杜梨组培生根过程中多胺、内源激素及相关氧化酶活性的变化
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摘要
【目的】探究杜梨组培苗生根诱导过程中内源激素、多胺类物质及相关氧化酶活性变化的生理响应机制。【方法】以长势相近的杜梨继代苗为试材,筛选优化了生根培养基配方,分别在生根诱导0、3、7、10、15 d后,测定分析了基部茎段多胺类物质、内源激素含量及关联酶活性。【结果】(1)激素配比2.0 mg·L~(-1)IAA+0.5 mg·L~(-1)IBA的生根效果良好,IAA和IBA两种激素共同诱导杜梨生根效果要优于单一激素处理;(2)在生根诱导0～15 d内,精胺(Spm)和亚精胺(Spd)含量呈现下降→升高→下降→升高的变化趋势,腐胺(Put)含量则呈现出下降→上升的变化趋势;内源激素IAA、ABA含量均呈现先下降后升高的趋势,最低值出现在第7天,GA含量也呈现先下降后上升的变化趋势,最低值出现在第3天,ZR含量呈现出下降→升高→下降的变化趋势;两种关联酶IAAO和PPO酶活性均呈现增高→降低→增高的趋势(;3)Spd与IAA、ABA、IAA/ABA IAA/ZR及IAAO酶活性均存在极显著正相关关系(p <0.01),IAA与Spd、GA、ABA及IAA/ABA、IAA/ZR、PPO酶活性在α=0.01水平达到极显著正相关。【结论】杜梨较适宜的生根培养基为:1/2MS+2.0 mg·L~(-1)IAA+0.5 mg·L~(-1)IBA+7.5 g·L~(-1)琼脂+25.0 g·L~(-1)蔗糖;IAA/ABA、IAA/ZR比值增大更有利于杜梨不定根的发生;多胺类物质中Spd含量与杜梨不定根发生紧密相关;生根诱导后期,GA含量增加对杜梨不定根发生并无明显的抑制作用。
【Objective】Pyrus betulifolia Bunge is widely used as a pear rootstock in China. Propagation with seeds has large genetic variation and poor consistency. Difficulty in rooting limitsits propagation by traditional vegetative propagation methods such as layering and cutting. With the increasingly perfecting of in vitro propagation technology, virus-free seedlings can be produced by micropropagation method. Root induction is one of the important processes in in vitro propagation. The transplanting survival rate and seedling quality are affected by rooting efficiency. There are many internal and external factors affecting the rooting process, including endogenous hormones, polyamines, related oxidases and exogenous substances such as ethylene and salicylic acid. The study aimed at revealing the roles of endogenous hormones, polyamines and related oxidases in root induction of tissue-cultured seedlings.【Methods】Subcultured plantlets of Pyrus betulifolia Bunge were cultured on rooting medium with1/2 MS, 7.5 g · L~(-1) agar, 25.0 g · L~(-1) sucrose and different ratios of hormones. A total of 24 treatments using five different concentrations(0, 0.2, 0.5, 1 and 2 mg · L~(-1)) of IAA and IBA were assessed in terms of rooting percentages at 40 days after transfer to the media. Each treatment consisted of 6 bottles each comprising 5 subcultured plantlets. Uniform subcultured plantlets were selected to be cultured on the optimized rooting medium. Approximately 500 plantlets were used to monitor the dynamic changes in polyamines, endogenous hormones and related enzyme activities during root regeneration. All of the cultures were incubated at 25 ℃ under a 16 h light/8 h dark cycle and 2 000-3 000 lx light intensity. The basal stem cuttings were taken 0, 3, 7, 10 and 15 days after transfer to media. Polyamines, endogenous hormones and related enzyme activities were determined.【Results】The treatment of 2.0 mg · L~(-1) IAA +0.5 mg · L~(-1) IBA increased rooting rate and the percentage of plantlets with ≥3 roots compared with the other treatments. The rooting rate was 86.7% and the percentage of plantlets with ≥3 roots was53.3%. The combination of IAA and IBA was better than single hormone. Callus was observed between the 7 th and 10 th day, growing fast from the 10 th to 15 th day. The roots generated 20 days after transfer to media. The contents of three polyamines declined after 3 days in the root induction medium. The content of spermine(Spm) and spermidine(Spd) fluctuated during root induction period. The content of putrescine declined in the first three days and rose between the 3 th day and the 15 th day. The contents of IAA and IBA decreased firstly and then increased, and were the lowest at the 7 th day. The content of GA was the lowest at the 3 rd day and increased rapidly from the 3 th to the 10 th day, but then the increasing speed slowed down. The variation in ZR content followed a pattern of "decrease→increase→decrease", lowest at the 7 th day. The ratio of IAA/ABA was relatively stable during the first seven days,and increased from the 7 th day to the 15 th day. The ratio