3种卫矛属植物叶片秋冬季转色期生理生化特征研究
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摘要
以疏花卫矛(Euonymus laxiflorus)、丝棉木(E.maackii)和卫矛(E.alatus)3种卫矛属植物为试验材料,测定叶片转色期叶色参数和相关生理生化指标,探讨各项指标的变化规律和内在联系,为优良色叶植物的筛选提供依据。结果表明:(1)转色期,疏花卫矛的明亮度参数L~*和色素参数b~*(黄/蓝)呈上升趋势,色素参数a~*(红/绿)变化不大;丝棉木的L~*和b~*呈先上升后下降的单峰曲线,a~*呈上升趋势;卫矛的L~*和b~*变化不大,a~*呈上升趋势。(2)3种植物的叶绿素含量在转色期呈明显下降趋势;疏花卫矛的花色素苷相对含量、花色素苷/叶绿素的值较为平稳,类胡萝卜素/叶绿素的值呈上升趋势;丝棉木和卫矛的花色素苷相对含量、花色素苷/叶绿素的值均呈上升趋势,类胡萝卜素/叶绿素的值保持平稳。(3)3种植物苯丙氨酸解氨酶(PAL)活性呈下降趋势,查尔酮异构酶(CHI)活性呈先上升后下降的单峰曲线;卫矛的过氧化物酶(POD)和多酚氧化酶(PPO)活性保持平稳,疏花卫矛和丝棉木的POD活性呈上升趋势,疏花卫矛的PPO活性保持平稳,丝棉木的PPO活性呈下降趋势。(4)疏花卫矛和卫矛的可溶性糖和淀粉的质量分数呈先上升后下降的单峰曲线,丝棉木的可溶性糖和淀粉的质量分数呈上升趋势。(5)相关性分析显示,疏花卫矛呈现黄色主要是因为叶绿素的分解,丝棉木和卫矛呈现红色是因为花色素苷的合成,可溶性糖、淀粉、CHI对花色素苷的合成有一定作用,POD对叶片呈现红色有促进作用,PAL和PPO活性对花色素苷的合成无显著影响。
With the perennial cutting seedling of Euonymus laxiflorus, E. maackii and E. alatus as experimental materials, we determined the leaf color parameters and related physiological and biochemical indexes of the leaves during the color change period. The changing rules and internal relations of various indicators were discussed, with a prospect to provide a scientific basis for selecting excellent color leaf plants. The results showed that:(1) the values of L~*(Gloss brightness) and b~*(Yellow/Blue) of E. laxiflorus showed an upward trend, and the value of a~*(Red/Green) did not change much. The values of L~* and b~* of E. maackii increased first and then decreased and the value of a~* is on the rise. The values of L~* and b~* of E. alatus did not changed much and the value of a~* is on the rise.(2) The chlorophyll content of the 3 species showed a significant downward trend during the color change period. The relative content of anthocyanins and the value of anthocyanin/chlorophyll of E. laxiflorus were relatively stable while its value of carotenoid/chlorophyll increased. The relative content of anthocyanin and the value of anthocyanin/chlorophyll of E. maackii and E. alatus increased, and their value of carotenoid/chlorophyll remained stable.(3) The phenylalanine ammonia lyase(PAL) activity of the three species showed a downward trend, and the activity of chalcone isomerase(CHI) increased first and then decreased. The activities of peroxidase(POD) and polyphenol oxidase(PPO) of E. alatus remained stable. The POD of E. laxiflorus and E. maackii increased. The PPO of E. laxiflorus remained stable and the PPO of E. maackii showed a downward trend.(4) The mass fraction of soluble sugar and starch of E. laxiflorus and E. alatus increased first and then decreased. The mass fraction of soluble sugar and starch of E. maackii increased.(5) According to the correlation analysis, the yellow color of E. laxiflorus was mainly due to the decomposition of chlorophyll. The red color of E. maackii and E. alatus were mainly due to the synthesis of anthocyanins. Soluble sugar, starch, CHI had a certain effect on the synthesis of anthocyanins. POD promoted the appearance of red leaves. PAL and PPO had no significant relationship with the synthesis of anthocyanins.
