绒毛白蜡引种及白蜡属内种间杂交育种研究
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摘要
我国有丰富的白蜡属树种资源,绒毛白蜡(Fraxinus velutina Torr.)是其中引种成功的树种之一,由于其耐盐性强、观赏性强和分布范围广等特点,被国内许多省市广泛引种栽培。目前已成为部分沿海城市及盐碱地区造林和绿化的优良树种,但黑龙江尚未引种。本文以绒毛白蜡引种为主线,从苗木引种、种子育苗、耐盐碱试验、嫁接技术等几个方面进行了深入研究,以期为黑龙江地区引种提供基础资料和技术保障。此外,为了改善白蜡属树种的品质,整合优良的基因资源,本文着重开展了水曲柳与白蜡属内种间三个树种的杂交育种技术研究,主要研究内容如下:
(1)苗木引种在黑龙江大庆、帽儿山、哈尔滨等地进行了绒毛白蜡树种的引种栽培和区域性试验。经过几年的引种驯化,苗木的抗寒能力逐渐提高,在上述试验地点均能正常生长和越冬,苗木长势良好。与当地水曲柳相比,绒毛白蜡生长较快,冠幅较大,观赏性强。此外,在大庆、肇州试验点引种的苗木表现出较强的耐盐碱特点。
(2)种子育苗在黑龙江的哈尔滨、肇州、宾西和带岭等地进行种子育苗引种试验,种子的出苗率较高,苗木在各试验点生长良好,种子苗不用任何防寒措施就能正常越冬。与苗木直接引种相比,种子苗在萌发和发育过程中耐寒性逐渐加强,更容易适应新的环境。
(3)耐盐碱试验模拟大庆碱斑土盐分的离子种类和浓度,将5种中性盐和2种碱性盐按不同比例混合,对2年生绒毛白蜡引种苗木进行混合盐胁迫,测定株高、叶绿素含量、SOD、POD、MDA及脯氨酸等相关生理指标。初步研究表明,苗木能够在相当大庆碱斑土浓度的混合盐胁迫下继续生长,与其在光合、抗氧化酶活性、膜透性及渗透调节物质等方面的协调一致密切相关,表现出较强的耐盐碱性,与大庆、肇州等地的实际引种试验结果是一致的,初步说明绒毛白蜡在大庆等盐碱地区引种是可行的。
(4)嫁接技术以水曲柳为砧木嫁接绒毛白蜡亲和力强,采用嵌芽接和劈接均可以成功。相比之下,嵌芽接方法节省接穗,嫁接效率更高,嵌芽接成活率可达80%以上。嫁接苗木长势和越冬情况良好,抗寒能力得到很大提高,说明通过嫁接在黑龙江引种也是可行的。
(5)杂交育种开展了水曲柳与白蜡属的绒毛白蜡、小叶白蜡和美国白蜡不同种源及不同无性系间的属内种间杂交育种技术研究。
a.花粉技术在单因素试验和正交试验初步确定花粉离体萌发培养条件的基础上,采用基于BBD的响应面法优化了白蜡花粉离体萌发的培养基成分,使花粉萌发率有较大幅度提高,为杂交亲本的选择提供了快速有效的方法,同时提高了杂交育种的效率。将优化理论应用于植物花粉培养基研究为植物学的相关研究提供了新的思路。
b.杂交亲本生态差异显著,优缺点互补,杂种优势明显。通过白蜡属种间杂交共获得140个杂交组合的16361粒种子。不同组合的杂交种子性状变异丰富,为杂种F1优良基因型的选育及未来杂交新品种的培育奠定了基础。
c.采用适宜场强的高压静电场处理绒毛白蜡花粉,提高了花粉萌发率和杂交坐果率,克服了白蜡属内种间杂交不亲和性的难题,为林木种间杂交提供了有效方法。
d.采用石蜡制片法观察了水曲柳×绒毛白蜡杂种胚胎的发育过程,揭示了由原胚、球形胚、心形胚、鱼雷胚到子叶胚的发育过程,受精后胚乳先发育,由核型胚乳逐渐发育为细胞形胚乳,为白蜡属种间杂交育种提供了形态学依据。
e.通过杂种幼胚的离体培养克服了白蜡属种间杂种的胚败育,获得各杂交组合的组培苗,炼苗移栽提前获得了杂种植株。从幼胚培养到成苗仅用3个月的时间,与传统育苗方法相比,育种周期缩短了1年。
f.在杂种幼胚离体培养的基础上建立了杂种苗组培再生的技术平台,获得了杂种苗茎段腋芽增殖的培养基激素组合和生根条件,为优良杂种的无性繁育提供了技术保障。
In China the tree species resource of Fraxinus is abundant.Fraxinus velutina Torr.is one of the successful introduction of exotics.Due to its charactefistcs of good salt-resistance,good watchfulness and wide distributions,it has been introduced by many regions in China.It has also become the excellent tree species of foresting and greening by part of the coastal cities and saline and alkaline areas.Unfortunately F.velutina has not been introduced to Heilongjiang province yet.
