榛子扦插生根机理与繁殖技术的研究
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摘要
本文以平榛、杂交榛、欧榛为研究对象,通过硬枝和嫩枝扦插试验,研究了不同的扦插基质、不同生长素种类和浓度处理的榛子插条生根指标(生根率、根数、根长、生根指数)差异;用石蜡切片法研究了不定根发生的解剖学特性;追踪生根过程研究了扦插生根相关的中生理生化指标(氧化酶类、枝条的营养物质、以及内源激素)的动态变化;并对扦插苗年生长规律进行研究。旨在为榛子扦插生根和苗木培育提供理论依据和技术支撑。主要试验结果如下:
榛子硬枝、嫩枝插穗刺激愈伤组织和自由愈伤组织均能生根,但绝大多数生根部位为刺激愈伤组织。榛子扦插生根过程可分为不定根诱导期、表达期和伸长期3个阶段。
欧榛嫩枝生根解剖学研究表明,生根类型属于愈伤部位生根型。插穗扦插前不存在潜伏根原基,诱生的不定根原始体起源于木质部和韧皮部之间形成层薄壁细胞。
3种氧化酶活性在欧榛硬枝和嫩枝扦插生根过程中总体呈现出“升高-降低”的变化趋势。生根期间POD活性出现2个高峰,在不定根诱导期和表达期上升,在伸长期下降。PPO和IAAO活性在不定根诱导期上升,在表达期和伸长期活性下降。IBA处理改变了生根过程中POD、PPO和IAAO的活性和其之间的相互关系,从而影响了插穗生根,表现出不同的生根效果。
可溶性糖含量在硬枝扦插生根中呈现逐渐降低的趋势,在嫩枝扦插生根中呈现“降低-升高-降低”的变化趋势,经IBA处理的插穗可溶性糖含量显著高于对照。可溶性糖含量高,C/N比高,有利于硬枝插条生根,嫩枝插条相反,C/N比低有利于生根,C/N比理论不适用于榛子嫩枝扦插。
3种内源激素(IAA、ABA、GA3)在生根过程中变化趋势不同。硬枝插穗中IAA绝对含量明显高于嫩枝插穗,但均表现出不定根表达期、伸长期下降趋势。ABA和GA3含量与生根负相关。IAA和ABA在生根过程中含量变化互为消长关系,IAA/ABA比值可以作为衡量树木嫩枝扦插生根难易的标准。
扦插基质影响生根。不论是欧榛硬枝还是嫩枝扦插,在200mg·kg-1IBA处理条件下,3种基质(河沙、珍珠岩、蛭石和珍珠岩组成的混合基质)以混合基质生根效果最好。欧榛硬枝扦插在河沙或珍珠岩基质中不生根,在混合基质中生根率可达60%。嫩枝扦插在河沙或珍珠岩基质中生根率较低,分别为13.3%、9.17%,在混合基质中生根率为42.5%。
生长素种类和浓度影响生根效果。混合基质条件下,用3种生长素(IBA、α-NAA、ABT1)、4种浓度(0、50、200、1000mg·kg-1)处理欧榛硬枝结果表明,生长素处理可提高插穗生根率,但生根率多在50%以下,而对照不生根。IBA处理效果好于α-NAA和ABT1,浓度以200mg·kg-1处理的生根率最高,达56.7%。
不同种类榛子对IBA处理反应不同。经不同浓度(0、100、200、300、400、500、600mg·kg-1)IBA处理的榛子(平榛、杂交榛、欧榛)嫩枝扦插试验得出,500mg·kg-1IBA处理的平榛插条生根率最高,为20%,对照不生根;100和500mg·kg-1IBA处理的杂交榛插条生根率均为48.3%,对照生根率为20%。400mg·kg-1IBA处理的欧榛插穗生根率最高,可达73.3%,对照生根率仅为8.3%。3种榛子中平榛生根效果最差,杂交榛和欧榛生根效果较好,生产中育苗可用扦插繁殖。
不同粗度(0.5~1.0cm、1.0~1.5cm)、不同部位(形态学上端、形态学下端)的欧榛嫩枝扦插(200mg·kg-1IBA处理,基质为河沙)生根率差异不显著。细枝条和粗枝条插穗生根率分别为8.3%和18.3%;上端、下端枝条的插穗生根率分别为10%和8.3%。插穗的粗度、部位不是影响插条生根的主要因素。
欧榛扦插苗苗高生长曲线呈现“S”型。苗高净生长呈现“慢-快-慢-停”的生长节律。高生长和地径生长分别在7月和8月期间达到最大值,6~8月为苗木速生期,是加强栽培管理的关键时期。
Rooting mechanism and cutting propagation technique of Corylus such as herterophylla, hybrid hazelnut and avellana were studied with different auxins and concentrations on soft and hardwood cuttings under different media. The research including: Rooting capacities(rooting rate, root number, root length) of soft and hardwood cuttings; anatomical proerty of adventitious roots using paraffin section technology; dynamic changes of rooting-related physiological biochemical indexes(oxidases, nutriments, endogenesis hormones) and annual growth rules of cuttage seedlings. The aim of studies is to provide scientific basis and technological guide for Corylus propagation.
