Effect of bloomless stock cultivar on the growth and mineral uptake of cucumber(Cucumis sativus)plants.Research Bulletin of the Aichi-ken Agricultural Research Center,29:127-133.
Shen Qiong.2013.Studies on bloom formation on fruit surface of cucumber and the relationship with absorption and distribution of silicon[M.D.Dissertation].Tai’an:Shandong Agricultural University.(in Chinese)沈琼.2013.黄瓜果实表面蜡粉形成及其硅吸收分配的关系研究[硕士论文].泰安:山东农业大学.
Sun Hong,Su Tong-bing,Yu Shuan-cang,Zhang Feng-lan,Yu Yang-jun.2017.Analysis of the compositions and synthetic genes related to wax tablets in leaves of Chinese cabbage.China Vegetables,(7):42-48.(in Chinese)孙红,苏同兵,于拴仓,张凤兰,余阳俊.2017.白菜叶片表皮蜡粉成分及合成相关基因表达分析.中国蔬菜,(7):42-48.
Tang Jun,Liu Dong-ming,Liu Ze-zhou,Yang Li-mei,Fang Zhi-yuan,Liu Yu-mei,Zhuang Mu,Zhang Yang-yong,Sun Pei-tian.2015.Studies on characteristics of several glossy mutants in cabbage.Acta Horticulturae Sinica,42(6):1093-1102.(in Chinese)唐俊,刘东明,刘泽洲,杨丽梅,方智远,刘玉梅,庄木,张扬勇,孙培田.2015.几份甘蓝蜡粉缺失突变体特征特性的研究.园艺学报,42(6):1093-1102.
Wang Dong-xu.2014.Cloning,expression and transformation of silicon transport gene CSi T-1 and CSi T-2 in cucumber[M.D.Dissertation].Hangzhou:Zhejiang Agriculture and Forestry University.(in Chinese)王东旭.2014.硅转运相关基因Csi T-1和Csi T-2克隆表达与转化黄瓜研究[硕士论文].杭州:浙江农林大学.
Wang Hui,Zhao Sheng,Wei Min,Wang Xiu-feng,Shi Qing-hua,Yang Feng-juan.2015.Cloning and expression analysis of silicon transporter gene Cm Lsi3 in roots of pumpkin.Acta Horticulturae Sinica,42(10):2075-2082.(in Chinese)王慧,赵升,魏珉,王秀峰,史庆华,杨凤娟.2015.砧木南瓜硅转运蛋白基因Cm Lsi3的克隆与表达分析.园艺学报,42(10):2075-2082.
Wang Tie-chen,Wang Hai-rong,Li Hong-ling,Zhao Shi-chun,Wang Shi-kai.2010.Effect of grafting on different rootstocks on wax characteristics of cucumber.China Vegetables,(8):77-79.(in Chinese)王铁臣,王海荣,李红岭,赵士春,王世凯.2010.不同砧木嫁接对黄瓜蜡粉性状的影响.中国蔬菜,(8):77-79.
Wang Tie-chen,Wang Hai-rong,Zhao Shi-chun,Li Hong-ling,Chi Mei-na.2011.Preliminary report about effects of cucumber grafted with different double rootstock on growth,yield and wax powers characters.China Vegetables,(6):54-57.(in Chinese)王铁臣,王海荣,赵士春,李红岭,池美娜.2011.不同双砧木嫁接对黄瓜长势、产量及蜡粉性状的影响初报.中国蔬菜,(6):54-57.
Yamaji N,Mitatni N,Ma J F.2008.A transporter regulating silicon distribution in rice shoots.Plant Cell,20(5):1381-1389.
Zhang Qian,Chen Jia-na,Li Jian-min.2013.Advances of researches on the silicon uptake and transporters in plants.Chinese Agricultural Science Bulletin,29(12):150-156.(in Chinese)张倩,陈佳娜,李建民.2013.植物硅转运蛋白的研究进展.中国农学通报,29(12):150-156.