In this thesis the F.velutina introduction was taken as the major clue.The related techniques,such as seedling introduction,seed seedling,experiments of salt resistance and the grafting techniques were systematically investigated,which were expected to provide research basis.In order to improve the quality of the tree species of Fraxinus and also to integrate the excellent gene resources,the interspecific crossbreedings for Fraxinus were mainly focused in this thesis.The main research contents are as follows:
Seedling Introduction The F.velutina introduction,cultivation and the regional experiments have been carried out in the areas of Daqing,Maoershan and Harbin of Heilongjiang province.The cold-resistance has been gradually enhanced after three years of introduction and domestication for F.velutina.The species can grow up and overwinter normally with good growth vigor.Compared to the local species of Fraxinus mandshurica Rupr,the introduced species exhibited faster growth,bigger crown width as well as better watchfullness.In addition,the more powerful saline-alkali tolerant capability was exposed for the seedling in Daqing and Zhaozhou experimental areas.
Seed planting Seed planting experiments were conducted in the areas of Harbin, Zhaozhou and Binxi of Heilongjiang province.The germination rates of the seeds were relatively high,and the seedling vagors were good in all experimental areas.The seedling can overwinter without any extra cold-proof measures.Compared to the introduced seedlings before,the cold-resistance capability for the seeds was gradually enhanced during the course of germinating and growth.It is easier to adapt to the new environment.So the seed introduction of F.velutina in Heilongjiang is more appropriate.
Experiments of salt resistance To simulate the ion,concentration and pH value of salinity of the alkaline soil in Daqing area,five neutral salts of NaCl,Na_2SO_4,MgCl_2,KCl, CaCl_2 and two alkaline salts of NaHCO_3 and Na_2CO_3 were blended into mixed-salts solutions. Three concentrations which were equivalent-to one time,two times and three times concentration of the alkaline soil were used to stress the introduced species of Fraxinus velutina Torr.A comparison was presented between the growth and the physiological indices of SPAS,SOD,POD,MDA as well as the Proline for the species.The analysis results showed that the species could survive under the stress with concentration by one time mainly due to the growth in harmony to the changes of the physiological indices.
Grafting techniques As the strong affinity of F.mandshurica,it could be successfully grafted with the scion of F.velutina by both methods of chip budding and cleft grafting.In comparison,the grafting efficiency was high for the chip budding method because it can save scions effectively and the survival rate of chip budding can reach above 80%.The seedling growth and the overwinter ability were so good that the cold-resistance can be enhanced greatly, which Verifies that it is feasible to introduce to Heilongjiang by grafting.
Crossbreeding techniques The interspecific crossbreeding between F.manchurica and Fraxinus velutina,,Fraxinus sogdiana and F.arnericana with different provenances and clones were studied in details as follows:
Based on the preliminary single-factor experiments and the orthogonal arrays the culture medium condition for pollen germinating in vitro has been determined,the culture medium ingredients of pollen germination in vitro for Fraxinus were optimized by employing the BBD(Box-Benkhen Design) based response surface methodology so that the pollen germination rate had been greatly enhanced,which provided effective method for crossing parents selection.In addition,this is as well a new way to apply the optimum theory into the study of plant pollen in the research fields of botany.
The crossing parents exhibited significant biological difference,and were complementary in advantages and disadvantages as well as the obvious heterosis.By interspecific crossing a total of 16361 seeds with 140 crossing combinations were obtained.This research will provide a basis for the cultivating of valuable hybrid F1 gene and future new crossing species.
The pollen of F.velutina is treated by the proper intensity of high-pressure electrostatic field,the high pollen germination rate and high setting percentage of crossing are enhanced, which overcome the difficulty of interspecific incompatibility of Fraxinus and provide effective way for interspecific crossing of forest.
Based on the conventional paraffin sectioning method,the development process of the hybrid embryos between F.mandshurian and F.velutina was observed at the optical level,which reveals the following processes:the development process of proembryo,globular embryo, heart-shape embryo,fish-torpedo-shape embryo and cotyledon embryo.The endosperm is developed firstly after fertilization.The nuclear endosperm is gradually developed into cellular shape endosperm in the course of the development.This work provides proof of morphology for the interspecific crossbreedings of Fraxinus.
The embryo abortion of interspecific crossbreeding of Fraxinus can be overcomed by in vitro culture of immature hybrid embryos.The tissue culture seedling for all crossing combinations can be obtained,and the hybrid plants can be obtained in early stage by acclimatization and transplants.It takes only 3 months from embryo culture to seedling formation.This method can reduce nearly one year comparing to the traditional breeding methods.