of IAA/ZR decreased slightly within the first 3 days and then increased from the 3 th day to the 15 th day. The activity of IAAO remained stable from the 7 th day to the 10 th day, and increased rapidly from the 10 th to the 15 th day, indicating that decomposition of IAA was accelerated during this period. There was a significant negative correlation between Spm and ZR contents p < 0.01, r=-0.898). Spd content had significant positive correlations(p < 0.01)with the contents of IAA and ABA, the ratios of IAA/ABA and IAA/ZR and the activity of IAAO. Put content was positively correlated with GA(p < 0.01, r=0.646), IAA and ABA(p < 0.05). IAAO activity was significantly positively correlated with PPO activity(r=0.843), Spd content(r=0.643) and IAA/ZR ratio(r=0.662). PPO enzyme activity was significantly positively correlated with IAA and ABA contents and the ratio of IAA/ZR(p < 0.01). The content of IAA had moderate correlations with the contents of Spd, GA and ABA, PPO activity and the ratios of IAA/ZR and IAA/ABA. The content of ABA had moderate correlations with the contents of IAA, Spd and GA and the ratios of IAA/ZR and IAA/ABA. The results showed that Spd, IAA and ABA played important regulation roles in rooting process.【Conclusion】The suitable rooting medium for Pyrus betulifolia Bunge was 1/2 MS + 2.0 mg· L~(-1) IAA +0.5 mg · L~(-1) IBA + 7.5 g · L~(-1) agar + 25.0 g · L~(-1) sucrose. The increase in IAA/ABA and IAA/ZR ratios induced adventitious roots formation. Spd might be closely related to the adventitious root formation. In the late stage of root induction, the increase in GA content had no significant effect on the adventitious root formation.
【Objective】Pyrus betulifolia Bunge is widely used as a pear rootstock in China. Propagation with seeds has large genetic variation and poor consistency. Difficulty in rooting limitsits propagation by traditional vegetative propagation methods such as layering and cutting. With the increasingly perfecting of in vitro propagation technology, virus-free seedlings can be produced by micropropagation method. Root induction is one of the important processes in in vitro propagation. The transplanting survival rate and seedling quality are affected by rooting efficiency. There are many internal and external factors affecting the rooting process, including endogenous hormones, polyamines, related oxidases and exogenous substances such as ethylene and salicylic acid. The study aimed at revealing the roles of endogenous hormones, polyamines and related oxidases in root induction of tissue-cultured seedlings.【Methods】Subcultured plantlets of Pyrus betulifolia Bunge were cultured on rooting medium with1/2 MS, 7.5 g · L~(-1) agar, 25.0 g · L~(-1) sucrose and different ratios of hormones. A total of 24 treatments using five different concentrations(0, 0.2, 0.5, 1 and 2 mg · L~(-1)) of IAA and IBA were assessed in terms of rooting percentages at 40 days after transfer to the media. Each treatment consisted of 6 bottles each comprising 5 subcultured plantlets. Uniform subcultured plantlets were selected to be cultured on the optimized rooting medium. Approximately 500 plantlets were used to monitor the dynamic changes in polyamines, endogenous hormones and related enzyme activities during root regeneration. All of the cultures were incubated at 25 ℃ under a 16 h light/8 h dark cycle and 2 000-3 000 lx light intensity. The basal stem cuttings were taken 0, 3, 7, 10 and 15 days after transfer to media. Polyamines, endogenous hormones and related enzyme activities were determined.