Most adventitious roots of Corylus cuttings(hard or softwood cuttings) came from stimulated callus. Both stimulated and free callus could root. The adventitious roots developmental process can be divided into 3 distinct phases, i.e., induction, expression and elongation.
The anatomical observation on softwood cuttings showed that the rooting type of avellana originated from callus. There was no induced root primordium in cuttings. The induced root primorbium which originated from the parenchyma cell of cambium between xylem and phloem developed into adventitious roots.
The activities of POD, PPO and IAAO appeared tendency of“enhancing-descending”. The activity of POD had two peaks during rooting process. The activity of POD increased up during induction and expression phase and declined at the elongation phase. The activities of PPO and IAAO increased up during induction and declined at the expression elongation phase. IBA could change the activities of the oxidases and promoted rooting process.
The contents of soluble sugar, total N and the ratio of C/N appeared different change laws. The content of soluble sugar decreased gradually during hardwood cutting rooting. The content of soluble sugar in cuttings treated by IBA had significant differences in contrast with control. The higher content of soluble sugar and ratio of C/N of hardwood cuttings was favorable to root. On the contrary, the lower ratio of C/N was benefit to softwood cuttings root. The content of soluble sugar in cuttings by IBA treatment is lower than that of control. So the theory of C/N was inapplicabe in Corylus softwood cutting.
IAA, ABA and GA3 had different change rules during rooting process. The content of IAA in hardwood cutting had significant differences in contrast with that of softwood cutting. The content of IAA in both hard and softwood declined at the expression and elongation phase. Both ABA and GA3 restrained cutting rooting. The changes of IAA content were contrary to ABA. The ratio of IAA and ABA can be as a scale estimating softwood cutting rooting capacities.
The media can affect cutting rooting. The results showed that only mixed media could enhance the percentage of hardwood cuttings rooting by IBA treatment in media experiment with river sand, perlite and mixed media. The hardwood cuttings of avellana could not root under river sand or perlite media. Rooting percentage of cutting by 200mg·kg-1IBA treatment under mixed media reached 60%. The rooting rate of softwood cutting in both river sand and perlite only reached 13.3% and 9.7%, and 42.5% in mixed media.
The kinds and concentrations of auxin affected rooting. Auxin treatment could increase rooting percentage, but the rooting rate was below 50% and control could not root. IBA had the most significant effect on rooting rate of cuttings in avellana thanα-NAA and ABT1, and the rooting rate could reach 56.7% by 200mg·kg-1IBA treatment.
Different Corylus varieties had different reaction to IBA treatment. The rooting rate reached the maximum 20% by 500mg·kg-1IBA treatment in Heterophylla, and control could not root. The rooting rate reached the maximum 48.3% by 100 or 500mg·kg-1IBA treatment in hybrid hazelnut, and the rooting percentage of cutting only reached 20% in control. Avellana reached the maximum percentage of 73.3% by 400mg·kg-1IBA treatment, and control only reached 8.3%. Heterophylla had the worst results among three varieties. Hybrid hazelnut and Avellana can be propagated by cutting.
The rooting percentage of cutting in different ends(upper and under end in morphology) and diameters(0.5~1.0cm and 1.0~1.5cm) by IBA treatment had not significant difference in Avellana. The rooting rate reached 8.3% and 18.3% in leptos and stout stress espectively, and the rooting rate reached 10% and 8.3% in upper and under end in morphology. Diameters and ends were not primary influenced factors.
