厚竹群落孕笋成竹期内源激素动态变化与调控作用
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摘要
为了研究厚竹群落在孕笋成竹期内源激素的分布特征和动态变化规律,探讨内源激素及其相互作用对厚竹孕笋成竹的影响,采集不同时期的竹鞭、竹笋和母竹样品,通过酶联免疫吸附法(ELISA),测定吲哚-3-乙酸(IAA)、赤霉素(GA)、玉米素核苷(ZR)和脱落酸(ABA)的质量分数,通过方差分析和多重比较方法分析内源激素质量分数及其比值的差异。结果表明:厚竹群落中IAA和ABA质量分数显著高于ZR和GA,但IAA和ABA之间及GA和ZR之间的差异因生长系统及发育期不同而程度不同。笋芽膨大期,竹鞭、竹笋和母竹中内源激素质量分数均是ABA> IAA> GA> ZR,竹鞭中促进生长类激素偏高,ABA显著偏低,冬笋中ZR和GA质量分数占内源激素总质量分数的比例显著高于竹鞭和母竹。竹笋生长期,竹鞭和母竹中是ABA>IAA> ZR> GA,春笋中是IAA> ABA> ZR> GA;春笋中的ZR质量分数及所占比例显著高于竹鞭,ABA的质量分数及所占比例显著低于竹鞭,GA/IAA值显著高于母竹;母竹中的内源激素总量明显偏多,并且是促进生长类激素比例较高,ZR/GA值显著高于竹鞭。新竹长成期,3个生长系统中均是ABA> IAA> ZR> GA;母竹中IAA质量分数显著偏低,ABA质量分数显著偏高。在孕笋成竹过程中,母竹和竹鞭中的IAA和ABA动态变化规律刚好相反。在新竹个体发育中,GA/ABA和GA/IAA值逐渐降低,其他激素间的比值均呈抛物线变化。综合分析得出:厚竹群落中IAA和ABA的质量分数显著偏高;在孕笋成竹过程中,不同生长系统及不同内源激素的分布特征和动态变化规律不同;内源激素及相互间的协同或拮抗作用对孕笋成竹有显著调控作用,相对较高的IAA和ZR质量分数可能是竹壁增厚的原因之一。
In orderto study the distribution characteristics and dynamic changes of endogenous hormones in the Phyllostachys edulis‘Pachyloen' community, and to explore the regulation of endogenous hormones and their interactions on Ph. edulis ‘Pachyloen'during the developing process bamboo shoots, the bamboo rhizomes, bamboo shoots and maternal bamboos from different periods were collected. Bamboo samples were assayed for the mass fraction of indole-3-acetic acid(IAA), gibberellin(GA), zeatin nucleoside(ZR)and abscisic acid(ABA) by enzyme-linked immunosorbent assay(ELISA). Analysis of Variance and multiple comparison methods were used to analyze differences in endogenous hormone mass fractions and their ratios. The results showed that the IAA and ABA mass fractions in Ph. edulis ‘Pachyloen' were significantly higher than those in GA and ZR, but the differences between IAA and ABA and between GA and ZR were different depending on the growth systems and developmental period. During the bamboo shoots sprouts period,the mass fractions of endogenous hormones in the three growth systems were all ABA > IAA > ZR > GA. The promoted hormone were high in bamboo rhizomes, and ABA was significantly lower. The ratio of ZR and GA mass fractions to total endogenous hormone mass fractions in winter shoots was significantly higher than that of bamboo rhizomes and maternal bamboos. In the bamboo shoots growing period, the mass fractions of endogenous hormones in bamboo rhizomes and maternal bamboos were all ABA > IAA > ZR > GA, and in spring shoots was IAA > ABA > ZR > GA.The ZR mass fraction and proportion in the spring shoots were significantly higher than those in the bamboo rhizomes. The mass fraction and proportion of ABA in spring shoots were significantly lower than that of bamboo rhizomes, and the ratio of GA/IAA was significantly higher than that of maternal bamboo. The total amount of endogenous hormones in maternal bamboo were significantly higher, especially the relative content of promote growth hormones were on the high side, and ZR/GA was significantly higher than that of bamboo rhizomes. During the new bamboo formation period, the mass fractions of endogenous hormones in the three growth systems were all ABA > IAA > ZR > GA. In maternal bamboo, IAA was significantly lower than that in bamboo rhizome and new bamboo, and ABA was significantly higher than that in bamboo rhizomes and new bamboos. In the process of growing bamboo shoots, the dynamic changes of IAA and ABA in maternal bamboos and bamboo rhizomes were just the opposite. In the development of new bamboos, the values of GA/ABA and GA/IAA gradually decreased, and the ratio of other endogenous hormones showed parabolic changes. According to the comprehensive analysis,the mass fractions of IAA and ABA in Ph. edulis ‘Pachyloen'community were significantly higher. In the process of growing bamboo shoots, the distribution characteristics and dynamic changes of different growth systems and different endogenous hormones were different. Endogenous hormones and their synergistic or antagonistic effects had a significant regulatory effect on the developing process of bamboo shoots, and the relatively high IAA and ZR mass fractions may be one of the reasons for the thickening of the bamboo wall.