【Results】The treatment of 2.0 mg · L~(-1) IAA +0.5 mg · L~(-1) IBA increased rooting rate and the percentage of plantlets with ≥3 roots compared with the other treatments. The rooting rate was 86.7% and the percentage of plantlets with ≥3 roots was53.3%. The combination of IAA and IBA was better than single hormone. Callus was observed between the 7 th and 10 th day, growing fast from the 10 th to 15 th day. The roots generated 20 days after transfer to media. The contents of three polyamines declined after 3 days in the root induction medium. The content of spermine(Spm) and spermidine(Spd) fluctuated during root induction period. The content of putrescine declined in the first three days and rose between the 3 th day and the 15 th day. The contents of IAA and IBA decreased firstly and then increased, and were the lowest at the 7 th day. The content of GA was the lowest at the 3 rd day and increased rapidly from the 3 th to the 10 th day, but then the increasing speed slowed down. The variation in ZR content followed a pattern of "decrease→increase→decrease", lowest at the 7 th day. The ratio of IAA/ABA was relatively stable during the first seven days,and increased from the 7 th day to the 15 th day. The ratio of IAA/ZR decreased slightly within the first 3 days and then increased from the 3 th day to the 15 th day. The activity of IAAO remained stable from the 7 th day to the 10 th day, and increased rapidly from the 10 th to the 15 th day, indicating that decomposition of IAA was accelerated during this period. There was a significant negative correlation between Spm and ZR contents p < 0.01, r=-0.898). Spd content had significant positive correlations(p < 0.01)with the contents of IAA and ABA, the ratios of IAA/ABA and IAA/ZR and the activity of IAAO. Put content was positively correlated with GA(p < 0.01, r=0.646), IAA and ABA(p < 0.05). IAAO activity was significantly positively correlated with PPO activity(r=0.843), Spd content(r=0.643) and IAA/ZR ratio(r=0.662). PPO enzyme activity was significantly positively correlated with IAA and ABA contents and the ratio of IAA/ZR(p < 0.01). The content of IAA had moderate correlations with the contents of Spd, GA and ABA, PPO activity and the ratios of IAA/ZR and IAA/ABA. The content of ABA had moderate correlations with the contents of IAA, Spd and GA and the ratios of IAA/ZR and IAA/ABA. The results showed that Spd, IAA and ABA played important regulation roles in rooting process.【Conclusion】The suitable rooting medium for Pyrus betulifolia Bunge was 1/2 MS + 2.0 mg· L~(-1) IAA +0.5 mg · L~(-1) IBA + 7.5 g · L~(-1) agar + 25.0 g · L~(-1) sucrose. The increase in IAA/ABA and IAA/ZR ratios induced adventitious roots formation. Spd might be closely related to the adventitious root formation. In the late stage of root induction, the increase in GA content had no significant effect on the adventitious root formation.
引文
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[2]DANIEL I P,MONICA L P,JOHN D B,JOSELI S,TIBERIA LP,MARIUSZ K,LAURENT G,EMILIE C,SALMA C,KAR-IN L,ARTHUR G F,DORU P,CATHERINE B.Identification of new adventitious rooting mutants amongst suppressors of the Arabidopsis thaliana superroot2 mutation[J].Journal of Experimental Botany,2014,65(6):1605-1618.
[3]王荣.苹果砧木茎源根系发生中次生代谢、内源激素和转录组差异分析[D].泰安:山东农业大学,2016.WANG Rong.Secondary metabolism endogenous hormone and transcriptome analysis of apple stock cutting root system[D].Tai’an:Shandong Agricultural University,2016.
[4]汪天,王素平,郭世荣.外源乙烯对黄瓜幼苗根系生长及内源多胺含量的影响[J].沈阳农业大学学报,2006(3):473-475.WANG Tian,WANG Supping,GUO Shirong.Effects of exogenous ethylene on growth of root system and contents of endogenous polyamines in cucumber seedlings[J]Journal of Shenyang Agricultural University,2006(3):473-475.
[5]王金祥,严小龙,潘瑞炽.不定根形成与植物激素的关系[J].植物生理学通讯,2005(2):133-142.WANG Jinxiang,YAN Xiaolong,PAN Ruizhi.Relationship between adventitious root formation and plant hormones[J].Plant Physiology Communications,2005(2):133-142.
[6]杨洪强,黄天栋.多胺和精氨酸对苹果实生根系的影响[J].植物学通报,1996(1):52-54.YANG Hongqiang,HUANG Tiandong.Effects of polyamines and agrinine on seedling root of apple[J].Chinese Bulletin of Botany,1996(1):52-54.
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