With the perennial cutting seedling of Euonymus laxiflorus, E. maackii and E. alatus as experimental materials, we determined the leaf color parameters and related physiological and biochemical indexes of the leaves during the color change period. The changing rules and internal relations of various indicators were discussed, with a prospect to provide a scientific basis for selecting excellent color leaf plants. The results showed that:(1) the values of L~*(Gloss brightness) and b~*(Yellow/Blue) of E. laxiflorus showed an upward trend, and the value of a~*(Red/Green) did not change much. The values of L~* and b~* of E. maackii increased first and then decreased and the value of a~* is on the rise. The values of L~* and b~* of E. alatus did not changed much and the value of a~* is on the rise.(2) The chlorophyll content of the 3 species showed a significant downward trend during the color change period. The relative content of anthocyanins and the value of anthocyanin/chlorophyll of E. laxiflorus were relatively stable while its value of carotenoid/chlorophyll increased. The relative content of anthocyanin and the value of anthocyanin/chlorophyll of E. maackii and E. alatus increased, and their value of carotenoid/chlorophyll remained stable.(3) The phenylalanine ammonia lyase(PAL) activity of the three species showed a downward trend, and the activity of chalcone isomerase(CHI) increased first and then decreased. The activities of peroxidase(POD) and polyphenol oxidase(PPO) of E. alatus remained stable. The POD of E. laxiflorus and E. maackii increased. The PPO of E. laxiflorus remained stable and the PPO of E. maackii showed a downward trend.(4) The mass fraction of soluble sugar and starch of E. laxiflorus and E. alatus increased first and then decreased. The mass fraction of soluble sugar and starch of E. maackii increased.(5) According to the correlation analysis, the yellow color of E. laxiflorus was mainly due to the decomposition of chlorophyll. The red color of E. maackii and E. alatus were mainly due to the synthesis of anthocyanins. Soluble sugar, starch, CHI had a certain effect on the synthesis of anthocyanins. POD promoted the appearance of red leaves. PAL and PPO had no significant relationship with the synthesis of anthocyanins.
引文
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[7] OREN-SHAMIR M.Does anthocyanin degradation play a significant role in determining pigment concentration in plants [J].Plant Science,2009,177(4):310-316.
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[10] ASSIS J S,MALDONADO R,MUNOZ T,et al.Effect of high carbon dioxide concentration on PAL activity and phenolic contents in ripening cherimoya fruit[J].Postharvest Biotechnology,2001,23:33-39.
[11] LISTER C E,LANCASTER J E.Developmental changes in enzymes of flavonoid biosynthesis in the skins of red and green apple cultivars[J].Journal of Science of Food Agriculture,1996,71:313-320.
[12] 宋丽红,曹帮华.光叶楮扦插生根的吲哚乙酸氧化酶、多酚氧化酶、过氧化物酶活性变化研究[J].武汉植物学研究,2005,(4):347-350.SONG L H,CAO B H.Studies on activities of indoleacetic acid oxidase,polyphenol oxidase and peroxidase in cuttings of Broussonetia papyrifera during rooting process[J].Journal of Wuhan Botanical Research,2005,(4):347-350.
[13] HARBORNE J B.Recent advances in chemical ecology[J].Natural Product Reports,1986,3:323-344.
[14] MATILE P.Biochemistry of Indian summer:physiology of autumnal leaf coloration[J].Experimental Gerontology,2000,35(2):145-158.
[15] 张亚平,曾艳,林雪莹,等.榉树叶色和色素组成的相互关系研究[J].河南农业科学,2017,46(6):116-119.ZHANG Y P,ZENG Y,LIN X Y,et al.Correlation between leaf color and pigments composition of Zelkova schneideriana[J].Journal of Henan Agricultural Sciences,2017,46(6):116-119.
[16] 冯露,吴际洋,鞠易倩,等.紫叶紫薇呈色生理及光合特性研究[J].北京林业大学学报,2017,39(12):93-101.FENG L,WU J Y,JU Y Q,et al.Leaf color characteristics and photosynthetic characteristics of purple-leafed Lagerstroemia indica ‘Ebony Ember’[J].Journal of Beijing Forestry University,2017,39(12):93-101.