In orderto study the distribution characteristics and dynamic changes of endogenous hormones in the Phyllostachys edulis‘Pachyloen' community, and to explore the regulation of endogenous hormones and their interactions on Ph. edulis ‘Pachyloen'during the developing process bamboo shoots, the bamboo rhizomes, bamboo shoots and maternal bamboos from different periods were collected. Bamboo samples were assayed for the mass fraction of indole-3-acetic acid(IAA), gibberellin(GA), zeatin nucleoside(ZR)and abscisic acid(ABA) by enzyme-linked immunosorbent assay(ELISA). Analysis of Variance and multiple comparison methods were used to analyze differences in endogenous hormone mass fractions and their ratios. The results showed that the IAA and ABA mass fractions in Ph. edulis ‘Pachyloen' were significantly higher than those in GA and ZR, but the differences between IAA and ABA and between GA and ZR were different depending on the growth systems and developmental period. During the bamboo shoots sprouts period,the mass fractions of endogenous hormones in the three growth systems were all ABA > IAA > ZR > GA. The promoted hormone were high in bamboo rhizomes, and ABA was significantly lower. The ratio of ZR and GA mass fractions to total endogenous hormone mass fractions in winter shoots was significantly higher than that of bamboo rhizomes and maternal bamboos. In the bamboo shoots growing period, the mass fractions of endogenous hormones in bamboo rhizomes and maternal bamboos were all ABA > IAA > ZR > GA, and in spring shoots was IAA > ABA > ZR > GA.The ZR mass fraction and proportion in the spring shoots were significantly higher than those in the bamboo rhizomes. The mass fraction and proportion of ABA in spring shoots were significantly lower than that of bamboo rhizomes, and the ratio of GA/IAA was significantly higher than that of maternal bamboo. The total amount of endogenous hormones in maternal bamboo were significantly higher, especially the relative content of promote growth hormones were on the high side, and ZR/GA was significantly higher than that of bamboo rhizomes. During the new bamboo formation period, the mass fractions of endogenous hormones in the three growth systems were all ABA > IAA > ZR > GA. In maternal bamboo, IAA was significantly lower than that in bamboo rhizome and new bamboo, and ABA was significantly higher than that in bamboo rhizomes and new bamboos. In the process of growing bamboo shoots, the dynamic changes of IAA and ABA in maternal bamboos and bamboo rhizomes were just the opposite. In the development of new bamboos, the values of GA/ABA and GA/IAA gradually decreased, and the ratio of other endogenous hormones showed parabolic changes. According to the comprehensive analysis,the mass fractions of IAA and ABA in Ph. edulis ‘Pachyloen'community were significantly higher. In the process of growing bamboo shoots, the distribution characteristics and dynamic changes of different growth systems and different endogenous hormones were different. Endogenous hormones and their synergistic or antagonistic effects had a significant regulatory effect on the developing process of bamboo shoots, and the relatively high IAA and ZR mass fractions may be one of the reasons for the thickening of the bamboo wall.
引文
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[11]于婷,李建贵,侍瑞,等.骏枣果实中的内源激素含量与其生理落果的关系[J].经济林研究,2016,34(2):45-48.
[12]ZHU W Z,WANG S G,YU D Z,et al.Elevational patterns of endogenous hormones and their relation to resprouting ability of Quercus aquifolioides plants on the eastern edge of the Tibetan Plateau[J].Trees-Structure and Function,2014,28(2):359-372.
[13]MIAO Y Y,ZHU Z B,GUO Q S,et al.Dynamic changes in carbohydrate metabolism and endogenous hormones during Tulipa edulis stolon development into a new bulb[J].Journal of Plant Biology,2016,59(2):121-132.
