试管苗远缘嫁接技术研究初探及嫁接苗的ISSR分析
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摘要
目前,嫁接已被广泛应用于农业、林业以及园艺植物生产中。自20世纪80年代,孟昭璜把豆科的绿豆(Phaseolus radiatus Linn.)嫁接在旋花科甘薯(Dioscorea esculenta(Lour.)Burkill)上面,得到了嫁接杂种种子,经长期的选择和培育最终获得了性状稳定遗传的高产优质的薯绿豆,并创造了极大的价值之后,利用远缘嫁接进行植物品种改良的方法开始受到人们的青睐,成为了育种研究工作的热点之一。以组织培养为基础的试管嫁接由于嫁接后在人工控制的环境条件下对嫁接植株进行培养,排除了室外环境因素对嫁接成活的影响,可以方便的改变条件以研究各种因素对嫁接的影响。因此试管内嫁接技术也成为生产和各种基础理论研究的重要工具。
本论文以花卉植物为主要研究对象,嫁接组合涉及11个科,包括草本与木本、单子叶植物与双子叶植物、科与科等,进行了试管内远缘嫁接的研究,这对远缘嫁接系统研究具有重要的理论和实践意义。其主要研究结果如下:
不同嫁接组合研究结果表明:在所研究的组合当中,成活率达到30%及以上的嫁接组合:鸡冠花(Celosia cristata L.)/红掌(Anthurium andraeanum)(36.7%),鸡冠花/大岩桐(Sinningiaspeciosa Benth.et Hook)(33.3%),长寿花(Kalanchoe blossfeldiana V.Poelln.)/红掌(30.0%),大岩桐/鸡冠花(30.0%);成活率在20%～30%之间的嫁接组合:长寿花/大岩桐(26.7%),长寿花/鸡冠花(23.4%),大岩桐/长寿花(20.0%),大岩桐/红掌(20.0%);成活率为10%～20%的组合:鸡冠花/长寿花(16.7%),鸡冠花/景天(Crassulaceae Sedum Spectabile L.)(16.7%),百合(Lilium brownii F.E.Br.var.viridulum Baker)/长寿花(15.0%),月见草(Oenothera erythrosepalaBorb.)/大岩桐(13.3%),长寿花/景天(13.3%),马蹄莲(Zantedeschia eathiopica)/鸡冠花(10%),长寿花/宝塔菜(Stachys arrecta L.H.Bailey)(10.0%);成活率在10%以下的组合:马蹄莲/大岩桐(6.7%),大岩桐/宝塔菜(6.7%),长春花(Catharanthus roseus(L.)G.Don)/鸡冠花(6.7%),百合/鸡冠花(5.0%),月见草/鸡冠花(3.3%)。其余各嫁接组合均未得到嫁接成活苗。
其中草本与木本嫁接组合未得到嫁接成活苗;单子叶植物与双子叶植物、草本植物科间嫁接的的成活率分别为10.9%和12.9%。
嫁接方法对嫁接成活率有一定的影响。对于长寿花/鸡冠花组合来说,用插接法进行嫁接时成活率最高,达到27.5%。且与其他两种嫁接方法相比差异显著。
砧穗苗龄是影响嫁接成功的关键因素之一,应该依据材料的不同选择适合苗龄的植株。用鸡冠花组培苗和实生苗嫁接大岩桐均以10 d苗龄的鸡冠花做接穗最为适宜,成活率分别达到30%和35%;鸡冠花组培苗和实生苗作砧木时,分别以20 d和25 d苗龄左右的嫁接成活率最高,分别达到27.5%和32.5%,其砧穗生长也较好;砧木和接穗用组培苗和用实生苗成活率差异不大。
接穗茎尖切取大小对嫁接成功十分重要。对于长寿花/大岩桐嫁接组合,当接穗大小为6mm左右时,嫁接成活率达到最高,达到30%,切取的茎尖小于3mm或大于10mm都不利于试管嫁接的成活。
砧木留叶有利于试管微嫁接的成功。本实验中月见草/大岩桐组合的嫁接,当砧木留有一定的叶片时的嫁接成活率为22.5%,远远高于切去所有叶片时的成活率7.5%。
本试验还首次尝试以红掌的愈伤组织作为砧木嫁接鸡冠花等接穗,结果表明,其成活率远高于用组培苗作砧木的嫁接。鸡冠花接在红掌愈伤组织上的嫁接成活率为27.5%,和接在组培苗上面相比,差异显著。
用ISSR分析检测到科间组合鸡冠花/大岩桐、大岩桐/鸡冠花、长寿花/鸡冠花、长寿花/大岩桐、鸡冠花/景天及单双子叶间组合长寿花/红掌、鸡冠花/红掌等儿种嫁接组合均在多种引物下出现了有差异的位点。鸡冠花/大岩桐组合与大岩桐/鸡冠花组合嫁接后接穗甚至出现了来自于砧木的特异带纹,对其增值后代重新检测发现此带纹仍然存在。这就支持了传统的砧木与接穗之间有可能是发生了DNA等大分子物质的转移,产生了基因交流的观点。但目前尚无分子生物学方面的直接证据证明此推测,实验结果所示的位点新增与缺失等现象,或许也与远缘嫁接逆境引起转座子活动而导致基因突变有关。
Grafting has beening widely applied to agriculture,forestry and horticulture,and micrografting based on tissue culture are used as an important tool for production and science research.It is being regarded as important that to improve plants' traits by distant grafting since the breeding specialist, Meng-Zhaohuang,succeeded in grafting mung bean onto sweet potato,and obtained a prolific,high quality new mung bean variety,which created great value.Further studies on the application of the distant grafting technique may contribute to a new way for plant breeding.The grafting plant was cultivated in manual manipulation condition,which excluded the effect of environmental agents,and makes the study much more convenient,therefore,micrografting is becoming an important tool in production and primary theory reaserch.