The technical platform of tissue culture and regeneration of hybrid are set up based on the culture of immature hybrid embryos,the hormone combinations and rooting conditions for culture medium for stem reproduction of hybrid canbe obtained.These can provide significant technical supports for the asexual breeding of valuable hybrids.
(1)苗木引种在黑龙江大庆、帽儿山、哈尔滨等地进行了绒毛白蜡树种的引种栽培和区域性试验。经过几年的引种驯化,苗木的抗寒能力逐渐提高,在上述试验地点均能正常生长和越冬,苗木长势良好。与当地水曲柳相比,绒毛白蜡生长较快,冠幅较大,观赏性强。此外,在大庆、肇州试验点引种的苗木表现出较强的耐盐碱特点。
(2)种子育苗在黑龙江的哈尔滨、肇州、宾西和带岭等地进行种子育苗引种试验,种子的出苗率较高,苗木在各试验点生长良好,种子苗不用任何防寒措施就能正常越冬。与苗木直接引种相比,种子苗在萌发和发育过程中耐寒性逐渐加强,更容易适应新的环境。
(3)耐盐碱试验模拟大庆碱斑土盐分的离子种类和浓度,将5种中性盐和2种碱性盐按不同比例混合,对2年生绒毛白蜡引种苗木进行混合盐胁迫,测定株高、叶绿素含量、SOD、POD、MDA及脯氨酸等相关生理指标。初步研究表明,苗木能够在相当大庆碱斑土浓度的混合盐胁迫下继续生长,与其在光合、抗氧化酶活性、膜透性及渗透调节物质等方面的协调一致密切相关,表现出较强的耐盐碱性,与大庆、肇州等地的实际引种试验结果是一致的,初步说明绒毛白蜡在大庆等盐碱地区引种是可行的。
(4)嫁接技术以水曲柳为砧木嫁接绒毛白蜡亲和力强,采用嵌芽接和劈接均可以成功。相比之下,嵌芽接方法节省接穗,嫁接效率更高,嵌芽接成活率可达80%以上。嫁接苗木长势和越冬情况良好,抗寒能力得到很大提高,说明通过嫁接在黑龙江引种也是可行的。
(5)杂交育种开展了水曲柳与白蜡属的绒毛白蜡、小叶白蜡和美国白蜡不同种源及不同无性系间的属内种间杂交育种技术研究。
a.花粉技术在单因素试验和正交试验初步确定花粉离体萌发培养条件的基础上,采用基于BBD的响应面法优化了白蜡花粉离体萌发的培养基成分,使花粉萌发率有较大幅度提高,为杂交亲本的选择提供了快速有效的方法,同时提高了杂交育种的效率。将优化理论应用于植物花粉培养基研究为植物学的相关研究提供了新的思路。
b.杂交亲本生态差异显著,优缺点互补,杂种优势明显。通过白蜡属种间杂交共获得140个杂交组合的16361粒种子。不同组合的杂交种子性状变异丰富,为杂种F1优良基因型的选育及未来杂交新品种的培育奠定了基础。
c.采用适宜场强的高压静电场处理绒毛白蜡花粉,提高了花粉萌发率和杂交坐果率,克服了白蜡属内种间杂交不亲和性的难题,为林木种间杂交提供了有效方法。
d.采用石蜡制片法观察了水曲柳×绒毛白蜡杂种胚胎的发育过程,揭示了由原胚、球形胚、心形胚、鱼雷胚到子叶胚的发育过程,受精后胚乳先发育,由核型胚乳逐渐发育为细胞形胚乳,为白蜡属种间杂交育种提供了形态学依据。
e.通过杂种幼胚的离体培养克服了白蜡属种间杂种的胚败育,获得各杂交组合的组培苗,炼苗移栽提前获得了杂种植株。从幼胚培养到成苗仅用3个月的时间,与传统育苗方法相比,育种周期缩短了1年。
f.在杂种幼胚离体培养的基础上建立了杂种苗组培再生的技术平台,获得了杂种苗茎段腋芽增殖的培养基激素组合和生根条件,为优良杂种的无性繁育提供了技术保障。
In China the tree species resource of Fraxinus is abundant.Fraxinus velutina Torr.is one of the successful introduction of exotics.Due to its charactefistcs of good salt-resistance,good watchfulness and wide distributions,it has been introduced by many regions in China.It has also become the excellent tree species of foresting and greening by part of the coastal cities and saline and alkaline areas.Unfortunately F.velutina has not been introduced to Heilongjiang province yet.