Height growth curve of Avellana cutting seedling could be fitted into‘S’curve. Net height growth appeared tendency of“slow-fast-slow-stopping”. The growth of height and ground diameter reached the maximum on July and August. The cutting seedling should be managed intensively during June to August.
榛子硬枝、嫩枝插穗刺激愈伤组织和自由愈伤组织均能生根,但绝大多数生根部位为刺激愈伤组织。榛子扦插生根过程可分为不定根诱导期、表达期和伸长期3个阶段。
欧榛嫩枝生根解剖学研究表明,生根类型属于愈伤部位生根型。插穗扦插前不存在潜伏根原基,诱生的不定根原始体起源于木质部和韧皮部之间形成层薄壁细胞。
3种氧化酶活性在欧榛硬枝和嫩枝扦插生根过程中总体呈现出“升高-降低”的变化趋势。生根期间POD活性出现2个高峰,在不定根诱导期和表达期上升,在伸长期下降。PPO和IAAO活性在不定根诱导期上升,在表达期和伸长期活性下降。IBA处理改变了生根过程中POD、PPO和IAAO的活性和其之间的相互关系,从而影响了插穗生根,表现出不同的生根效果。
可溶性糖含量在硬枝扦插生根中呈现逐渐降低的趋势,在嫩枝扦插生根中呈现“降低-升高-降低”的变化趋势,经IBA处理的插穗可溶性糖含量显著高于对照。可溶性糖含量高,C/N比高,有利于硬枝插条生根,嫩枝插条相反,C/N比低有利于生根,C/N比理论不适用于榛子嫩枝扦插。
3种内源激素(IAA、ABA、GA3)在生根过程中变化趋势不同。硬枝插穗中IAA绝对含量明显高于嫩枝插穗,但均表现出不定根表达期、伸长期下降趋势。ABA和GA3含量与生根负相关。IAA和ABA在生根过程中含量变化互为消长关系,IAA/ABA比值可以作为衡量树木嫩枝扦插生根难易的标准。
扦插基质影响生根。不论是欧榛硬枝还是嫩枝扦插,在200mg·kg-1IBA处理条件下,3种基质(河沙、珍珠岩、蛭石和珍珠岩组成的混合基质)以混合基质生根效果最好。欧榛硬枝扦插在河沙或珍珠岩基质中不生根,在混合基质中生根率可达60%。嫩枝扦插在河沙或珍珠岩基质中生根率较低,分别为13.3%、9.17%,在混合基质中生根率为42.5%。
生长素种类和浓度影响生根效果。混合基质条件下,用3种生长素(IBA、α-NAA、ABT1)、4种浓度(0、50、200、1000mg·kg-1)处理欧榛硬枝结果表明,生长素处理可提高插穗生根率,但生根率多在50%以下,而对照不生根。IBA处理效果好于α-NAA和ABT1,浓度以200mg·kg-1处理的生根率最高,达56.7%。
不同种类榛子对IBA处理反应不同。经不同浓度(0、100、200、300、400、500、600mg·kg-1)IBA处理的榛子(平榛、杂交榛、欧榛)嫩枝扦插试验得出,500mg·kg-1IBA处理的平榛插条生根率最高,为20%,对照不生根;100和500mg·kg-1IBA处理的杂交榛插条生根率均为48.3%,对照生根率为20%。400mg·kg-1IBA处理的欧榛插穗生根率最高,可达73.3%,对照生根率仅为8.3%。3种榛子中平榛生根效果最差,杂交榛和欧榛生根效果较好,生产中育苗可用扦插繁殖。
不同粗度(0.5~1.0cm、1.0~1.5cm)、不同部位(形态学上端、形态学下端)的欧榛嫩枝扦插(200mg·kg-1IBA处理,基质为河沙)生根率差异不显著。细枝条和粗枝条插穗生根率分别为8.3%和18.3%;上端、下端枝条的插穗生根率分别为10%和8.3%。插穗的粗度、部位不是影响插条生根的主要因素。
欧榛扦插苗苗高生长曲线呈现“S”型。苗高净生长呈现“慢-快-慢-停”的生长节律。高生长和地径生长分别在7月和8月期间达到最大值,6~8月为苗木速生期,是加强栽培管理的关键时期。
Rooting mechanism and cutting propagation technique of Corylus such as herterophylla, hybrid hazelnut and avellana were studied with different auxins and concentrations on soft and hardwood cuttings under different media. The research including: Rooting capacities(rooting rate, root number, root length) of soft and hardwood cuttings; anatomical proerty of adventitious roots using paraffin section technology; dynamic changes of rooting-related physiological biochemical indexes(oxidases, nutriments, endogenesis hormones) and annual growth rules of cuttage seedlings. The aim of studies is to provide scientific basis and technological guide for Corylus propagation.