Flowering plants involved in 11 families included grafting combinations of herb and wood, dicotyledon and monocotyl,different families,were the main materials in this study.By the method of micrografting,a serious experiments were carried out in an attempt to approach the influence on grafting of several factors.The main results showed as follow.
The study on different grafting combinations indicated that combinations whose survival rates achieved 30%and above included Celosia cristata/Anthurium andraeanum(36.7%),Celosia cristata/Sinningia speciosa(33.3%),Kalanchoe blossfeldiana/Anthurium andraeanum(30.0%), Sinningia speciosa/Celosia cristata(30.0%);the combinations whose survival rates were 20%～30% included Kalanchoe blossfeldiana/Sinningia speciosa(26.7%),Kalanchoe blossfeldiana/Celosia cristata(23.4%),Sinningia speciosa/Kalanchoe blossfeldiana(20.0%),Sinningia speciosa/Anthurium andraeanum(20.0%);the combinations whose suvival rates were 10%～20%included Celosia cristata/Kalanchoe blossfeldiana(16.7%),Celosia cristata/Crassulaceae Sedum(16.7%),Lilium brownii/Kalanchoe blossfeldiana(15.0%),Oenothera erythrosepala/Sinningia speciosa(13.3%), Kalanchoe blossfeldiana/Crassulaceae Sedum(13.3%),Zantedeschia eathiopica/Celosia cristata (10.0%),Kalanchoe blossfeldiana/Stachys arrecta(10.0%);the combinations whose suevival rates were below 10%included Zantedeschia eathiopica/Sinningia speciosa(6.7%),Sinningia speciosa/ Stachys arrecta(6.7%),Catharanthus roseus/Celosia cristata(6.7%),Lilium brownii/Celosia cristata(5.0%),Oenothera erythrosepala/Celosia cristata(3.3%);and none survival grafting plants was obtained in the other combinations.
There was no survival grafting plant came from conbinations of herb and wood.The survival rate of combinations of dicotyledon and monocotyl,family and family,were respectively 10.9%and 13.9%.
The method of grafting has a certain influence on grafting survival rate.To Kalanchoe blossfeldiana/Celosia cristata,by method C,the rate was 27.5%,which is the top of the three.And it made significant difference contrasted with the other two methods.
Age of the stock and scion is one of the crucial factors for a successful grafting.Source of scion and stock in proper age should be choosed according to different plant.Celosia cristata L.cultured in vitro and seeding plant of 10 d grafted onto Sinningia speciosa were the most suitable,and the survival rate were respectively 30%and 35%;as stocks,Celosia cristata L.cultured in vitro of 20ds and seeding plant of 25 d were better to be grafted in Kalanchoe blossfeldiana,and their survival rate were 27.5%and 32.5%.And there was little difference between tissue culture plant and seeding plant.
Size of stem apex cut as scion is also an important factor for a successful grafting.To Kalanchoe blossfeldiana/Sinningia speciosa,stem apex of 6 mm got the highest survival rate,30.%. Larger or smaller size both do harm to micrografting.
To remain some stock leaves benefits to micrografting.In this experiment,the survival rate of Oenothera erythrosepala/Sinningia speciosa reached 22.5%when using stocks with leaves,which was far higher than using stocks without leaves.
In this experiment,scions were grafted in Anthurium andraeanum callus first time.Celosia cristata was grafted in Anthurium andraeanum callus,the survival rate was 27.5%,and the result made it clear that it was much higher than using tissue culture plants as stocks.