[14]黄坚钦,刘力,章滨森,等.雷竹地下鞭侧芽内源激素的动态变化研究[J].林业科学,2002,38(3):38-41.
[15]王波,汪宏奎,李琴,等.春笋冬出毛竹的营养成分、氨基酸和激素的比较研究[J].浙江林业科技,2015,35(3):13-16.
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[18]张艳华,雷先高,黎祖尧,等.厚竹内源激素及新竹发育对繁殖方式的响应[J].竹子学报,2016,35(4):14-19.
[19]张艳华,杨光耀,黎祖尧,等.厚竹新竹发育过程中母竹内源激素动态变化[J].江西农业大学学报,2017,39(4):731-738.
[20]KOF E M,VINOGRADOVA I A,OORZHAK A S,et al.ABAcontent in shoots and roots of pea mutants af and tl as related to their growth and morphogenesis[J].Russian Journal of Plant Physiology,2006,53(3):359-365.
[21]JONG M D,MARIANI C,VRIEZEN W H.The role of auxin and gibberellin in tomato fruit set[J].Journal of Experimental Botany.2009,60(5):1523-1532.
[22]杨荣超,张海军,王倩,等.植物激素对种子休眠和萌发调控机理的研究进展[J].草地学报,2012,20(1):1-9.
[23]JAVID M G,SOROOSHZADEH A,SANAVY S A M M,et al.Effects of the exogenous application of auxin and cytokinin on carbohydrate accumulation in grains of rice under salt stress[J].Plant Growth Regulat,2011,65(2):305-313.
[24]WANG H Y,CUI K H,CAI Y,et al.Endogenous hormonal equilibrium linked to bamboo culm development.[J].Genetics and Molecular Research,2015,14(3):11312-11323.
[25]郑郁善,洪伟,邱尔发,等.毛竹出笋期各器官激素分布特征的研究[J].林业科学,1998,34(1):100-104.
[26]潘瑞炽,王小菁,李娘辉.植物生理学[M].北京:高等教育出版社,2012:211-215.
[27]ZHANG F,WAN X Q,ZHANG H Q,et al.The effect of cold stress on endogenous hormones and CBF 1 homolog in four contrasting bamboo species[J].Journal of Forest Research,2012,17(1):72-78.
[28]LIU X H,WU Y.Responding relationship between endogenous hormones and hyperspectra reflectance for the different flowering stages of the Giant Panda’s staple-food bamboos in Qinling[J].Spectroscopy and Spectral Analysis,2013,33(9):2512-2518.
[29]吴耀荣,谢旗.ABA与植物胁迫抗性[J].植物学通报,2006,23(5):511-518.
[30]王兴,于晶,杨阳,等.低温条件下不同抗寒性冬小麦内源激素的变化[J].麦类作物学报,2009,29(5):827-831.
[31]李丽霞,梁宗锁,魏宇昆,等.土壤干旱胁迫下沙棘休眠、萌芽期内源激素变化及外源GA3的调节[J].西北林学院学报,2001,16(2):10-14.
[32]何奇江,汪奎宏,华锡奇.秋季出笋雷竹的氨基酸、激素和营养成分的分析研究[J].科技通报,2006,22(4):477-481.
[2]陈建华,晏存育,王艳梅,等.6种不同变异类型毛竹叶片的生理特性研究[J].中南林业科技大学学报,2011,31(4):70-73.
[3]施建敏,杨光耀,杨清培,等.厚壁毛竹光合作用对环境因子响应的季节变化[J].林业科学研究,2009,22(6):872-877.
[4]李建,杨清培,施建敏,等.当年生厚壁毛竹光合生理动态研究[J].江西农业大学学报,2010,32(4):763-767.
[5]李健,杨清培,方楷,等.厚壁毛竹与毛竹的抗寒性生理比较研究[J].江西农业大学学报,2011,33(3):537-541.
[6]杜天真,杨光耀,郭起荣,等.厚皮毛竹春笋营养成分研究[J].江西林业科技,1997(6):1-2.
[7]李苑,张令,黎祖尧,等.不同产地厚竹笋的营养成分及其对土壤养分的响应特征[J].经济林研究,2017,35(4):147-154.
[8]孙娅东,张文根,黎祖尧,等.不同引种地厚竹纤维形态和含量比较[J].中南林业科技大学学报,2017,37(12):51-57.
[9]张雷,杨光耀,黎祖尧,等.不同产地厚竹秆形结构比较[J].竹子学报,2017,36(1):19-24.