Most adventitious roots of Corylus cuttings(hard or softwood cuttings) came from stimulated callus. Both stimulated and free callus could root. The adventitious roots developmental process can be divided into 3 distinct phases, i.e., induction, expression and elongation.
The anatomical observation on softwood cuttings showed that the rooting type of avellana originated from callus. There was no induced root primordium in cuttings. The induced root primorbium which originated from the parenchyma cell of cambium between xylem and phloem developed into adventitious roots.
The activities of POD, PPO and IAAO appeared tendency of“enhancing-descending”. The activity of POD had two peaks during rooting process. The activity of POD increased up during induction and expression phase and declined at the elongation phase. The activities of PPO and IAAO increased up during induction and declined at the expression elongation phase. IBA could change the activities of the oxidases and promoted rooting process.
The contents of soluble sugar, total N and the ratio of C/N appeared different change laws. The content of soluble sugar decreased gradually during hardwood cutting rooting. The content of soluble sugar in cuttings treated by IBA had significant differences in contrast with control. The higher content of soluble sugar and ratio of C/N of hardwood cuttings was favorable to root. On the contrary, the lower ratio of C/N was benefit to softwood cuttings root. The content of soluble sugar in cuttings by IBA treatment is lower than that of control. So the theory of C/N was inapplicabe in Corylus softwood cutting.
IAA, ABA and GA3 had different change rules during rooting process. The content of IAA in hardwood cutting had significant differences in contrast with that of softwood cutting. The content of IAA in both hard and softwood declined at the expression and elongation phase. Both ABA and GA3 restrained cutting rooting. The changes of IAA content were contrary to ABA. The ratio of IAA and ABA can be as a scale estimating softwood cutting rooting capacities.
The media can affect cutting rooting. The results showed that only mixed media could enhance the percentage of hardwood cuttings rooting by IBA treatment in media experiment with river sand, perlite and mixed media. The hardwood cuttings of avellana could not root under river sand or perlite media. Rooting percentage of cutting by 200mg·kg-1IBA treatment under mixed media reached 60%. The rooting rate of softwood cutting in both river sand and perlite only reached 13.3% and 9.7%, and 42.5% in mixed media.
The kinds and concentrations of auxin affected rooting. Auxin treatment could increase rooting percentage, but the rooting rate was below 50% and control could not root. IBA had the most significant effect on rooting rate of cuttings in avellana thanα-NAA and ABT1, and the rooting rate could reach 56.7% by 200mg·kg-1IBA treatment.
Different Corylus varieties had different reaction to IBA treatment. The rooting rate reached the maximum 20% by 500mg·kg-1IBA treatment in Heterophylla, and control could not root. The rooting rate reached the maximum 48.3% by 100 or 500mg·kg-1IBA treatment in hybrid hazelnut, and the rooting percentage of cutting only reached 20% in control. Avellana reached the maximum percentage of 73.3% by 400mg·kg-1IBA treatment, and control only reached 8.3%. Heterophylla had the worst results among three varieties. Hybrid hazelnut and Avellana can be propagated by cutting.
The rooting percentage of cutting in different ends(upper and under end in morphology) and diameters(0.5~1.0cm and 1.0~1.5cm) by IBA treatment had not significant difference in Avellana. The rooting rate reached 8.3% and 18.3% in leptos and stout stress espectively, and the rooting rate reached 10% and 8.3% in upper and under end in morphology. Diameters and ends were not primary influenced factors.
Height growth curve of Avellana cutting seedling could be fitted into‘S’curve. Net height growth appeared tendency of“slow-fast-slow-stopping”. The growth of height and ground diameter reached the maximum on July and August. The cutting seedling should be managed intensively during June to August.
引文
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