[17] 卓启苗,丁彦芬,余慧,等.欧洲卫矛秋冬转色期叶色变化的生理机制[J].西北植物学报,2018,38(6):1 072-1 079.ZHUO Q M,DING Y F,YU H,et al.Physiological characters of Euonymus europaea leaves during the color-changing period in autumn and winter[J].Acta Botanica Boreali-Occidentalia Sinica,2018,38(6):1 072-1 079.
[18] 黄小贞,赵德刚.植物苯丙氨酸解氨酶表达调控机理的研究进展[J].贵州农业科学,2017,45(4):16-20.HUANG X Z,ZHAO D G.Research progress in regulation and control mechanism of phenylalanine ammonia lyase in plants[J].Guizhou Agricultural Sciences,2017,45(4):16-20.
[19] 韩培培.3种杉科植物秋季叶色变化的生理生化研究[D].杭州:浙江农林大学,2014.
[20] 王淑娟.元宝枫秋叶转色期花色素苷消长规律及生理机制研究[D].山西晋中:山西农业大学,2014.
[21] JU Z,LIU C,YUAN Y,et al.Coloration potential,anthocyanin accumulation,and enzyme activity in fruit of commercial apple cultivars and their F1 progeny[J].Scientia Horticulturae (Amsterdam),1999,79(1-2):0-50.
[22] 王惠聪,黄旭明,胡桂兵,等.荔枝果皮花青苷合成与相关酶的关系研究[J].中国农业科学,2004,(12):2 028-2 032.WANG H C,HUANG X M,HU G B,et al.Studies on the relationship between anthocyanin biosynthesis and related enzymes in Litchi pericarp[J].Scientia Agricultura Sinica,2004,(12):2 028-2 032.
[24] 冯立娟.‘红叶短枝'卫矛变色期叶片中花青苷含量及其合成相关酶活性变化的研究[C]// 中国园艺学会观赏园艺专业委员会,张启翔.中国观赏园艺研究进展2012.北京:中国林业出版社,2012:5.
[25] 史宝胜.紫叶李叶色生理变化及影响因素研究[D].哈尔滨:东北林业大学,2006.
[26] 吴驭帆,于萍,祝遵凌.春季不同叶色鹅耳枥叶片生理生化特性的研究[J].西北农林科技大学学报(自然科学版),2016,44(5):120-126,132.WU Y F,YU P,ZHU Z L.Physiological and biochemical characteristics of Carpinus turczaninowii leaves with different colors in spring[J].Journal of Northwest A&F University(Natural Science Edition),2016,44(5):120-126,132.
[27] 陈桂信,潘东明,赖钟雄,等.园艺植物多酚氧化酶的分子生物学研究进展[J].江西农业大学学报,2003,(1):107-110.CHEN G X,PAN D M,LAI Z X,et al.Advances in studies on molecular biology of polyphenol oxidase in horticultural plants[J].Acta Agriculturae Universitatis Jiangxiensis,2003,(1):107-110.
[28] 王玲平,戴丹丽,胡海娇,等.不同基因型茄子果实发育过程中色素与可溶性糖的关系[J].中国蔬菜,2010,(22):41-46.WANG L P,DAI D L,HU H J,et al.Relationship between contents of pigments and soluble sugars during development of eggplant fruits with different genotypes[J].China Vegetables,2010,(22):41-46.
[29] GOLLOP R,FARHI S,PERL A.Regulation of the leucoanthocyanidin dioxygenase gene expression in Vitis vinifera[J].Plant Science,2001,161(3):579-588.
[30] 朱书香.李属彩叶植物叶片成色机理的研究[D].河北保定:河北农业大学,2010.
[31] 郭卫珍.五个山茶新品种叶色变化及景观灯对其影响研究[D].北京:北京林业大学,2014.
[32] 王振兴,于云飞,陈丽,等.彩叶植物叶片色素组成、结构以及光合特性的研究进展[J].植物生理学报,2016,52(1):1-7.WANG Z X,YU Y F,CHEN L,et al.Advances in leaf pigment composition,structure and photosynthetic characteristics of colored-leaf plants[J].Plant Physiology Journal,2016,52(1):1-7.
[33] 吴慧,王爱波,潘一展.彩叶植物叶片色素含量影响因素研究进展[J].北方园艺,2016,(8):197-200.WU H,WANG A B,PAN Y Z.Research advance on effect factor of leaf pigment content in colored-leaf plants[J].Northern Horticulture,2016,(8):197-200.