[10]李培旺,陈景震,张良波.等.蓖麻叶片内源激素变化对主花序开花结实的影响[J].经济林研究,2016,34(4):113-116.
[11]于婷,李建贵,侍瑞,等.骏枣果实中的内源激素含量与其生理落果的关系[J].经济林研究,2016,34(2):45-48.
[12]ZHU W Z,WANG S G,YU D Z,et al.Elevational patterns of endogenous hormones and their relation to resprouting ability of Quercus aquifolioides plants on the eastern edge of the Tibetan Plateau[J].Trees-Structure and Function,2014,28(2):359-372.
[13]MIAO Y Y,ZHU Z B,GUO Q S,et al.Dynamic changes in carbohydrate metabolism and endogenous hormones during Tulipa edulis stolon development into a new bulb[J].Journal of Plant Biology,2016,59(2):121-132.
[14]黄坚钦,刘力,章滨森,等.雷竹地下鞭侧芽内源激素的动态变化研究[J].林业科学,2002,38(3):38-41.
[15]王波,汪宏奎,李琴,等.春笋冬出毛竹的营养成分、氨基酸和激素的比较研究[J].浙江林业科技,2015,35(3):13-16.
[16]方楷,杨光耀,杨清培,等.毛竹成竹过程中内源激素动态变化[J].江西农业大学学报,2011,33(6):1107-1111.
[17]方楷,杨光耀,杨清培,等.厚壁毛竹展叶期竹秆内源激素特征[J].江西农业大学学报,2012,34(1):54-58.
[18]张艳华,雷先高,黎祖尧,等.厚竹内源激素及新竹发育对繁殖方式的响应[J].竹子学报,2016,35(4):14-19.
[19]张艳华,杨光耀,黎祖尧,等.厚竹新竹发育过程中母竹内源激素动态变化[J].江西农业大学学报,2017,39(4):731-738.
[20]KOF E M,VINOGRADOVA I A,OORZHAK A S,et al.ABAcontent in shoots and roots of pea mutants af and tl as related to their growth and morphogenesis[J].Russian Journal of Plant Physiology,2006,53(3):359-365.
[21]JONG M D,MARIANI C,VRIEZEN W H.The role of auxin and gibberellin in tomato fruit set[J].Journal of Experimental Botany.2009,60(5):1523-1532.
[22]杨荣超,张海军,王倩,等.植物激素对种子休眠和萌发调控机理的研究进展[J].草地学报,2012,20(1):1-9.
[23]JAVID M G,SOROOSHZADEH A,SANAVY S A M M,et al.Effects of the exogenous application of auxin and cytokinin on carbohydrate accumulation in grains of rice under salt stress[J].Plant Growth Regulat,2011,65(2):305-313.
[24]WANG H Y,CUI K H,CAI Y,et al.Endogenous hormonal equilibrium linked to bamboo culm development.[J].Genetics and Molecular Research,2015,14(3):11312-11323.
[25]郑郁善,洪伟,邱尔发,等.毛竹出笋期各器官激素分布特征的研究[J].林业科学,1998,34(1):100-104.
[26]潘瑞炽,王小菁,李娘辉.植物生理学[M].北京:高等教育出版社,2012:211-215.
[27]ZHANG F,WAN X Q,ZHANG H Q,et al.The effect of cold stress on endogenous hormones and CBF 1 homolog in four contrasting bamboo species[J].Journal of Forest Research,2012,17(1):72-78.
[28]LIU X H,WU Y.Responding relationship between endogenous hormones and hyperspectra reflectance for the different flowering stages of the Giant Panda’s staple-food bamboos in Qinling[J].Spectroscopy and Spectral Analysis,2013,33(9):2512-2518.
[29]吴耀荣,谢旗.ABA与植物胁迫抗性[J].植物学通报,2006,23(5):511-518.
[30]王兴,于晶,杨阳,等.低温条件下不同抗寒性冬小麦内源激素的变化[J].麦类作物学报,2009,29(5):827-831.
[31]李丽霞,梁宗锁,魏宇昆,等.土壤干旱胁迫下沙棘休眠、萌芽期内源激素变化及外源GA3的调节[J].西北林学院学报,2001,16(2):10-14.
[32]何奇江,汪奎宏,华锡奇.秋季出笋雷竹的氨基酸、激素和营养成分的分析研究[J].科技通报,2006,22(4):477-481.
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