ISSR analysis indicated that significant difference between the variation and the source of scion as well as stock was recognized in 7 combinations.2 of them may support the possibility that gene's transformation from stock to scion as traditional studies presumed for that a few PCR fragments amplified from the variety corresponded to those amplified from the stock.The genetic variation seems not to be caused simply by DNA transformation,since there is so far no cellular or molecular evidence revealing the possibility or exact mechanism of long distance movement of DNA.And transposition in response to environmental stress may also contribute to distant grafting-induced genetic diversity.
本论文以花卉植物为主要研究对象,嫁接组合涉及11个科,包括草本与木本、单子叶植物与双子叶植物、科与科等,进行了试管内远缘嫁接的研究,这对远缘嫁接系统研究具有重要的理论和实践意义。其主要研究结果如下:
不同嫁接组合研究结果表明:在所研究的组合当中,成活率达到30%及以上的嫁接组合:鸡冠花(Celosia cristata L.)/红掌(Anthurium andraeanum)(36.7%),鸡冠花/大岩桐(Sinningiaspeciosa Benth.et Hook)(33.3%),长寿花(Kalanchoe blossfeldiana V.Poelln.)/红掌(30.0%),大岩桐/鸡冠花(30.0%);成活率在20%～30%之间的嫁接组合:长寿花/大岩桐(26.7%),长寿花/鸡冠花(23.4%),大岩桐/长寿花(20.0%),大岩桐/红掌(20.0%);成活率为10%～20%的组合:鸡冠花/长寿花(16.7%),鸡冠花/景天(Crassulaceae Sedum Spectabile L.)(16.7%),百合(Lilium brownii F.E.Br.var.viridulum Baker)/长寿花(15.0%),月见草(Oenothera erythrosepalaBorb.)/大岩桐(13.3%),长寿花/景天(13.3%),马蹄莲(Zantedeschia eathiopica)/鸡冠花(10%),长寿花/宝塔菜(Stachys arrecta L.H.Bailey)(10.0%);成活率在10%以下的组合:马蹄莲/大岩桐(6.7%),大岩桐/宝塔菜(6.7%),长春花(Catharanthus roseus(L.)G.Don)/鸡冠花(6.7%),百合/鸡冠花(5.0%),月见草/鸡冠花(3.3%)。其余各嫁接组合均未得到嫁接成活苗。
其中草本与木本嫁接组合未得到嫁接成活苗;单子叶植物与双子叶植物、草本植物科间嫁接的的成活率分别为10.9%和12.9%。
嫁接方法对嫁接成活率有一定的影响。对于长寿花/鸡冠花组合来说,用插接法进行嫁接时成活率最高,达到27.5%。且与其他两种嫁接方法相比差异显著。
砧穗苗龄是影响嫁接成功的关键因素之一,应该依据材料的不同选择适合苗龄的植株。用鸡冠花组培苗和实生苗嫁接大岩桐均以10 d苗龄的鸡冠花做接穗最为适宜,成活率分别达到30%和35%;鸡冠花组培苗和实生苗作砧木时,分别以20 d和25 d苗龄左右的嫁接成活率最高,分别达到27.5%和32.5%,其砧穗生长也较好;砧木和接穗用组培苗和用实生苗成活率差异不大。
接穗茎尖切取大小对嫁接成功十分重要。对于长寿花/大岩桐嫁接组合,当接穗大小为6mm左右时,嫁接成活率达到最高,达到30%,切取的茎尖小于3mm或大于10mm都不利于试管嫁接的成活。
砧木留叶有利于试管微嫁接的成功。本实验中月见草/大岩桐组合的嫁接,当砧木留有一定的叶片时的嫁接成活率为22.5%,远远高于切去所有叶片时的成活率7.5%。
本试验还首次尝试以红掌的愈伤组织作为砧木嫁接鸡冠花等接穗,结果表明,其成活率远高于用组培苗作砧木的嫁接。鸡冠花接在红掌愈伤组织上的嫁接成活率为27.5%,和接在组培苗上面相比,差异显著。
用ISSR分析检测到科间组合鸡冠花/大岩桐、大岩桐/鸡冠花、长寿花/鸡冠花、长寿花/大岩桐、鸡冠花/景天及单双子叶间组合长寿花/红掌、鸡冠花/红掌等儿种嫁接组合均在多种引物下出现了有差异的位点。鸡冠花/大岩桐组合与大岩桐/鸡冠花组合嫁接后接穗甚至出现了来自于砧木的特异带纹,对其增值后代重新检测发现此带纹仍然存在。这就支持了传统的砧木与接穗之间有可能是发生了DNA等大分子物质的转移,产生了基因交流的观点。但目前尚无分子生物学方面的直接证据证明此推测,实验结果所示的位点新增与缺失等现象,或许也与远缘嫁接逆境引起转座子活动而导致基因突变有关。
Grafting has beening widely applied to agriculture,forestry and horticulture,and micrografting based on tissue culture are used as an important tool for production and science research.It is being regarded as important that to improve plants' traits by distant grafting since the breeding specialist, Meng-Zhaohuang,succeeded in grafting mung bean onto sweet potato,and obtained a prolific,high quality new mung bean variety,which created great value.Further studies on the application of the distant grafting technique may contribute to a new way for plant breeding.The grafting plant was cultivated in manual manipulation condition,which excluded the effect of environmental agents,and makes the study much more convenient,therefore,micrografting is becoming an important tool in production and primary theory reaserch.