[2] 李研南.中国卫矛属(Euonymus L.)的系统发育和演化研究[D].北京:北京林业大学,2014.
[3] 赵天鹏,丁彦芬,陈舒博.欧洲卫矛嫁接技术[J].东北林业大学学报,2016,44(7):1-3.ZHAO T P,DING Y F,CHEN S B.Grafting techniques for Euonymus europaea[J].Journal of Northeast Forestry University,2016,44(7):1-3.
[4] BAEK N I,LEE Y H,PARK J D,et al.Euonymoside a:a new cytotoxic cardenolide glycoside from the bark of Euonymus sieboldianus[J].Planta Medica,1994,60(1):26-29.
[5] 郭衍银,徐坤.矿质营养与植物病害机理研究进展 [J].甘肃农业大学学报,2003,38(4):385-393.GUO Y Y,XU K.Research progress on the mechanism of plant disease related to mineral nutrition[J].Journal of Gansu Agricultural University,2003,38(4):385-393
[6] DUBOS C,LE G J,BAUDRY A,et al.MYBL2 is a new regulator of flavonoid biosynthesis in Arabidopsis thaliana[J].The Plant Journal,2008,55:940-953.
[7] OREN-SHAMIR M.Does anthocyanin degradation play a significant role in determining pigment concentration in plants [J].Plant Science,2009,177(4):310-316.
[8] WANG L S,HASHIMOTO F,SHIRAISHI A,et al.Coloration and pigmentation of tree peony cultivars of the northwest of China[J].J.Jpn.Soc.Hort.Sci.,2000,69(2):233.
[9] 刘儒,原勤勤,袁小平,等.不同枫香家系叶片色素含量变化及其与叶色变化的关系[J].南方林业科学,2017,45(4):46-49.LIU R,YUAN Q Q,YUAN X P,et al.The relationship with change of pigment content in leaves of different Liquidambar formosana families and change of leaf color[J].South China Forestry Science,2017,45(4):46-49.
[10] ASSIS J S,MALDONADO R,MUNOZ T,et al.Effect of high carbon dioxide concentration on PAL activity and phenolic contents in ripening cherimoya fruit[J].Postharvest Biotechnology,2001,23:33-39.
[11] LISTER C E,LANCASTER J E.Developmental changes in enzymes of flavonoid biosynthesis in the skins of red and green apple cultivars[J].Journal of Science of Food Agriculture,1996,71:313-320.
[12] 宋丽红,曹帮华.光叶楮扦插生根的吲哚乙酸氧化酶、多酚氧化酶、过氧化物酶活性变化研究[J].武汉植物学研究,2005,(4):347-350.SONG L H,CAO B H.Studies on activities of indoleacetic acid oxidase,polyphenol oxidase and peroxidase in cuttings of Broussonetia papyrifera during rooting process[J].Journal of Wuhan Botanical Research,2005,(4):347-350.
[13] HARBORNE J B.Recent advances in chemical ecology[J].Natural Product Reports,1986,3:323-344.
[14] MATILE P.Biochemistry of Indian summer:physiology of autumnal leaf coloration[J].Experimental Gerontology,2000,35(2):145-158.
[15] 张亚平,曾艳,林雪莹,等.榉树叶色和色素组成的相互关系研究[J].河南农业科学,2017,46(6):116-119.ZHANG Y P,ZENG Y,LIN X Y,et al.Correlation between leaf color and pigments composition of Zelkova schneideriana[J].Journal of Henan Agricultural Sciences,2017,46(6):116-119.
[16] 冯露,吴际洋,鞠易倩,等.紫叶紫薇呈色生理及光合特性研究[J].北京林业大学学报,2017,39(12):93-101.FENG L,WU J Y,JU Y Q,et al.Leaf color characteristics and photosynthetic characteristics of purple-leafed Lagerstroemia indica ‘Ebony Ember’[J].Journal of Beijing Forestry University,2017,39(12):93-101.
[17] 卓启苗,丁彦芬,余慧,等.欧洲卫矛秋冬转色期叶色变化的生理机制[J].西北植物学报,2018,38(6):1 072-1 079.ZHUO Q M,DING Y F,YU H,et al.Physiological characters of Euonymus europaea leaves during the color-changing period in autumn and winter[J].Acta Botanica Boreali-Occidentalia Sinica,2018,38(6):1 072-1 079.
