能源油料植物光皮树花的发育研究
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摘要
光皮树[Swida Cornus wilsonian(Wanger.)Sojak]是山茱萸科徕木属落叶灌木或乔木,是一种生态适应性广的木本油料植物,其果实和种子油以油酸和亚油酸为主,是理想的生物柴油原料油料和食用油。因其适宜在瘠薄干旱的丘陵山地和非耕地种植,成为最具开发潜能的木本油料植物之一,近年来国家对该树种的推广栽培、加工利用给予了大力支持。
本文以生长在湖南省林业科学院试验林场—施公坡实验基地4年生同品种、树势健壮、植株大小及生长势基本一致的光皮树嫁接植株为研究对象,分别取其顶芽和腋芽,利用常规石蜡切片法与野外观测相结合对芽体的外部形态变化和解剖结构进行观测分析,了解花芽、叶芽形态结构变化规律；采集有花和无花光皮树植株一年生枝嫩叶,采用间接酶联免疫吸附法测定光皮树花芽分化过程中4种内源激素含量变化,采用苯酚-硫酸法测定叶内碳水化合物含量变化。首次较为系统地观测研究光皮树花芽的分化发育特点,探索花芽分化与内源激素及多糖含量变化的相关性,其研究结果如下：
1、光皮树的顶芽和腋芽上都可进行花芽分化,在湖南中部地区(如长沙)的气候条件下,花芽分化在2月下旬开始,4月下旬基本完成。花芽形态分化过程可划分为：潜伏(未)分化期、花序分化期、萼片分化期、花瓣分化期和雌、雄蕊分化期5个时期。
2、花芽分化不同时期4种内源激素含量存在显著差异。有花芽分化的光皮树保持相对较低和稳定的IAA.GA3含量,ABA含量高且变化幅度较大,ZR含量相对较低,但随着花的分化发育含量逐渐升高；而在相同的生长季内,无花光皮树IAA.GA3含量先升高后降低,ABA含量先逐渐升高,然后下降,而ZR含量呈现出先逐渐升高然后降低,再升高再下降的变化趋势。从被分析的4种内源激素的绝对值来看,IAA平均含量,显著低于所分析的其他3种内源激素,ABA的平均含量较高,且变化波动比较大。4种内源激素含量的动态变化影响光皮树的成花过程。
3、碳水化合物含量随着花芽分化而不断升高,并且高于花芽分化前期的水平,表明一定量碳水化合物的积累有利于花芽分化。
4、光皮树花芽分化过程中各类内源激素相对含量发生变化：ABA /GA3与ZR/IAA变化趋势相近,其含量较稳定；ABA /IAA及ABA /GA3均在花芽孕育期处于低水平,花芽形态分化期逐渐升高,随后在雌雄蕊形成期逐渐下降。较高水平的ABA /IAA及ABA /GA3有利于花芽形态的分化,而较低的ZR /GA3有利于花芽分化。
Swida Cornus wilsonian (Wanger.) Sojak is a kind of deciduous arbors or shrub belonging to Cornaceae,whose height is 5-18 m and diameter may reach 55 cm.The bark is purple or greyish white with green,which is smooth and maculosus abscission.Its crown is umbrella shape,the leaves are oblong,and its inflorescence is cyme with small white flower.The mature drupes of Swida wilsoniana are sphere with black pericarp,and the seeds are yellowish Swida wilsoniana mainly distributed south of the Huanghe River valley, particularly in these provinces such as Hunan,Jiangxi,Hubei.Its vertical distribution is below elevation 1000 m. Swida wilsoniana is a heliophilous plant which has a strong Suitability to different soil.And Swida wilsoniana is also one kind of oil plant with extensive ecological adaptability. Oil of the fruit and seed richening with oleic acid and linoleic acid was an ideal raw material for producing bio-diesel. Because of its suitable in infertile arid hill mountainous region and non-farming land,it is one of the bio-energy plants with great developing potentiality In recent years the State Government has been supporting the extensive cultivation,process and utilization of this tree species.
In this paper researches on floral differentiation of Swida wilsoniana was carried out at forest experimental farm of Hunan Academy of Forestry which located at Shi Gongpo,Changsha city.In the experiment,4-year old grafted,vigorous and healthy adult plants were selected as the research target. All the selected plants were the same variety, and had similar size,located at the same site conditions.