Flowering plants involved in 11 families included grafting combinations of herb and wood, dicotyledon and monocotyl,different families,were the main materials in this study.By the method of micrografting,a serious experiments were carried out in an attempt to approach the influence on grafting of several factors.The main results showed as follow.
The study on different grafting combinations indicated that combinations whose survival rates achieved 30%and above included Celosia cristata/Anthurium andraeanum(36.7%),Celosia cristata/Sinningia speciosa(33.3%),Kalanchoe blossfeldiana/Anthurium andraeanum(30.0%), Sinningia speciosa/Celosia cristata(30.0%);the combinations whose survival rates were 20%～30% included Kalanchoe blossfeldiana/Sinningia speciosa(26.7%),Kalanchoe blossfeldiana/Celosia cristata(23.4%),Sinningia speciosa/Kalanchoe blossfeldiana(20.0%),Sinningia speciosa/Anthurium andraeanum(20.0%);the combinations whose suvival rates were 10%～20%included Celosia cristata/Kalanchoe blossfeldiana(16.7%),Celosia cristata/Crassulaceae Sedum(16.7%),Lilium brownii/Kalanchoe blossfeldiana(15.0%),Oenothera erythrosepala/Sinningia speciosa(13.3%), Kalanchoe blossfeldiana/Crassulaceae Sedum(13.3%),Zantedeschia eathiopica/Celosia cristata (10.0%),Kalanchoe blossfeldiana/Stachys arrecta(10.0%);the combinations whose suevival rates were below 10%included Zantedeschia eathiopica/Sinningia speciosa(6.7%),Sinningia speciosa/ Stachys arrecta(6.7%),Catharanthus roseus/Celosia cristata(6.7%),Lilium brownii/Celosia cristata(5.0%),Oenothera erythrosepala/Celosia cristata(3.3%);and none survival grafting plants was obtained in the other combinations.
There was no survival grafting plant came from conbinations of herb and wood.The survival rate of combinations of dicotyledon and monocotyl,family and family,were respectively 10.9%and 13.9%.
The method of grafting has a certain influence on grafting survival rate.To Kalanchoe blossfeldiana/Celosia cristata,by method C,the rate was 27.5%,which is the top of the three.And it made significant difference contrasted with the other two methods.
Age of the stock and scion is one of the crucial factors for a successful grafting.Source of scion and stock in proper age should be choosed according to different plant.Celosia cristata L.cultured in vitro and seeding plant of 10 d grafted onto Sinningia speciosa were the most suitable,and the survival rate were respectively 30%and 35%;as stocks,Celosia cristata L.cultured in vitro of 20ds and seeding plant of 25 d were better to be grafted in Kalanchoe blossfeldiana,and their survival rate were 27.5%and 32.5%.And there was little difference between tissue culture plant and seeding plant.
Size of stem apex cut as scion is also an important factor for a successful grafting.To Kalanchoe blossfeldiana/Sinningia speciosa,stem apex of 6 mm got the highest survival rate,30.%. Larger or smaller size both do harm to micrografting.
To remain some stock leaves benefits to micrografting.In this experiment,the survival rate of Oenothera erythrosepala/Sinningia speciosa reached 22.5%when using stocks with leaves,which was far higher than using stocks without leaves.
In this experiment,scions were grafted in Anthurium andraeanum callus first time.Celosia cristata was grafted in Anthurium andraeanum callus,the survival rate was 27.5%,and the result made it clear that it was much higher than using tissue culture plants as stocks.
ISSR analysis indicated that significant difference between the variation and the source of scion as well as stock was recognized in 7 combinations.2 of them may support the possibility that gene's transformation from stock to scion as traditional studies presumed for that a few PCR fragments amplified from the variety corresponded to those amplified from the stock.The genetic variation seems not to be caused simply by DNA transformation,since there is so far no cellular or molecular evidence revealing the possibility or exact mechanism of long distance movement of DNA.And transposition in response to environmental stress may also contribute to distant grafting-induced genetic diversity.
引文
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