[18] 黄小贞,赵德刚.植物苯丙氨酸解氨酶表达调控机理的研究进展[J].贵州农业科学,2017,45(4):16-20.HUANG X Z,ZHAO D G.Research progress in regulation and control mechanism of phenylalanine ammonia lyase in plants[J].Guizhou Agricultural Sciences,2017,45(4):16-20.
[19] 韩培培.3种杉科植物秋季叶色变化的生理生化研究[D].杭州:浙江农林大学,2014.
[20] 王淑娟.元宝枫秋叶转色期花色素苷消长规律及生理机制研究[D].山西晋中:山西农业大学,2014.
[21] JU Z,LIU C,YUAN Y,et al.Coloration potential,anthocyanin accumulation,and enzyme activity in fruit of commercial apple cultivars and their F1 progeny[J].Scientia Horticulturae (Amsterdam),1999,79(1-2):0-50.
[22] 王惠聪,黄旭明,胡桂兵,等.荔枝果皮花青苷合成与相关酶的关系研究[J].中国农业科学,2004,(12):2 028-2 032.WANG H C,HUANG X M,HU G B,et al.Studies on the relationship between anthocyanin biosynthesis and related enzymes in Litchi pericarp[J].Scientia Agricultura Sinica,2004,(12):2 028-2 032.
[24] 冯立娟.‘红叶短枝'卫矛变色期叶片中花青苷含量及其合成相关酶活性变化的研究[C]// 中国园艺学会观赏园艺专业委员会,张启翔.中国观赏园艺研究进展2012.北京:中国林业出版社,2012:5.
[25] 史宝胜.紫叶李叶色生理变化及影响因素研究[D].哈尔滨:东北林业大学,2006.
[26] 吴驭帆,于萍,祝遵凌.春季不同叶色鹅耳枥叶片生理生化特性的研究[J].西北农林科技大学学报(自然科学版),2016,44(5):120-126,132.WU Y F,YU P,ZHU Z L.Physiological and biochemical characteristics of Carpinus turczaninowii leaves with different colors in spring[J].Journal of Northwest A&F University(Natural Science Edition),2016,44(5):120-126,132.
[27] 陈桂信,潘东明,赖钟雄,等.园艺植物多酚氧化酶的分子生物学研究进展[J].江西农业大学学报,2003,(1):107-110.CHEN G X,PAN D M,LAI Z X,et al.Advances in studies on molecular biology of polyphenol oxidase in horticultural plants[J].Acta Agriculturae Universitatis Jiangxiensis,2003,(1):107-110.
[28] 王玲平,戴丹丽,胡海娇,等.不同基因型茄子果实发育过程中色素与可溶性糖的关系[J].中国蔬菜,2010,(22):41-46.WANG L P,DAI D L,HU H J,et al.Relationship between contents of pigments and soluble sugars during development of eggplant fruits with different genotypes[J].China Vegetables,2010,(22):41-46.
[29] GOLLOP R,FARHI S,PERL A.Regulation of the leucoanthocyanidin dioxygenase gene expression in Vitis vinifera[J].Plant Science,2001,161(3):579-588.
[30] 朱书香.李属彩叶植物叶片成色机理的研究[D].河北保定:河北农业大学,2010.
[31] 郭卫珍.五个山茶新品种叶色变化及景观灯对其影响研究[D].北京:北京林业大学,2014.
[32] 王振兴,于云飞,陈丽,等.彩叶植物叶片色素组成、结构以及光合特性的研究进展[J].植物生理学报,2016,52(1):1-7.WANG Z X,YU Y F,CHEN L,et al.Advances in leaf pigment composition,structure and photosynthetic characteristics of colored-leaf plants[J].Plant Physiology Journal,2016,52(1):1-7.
[33] 吴慧,王爱波,潘一展.彩叶植物叶片色素含量影响因素研究进展[J].北方园艺,2016,(8):197-200.WU H,WANG A B,PAN Y Z.Research advance on effect factor of leaf pigment content in colored-leaf plants[J].Northern Horticulture,2016,(8):197-200.