Terminal bud and the axial bud from blossom and blossomless plants were collected separately from the selected plants to investigate their morphological and anatomic structures by using conventional wax slicing method,and direct field observations;Meanwhile tender leaves from one year branches of the same plants as buds were collected to analyze the endogenous hormones contents by means of indirect enzyme-linked immunosorbent assay;Also carbohydrate content in the leaves was determined by means of Phenol-sulfuric acid Method. It's the first time to study the flower buds differentiation and development of Swida wilsoniana,and to analyze the correlation between the flower differentiation and endogenous hormones and carbohydrate. The experimental results showed as followings:
1. Flower bud differentiation occurred both at terminal buds and axiliary buds. Under the climate conditions of Central part of Hunan Province,the flower bud differentiation started at late February and ended around the end of April. The whole process of flower bud differentiation might be divided into five stages,including pre-differentiation, Inflorescence differentiation,Sepal differentiation,Petal differentiation and Stamens and pistils differentiation stage.
2. There was a great difference in content change of endogenous hormones between flowering plants and non-flowering plants of Swida wilsoniana. During floral bud differentiation, IAA and GA3 were at relatively low and steady level,ABA was at a higher level with fluctuation change,ZR firstly was at a relatively low level and increased gradually along with the development of flower. Meanwhile,for the non-flowering plants, both IAA and ZR contents changed as increased and decresased,then increased and decreased again,ABA increased at initial time and then decreased gradually;GA3 firstly increased and then decreased.As to the absolute contents of the four kinds of endogenous hormone,the average content of IAA was obviously lower than that of the other three kinds of endogenous hormone which were analyzed. Of the analized endogene hormones,the average content of ABA was higher,and its change fluctuation was quite large.It's suggested that the dynamic changes of endogenous were related with the flowering process.
3. The content of carbohydrate increased unceasingly along with the differentiation of flower bud, and it was higher than that during the earlier stage of flower bud differentiation, which suggested that the accumulation of carbohydrate was advantageous to the differentiation of flower bud.
4. Changes of endogene hormone in the flower bud differentiation process of Swida wilsoniana indicated that:Changes of ABA/ GA3 and ZR/IAA were very similar, whose content were stable; the ABA /IAA and ABA /GA3 content were at the low levels during the floral initiation period, then increased gradually during flower bud morphological differentiation, and decreased gradually during the pistil and stamen formation period. Those suggested that the high level of ABA /IAA was advantageous to the morphological differentiation of flower bud, while the low level of ZR /GA3 was advantageous to flower bud initiation.
本文以生长在湖南省林业科学院试验林场—施公坡实验基地4年生同品种、树势健壮、植株大小及生长势基本一致的光皮树嫁接植株为研究对象,分别取其顶芽和腋芽,利用常规石蜡切片法与野外观测相结合对芽体的外部形态变化和解剖结构进行观测分析,了解花芽、叶芽形态结构变化规律；采集有花和无花光皮树植株一年生枝嫩叶,采用间接酶联免疫吸附法测定光皮树花芽分化过程中4种内源激素含量变化,采用苯酚-硫酸法测定叶内碳水化合物含量变化。首次较为系统地观测研究光皮树花芽的分化发育特点,探索花芽分化与内源激素及多糖含量变化的相关性,其研究结果如下：
1、光皮树的顶芽和腋芽上都可进行花芽分化,在湖南中部地区(如长沙)的气候条件下,花芽分化在2月下旬开始,4月下旬基本完成。花芽形态分化过程可划分为：潜伏(未)分化期、花序分化期、萼片分化期、花瓣分化期和雌、雄蕊分化期5个时期。
2、花芽分化不同时期4种内源激素含量存在显著差异。有花芽分化的光皮树保持相对较低和稳定的IAA.GA3含量,ABA含量高且变化幅度较大,ZR含量相对较低,但随着花的分化发育含量逐渐升高；而在相同的生长季内,无花光皮树IAA.GA3含量先升高后降低,ABA含量先逐渐升高,然后下降,而ZR含量呈现出先逐渐升高然后降低,再升高再下降的变化趋势。从被分析的4种内源激素的绝对值来看,IAA平均含量,显著低于所分析的其他3种内源激素,ABA的平均含量较高,且变化波动比较大。4种内源激素含量的动态变化影响光皮树的成花过程。
3、碳水化合物含量随着花芽分化而不断升高,并且高于花芽分化前期的水平,表明一定量碳水化合物的积累有利于花芽分化。
4、光皮树花芽分化过程中各类内源激素相对含量发生变化：ABA /GA3与ZR/IAA变化趋势相近,其含量较稳定；ABA /IAA及ABA /GA3均在花芽孕育期处于低水平,花芽形态分化期逐渐升高,随后在雌雄蕊形成期逐渐下降。较高水平的ABA /IAA及ABA /GA3有利于花芽形态的分化,而较低的ZR /GA3有利于花芽分化。