In this thesis the F.velutina introduction was taken as the major clue.The related techniques,such as seedling introduction,seed seedling,experiments of salt resistance and the grafting techniques were systematically investigated,which were expected to provide research basis.In order to improve the quality of the tree species of Fraxinus and also to integrate the excellent gene resources,the interspecific crossbreedings for Fraxinus were mainly focused in this thesis.The main research contents are as follows:
Seedling Introduction The F.velutina introduction,cultivation and the regional experiments have been carried out in the areas of Daqing,Maoershan and Harbin of Heilongjiang province.The cold-resistance has been gradually enhanced after three years of introduction and domestication for F.velutina.The species can grow up and overwinter normally with good growth vigor.Compared to the local species of Fraxinus mandshurica Rupr,the introduced species exhibited faster growth,bigger crown width as well as better watchfullness.In addition,the more powerful saline-alkali tolerant capability was exposed for the seedling in Daqing and Zhaozhou experimental areas.
Seed planting Seed planting experiments were conducted in the areas of Harbin, Zhaozhou and Binxi of Heilongjiang province.The germination rates of the seeds were relatively high,and the seedling vagors were good in all experimental areas.The seedling can overwinter without any extra cold-proof measures.Compared to the introduced seedlings before,the cold-resistance capability for the seeds was gradually enhanced during the course of germinating and growth.It is easier to adapt to the new environment.So the seed introduction of F.velutina in Heilongjiang is more appropriate.
Experiments of salt resistance To simulate the ion,concentration and pH value of salinity of the alkaline soil in Daqing area,five neutral salts of NaCl,Na_2SO_4,MgCl_2,KCl, CaCl_2 and two alkaline salts of NaHCO_3 and Na_2CO_3 were blended into mixed-salts solutions. Three concentrations which were equivalent-to one time,two times and three times concentration of the alkaline soil were used to stress the introduced species of Fraxinus velutina Torr.A comparison was presented between the growth and the physiological indices of SPAS,SOD,POD,MDA as well as the Proline for the species.The analysis results showed that the species could survive under the stress with concentration by one time mainly due to the growth in harmony to the changes of the physiological indices.
Grafting techniques As the strong affinity of F.mandshurica,it could be successfully grafted with the scion of F.velutina by both methods of chip budding and cleft grafting.In comparison,the grafting efficiency was high for the chip budding method because it can save scions effectively and the survival rate of chip budding can reach above 80%.The seedling growth and the overwinter ability were so good that the cold-resistance can be enhanced greatly, which Verifies that it is feasible to introduce to Heilongjiang by grafting.
Crossbreeding techniques The interspecific crossbreeding between F.manchurica and Fraxinus velutina,,Fraxinus sogdiana and F.arnericana with different provenances and clones were studied in details as follows:
Based on the preliminary single-factor experiments and the orthogonal arrays the culture medium condition for pollen germinating in vitro has been determined,the culture medium ingredients of pollen germination in vitro for Fraxinus were optimized by employing the BBD(Box-Benkhen Design) based response surface methodology so that the pollen germination rate had been greatly enhanced,which provided effective method for crossing parents selection.In addition,this is as well a new way to apply the optimum theory into the study of plant pollen in the research fields of botany.
The crossing parents exhibited significant biological difference,and were complementary in advantages and disadvantages as well as the obvious heterosis.By interspecific crossing a total of 16361 seeds with 140 crossing combinations were obtained.This research will provide a basis for the cultivating of valuable hybrid F1 gene and future new crossing species.
The pollen of F.velutina is treated by the proper intensity of high-pressure electrostatic field,the high pollen germination rate and high setting percentage of crossing are enhanced, which overcome the difficulty of interspecific incompatibility of Fraxinus and provide effective way for interspecific crossing of forest.
Based on the conventional paraffin sectioning method,the development process of the hybrid embryos between F.mandshurian and F.velutina was observed at the optical level,which reveals the following processes:the development process of proembryo,globular embryo, heart-shape embryo,fish-torpedo-shape embryo and cotyledon embryo.The endosperm is developed firstly after fertilization.The nuclear endosperm is gradually developed into cellular shape endosperm in the course of the development.This work provides proof of morphology for the interspecific crossbreedings of Fraxinus.
The embryo abortion of interspecific crossbreeding of Fraxinus can be overcomed by in vitro culture of immature hybrid embryos.The tissue culture seedling for all crossing combinations can be obtained,and the hybrid plants can be obtained in early stage by acclimatization and transplants.It takes only 3 months from embryo culture to seedling formation.This method can reduce nearly one year comparing to the traditional breeding methods.
The technical platform of tissue culture and regeneration of hybrid are set up based on the culture of immature hybrid embryos,the hormone combinations and rooting conditions for culture medium for stem reproduction of hybrid canbe obtained.These can provide significant technical supports for the asexual breeding of valuable hybrids.
引文
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