Swida Cornus wilsonian (Wanger.) Sojak is a kind of deciduous arbors or shrub belonging to Cornaceae,whose height is 5-18 m and diameter may reach 55 cm.The bark is purple or greyish white with green,which is smooth and maculosus abscission.Its crown is umbrella shape,the leaves are oblong,and its inflorescence is cyme with small white flower.The mature drupes of Swida wilsoniana are sphere with black pericarp,and the seeds are yellowish Swida wilsoniana mainly distributed south of the Huanghe River valley, particularly in these provinces such as Hunan,Jiangxi,Hubei.Its vertical distribution is below elevation 1000 m. Swida wilsoniana is a heliophilous plant which has a strong Suitability to different soil.And Swida wilsoniana is also one kind of oil plant with extensive ecological adaptability. Oil of the fruit and seed richening with oleic acid and linoleic acid was an ideal raw material for producing bio-diesel. Because of its suitable in infertile arid hill mountainous region and non-farming land,it is one of the bio-energy plants with great developing potentiality In recent years the State Government has been supporting the extensive cultivation,process and utilization of this tree species.
In this paper researches on floral differentiation of Swida wilsoniana was carried out at forest experimental farm of Hunan Academy of Forestry which located at Shi Gongpo,Changsha city.In the experiment,4-year old grafted,vigorous and healthy adult plants were selected as the research target. All the selected plants were the same variety, and had similar size,located at the same site conditions.
Terminal bud and the axial bud from blossom and blossomless plants were collected separately from the selected plants to investigate their morphological and anatomic structures by using conventional wax slicing method,and direct field observations;Meanwhile tender leaves from one year branches of the same plants as buds were collected to analyze the endogenous hormones contents by means of indirect enzyme-linked immunosorbent assay;Also carbohydrate content in the leaves was determined by means of Phenol-sulfuric acid Method. It's the first time to study the flower buds differentiation and development of Swida wilsoniana,and to analyze the correlation between the flower differentiation and endogenous hormones and carbohydrate. The experimental results showed as followings:
1. Flower bud differentiation occurred both at terminal buds and axiliary buds. Under the climate conditions of Central part of Hunan Province,the flower bud differentiation started at late February and ended around the end of April. The whole process of flower bud differentiation might be divided into five stages,including pre-differentiation, Inflorescence differentiation,Sepal differentiation,Petal differentiation and Stamens and pistils differentiation stage.
2. There was a great difference in content change of endogenous hormones between flowering plants and non-flowering plants of Swida wilsoniana. During floral bud differentiation, IAA and GA3 were at relatively low and steady level,ABA was at a higher level with fluctuation change,ZR firstly was at a relatively low level and increased gradually along with the development of flower. Meanwhile,for the non-flowering plants, both IAA and ZR contents changed as increased and decresased,then increased and decreased again,ABA increased at initial time and then decreased gradually;GA3 firstly increased and then decreased.As to the absolute contents of the four kinds of endogenous hormone,the average content of IAA was obviously lower than that of the other three kinds of endogenous hormone which were analyzed. Of the analized endogene hormones,the average content of ABA was higher,and its change fluctuation was quite large.It's suggested that the dynamic changes of endogenous were related with the flowering process.
3. The content of carbohydrate increased unceasingly along with the differentiation of flower bud, and it was higher than that during the earlier stage of flower bud differentiation, which suggested that the accumulation of carbohydrate was advantageous to the differentiation of flower bud.
4. Changes of endogene hormone in the flower bud differentiation process of Swida wilsoniana indicated that:Changes of ABA/ GA3 and ZR/IAA were very similar, whose content were stable; the ABA /IAA and ABA /GA3 content were at the low levels during the floral initiation period, then increased gradually during flower bud morphological differentiation, and decreased gradually during the pistil and stamen formation period. Those suggested that the high level of ABA /IAA was advantageous to the morphological differentiation of flower bud, while the low level of ZR /GA3 was advantageous to flower bud initiation.
引文
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