摘要
百合是百合属(Lilium)植物总称,近年来已成为我国最受欢迎和销量最高的切花之一。百合喜冷凉湿润气候,当温度高于28℃,就会严重影响切花质量并引起百合种球退化,因此百合越夏成为我国南方和北方地区生产优质鲜切花的难关之一。培育耐高温的百合品种,是解决夏季百合生产困难的主要途径。我国拥有丰富的百合属资源,但在国际新品种登录的5000多个百合品种中,来源于我国的仅有8种,因而充分利用自有资源对于培育适合我国的耐热百合品种具有十分重要的意义。因此本研究从体细胞无性系变异规律入手,选取了耐热性较好的原生种渥丹(Lilium concolor)、岷江百合(L.regale)和湖北百合(L.henryi),从形态学、细胞学和分子生物学三方面进行了较为系统的研究,并在此基础上对岷江百合开展了体细胞无性系耐热选育工作。主要结果如下:
(1)在三种百合的组织培养过程中,基因型在体细胞变异频率上起着决定性作用:渥丹的分子标记多态性变异率最高,为11.44%;岷江百合次之,为5.65%;湖北百合最低,为1.10%。在外植体的选择上,体细胞变异频率由高到低依次为叶片>鳞片>种子。分子标记结果表明,渥丹叶片外植体再生植株的变异率为46.48%,高于鳞片(32.74%)和种子(11.44%)外植体的变异率。岷江百合的叶片外植体再生植株的变异率29.70%,高于鳞片(21.20%)和种子(5.65%)外植体的变异率。湖北百合叶片外植体再生植株的变异率在三种百合种最低,为20.73%,高于鳞片(1.10%)和种子(14.11%)外植体的变异率。不同激素对遗传变异率影响不同。在渥丹、岷江百合的组织培养过程中,NAA、6-BA,IBA的添加对遗传稳定性影响不显著;在添加了2,4-D或TDZ后(M3～M6培养基),渥丹的分子标记多态性变异频率和对照相比增加了0.74～1.09倍,岷江百合则增加了0.28～0.42倍;湖北百合和对照相比无显著差异。在三种百合中,愈伤组织为再生途径得到无性系的变异频率均高于直接以不定芽途径得到的无性系变异频率。渥丹、岷江百合和湖北百合的愈伤组织再生途径的分子标记多态性变异率为42.10%、11.98%和8.56%,分别为不定芽再生途径变异率的3.68倍、2.12和7.79倍。继代时间对三种百合的体细胞变异率影响不同。根据分子检测结果,渥丹在继代5次、15次时,变异率分别为36.71%、10.28%;岷江百合在继代8次、15次时,分别为15.22%、5.41%:湖北百合在继代15次时,遗传变异率增加至4.88%。
在以保存种质资源为前提下,宜用种子作为外植体,通过不定芽直接再生途径,在含有NAA、6-BA或IBA的培养基中进行培养。渥丹、岷江百合以大于1cm的小鳞茎,湖北百合以小于1cm的鳞茎为宜。在突变体育种中,宜用叶片作为外植体,通过愈伤组织再生途径,在含有2,4-D或TDZ的培养基中进行培养。渥丹、岷江百合的小鳞茎在小于1cm,湖北百合大于1cm时进行突变育种为宜。
(2)随着温度的升高和处理时间的延长,移栽的岷江百合离体幼苗耐热指数不断下降,叶绿素含量减少,丙二醛(MDA)含量增加;42℃胁迫32 h后,游离脯氨酸为16.84 mg/g、可溶性蛋白含量为0.61 mg/g、SOD活性为5457.86 U/g,均达到最大值,比对照增加了0.66倍、1.77倍和9.78倍。但42℃高温胁迫48 h后,指标显著下降,这说明,42℃间断胁迫48 h对植株产生了不可逆的的热伤害作用。
(3)采用了高温胁迫(高温一次、多次筛选),化学胁迫(HYP一次、多次筛选)和复合胁迫(HYP+高温)5种筛选方法对岷江百合无性系进行筛选。通过对所筛选得到的幼苗进行鉴定和分析可知,高温筛选的植株在高温测试中存活率为26%,高于HYP筛选(17%)和复合筛选(23%)的存活率。在耐热性能上,高温筛选的耐热指数为0.17,高于HYP筛选(0.055)和复合筛选(0.07)的耐热指数。在高温测试中,存活的筛选植株SOD活性大幅度提高,游离脯氨酸和可溶性蛋白含量也均高于正常移栽苗。本研究从近5000棵扩繁的植株中,筛选出30棵耐热性能优良的突变体。综上,从实验过程的复杂度和效率来讲,起始材料宜通过不定芽直接再生途径来进行选择筛选,筛选方法宜用37℃或40℃高温多次筛选方法,这样可以较多的筛选到真正耐高温的岷江百合幼苗。
本研究为百合种质资源保存提供了理论依据,为耐热育种奠定了基础。
Lily,belonging to the genus Lilium,is becoming one of the most popular cut flowers.Lily likes cold humid climate,when the temperature is higher than 28℃,it will seriously affect the quality of cut flowers and cause the degradation of lily bulbs.So,lily production over summer in southern and northern areas has become one of the problems of producing perfect cut flowers.Given this,breeding heat-resistant lily varieties is the key factor to solve the problem.There has been only 8 cultivars from China among more than 5000 cultivars in International Registry of New Varieties,although our country is blessed with lily resources.Therefore,It is of profound significance to make full use of own resources for breeding heat-resistant cultivars grown well in China.So in this paper,firstly the law of somaclonal variation of Lilium concolor,L.regale and L.henryi were investigated systematically from three ways, including Morphology,cytology and molecular biology.Secondly,breeding heat-resistant cultivars for Lilium regale was carried out.The main results are as follows:
(1) In the process of lily tissue culture,the genotype plays a decisive role in somaclonal variation rate:L.concolor is the highest(11.44%),L.regale higher(5.65%) and L.henryi the lowest(1.10%) according to the results of ISSR molecular marker.In the matter of explant selection,the somaclonal variation rate is as follows:leaf>scale>seed.Somaclonal variation rate determined by ISSR molecular marker of clones regenerated from leaf in L.concolor was 46.48%higher than that of clones regenerated from scales(32.74%) and seed(11.44%).In L.regale somaclonal variation rate of clones regenerated from leaf was 29.70%higher than that of clones regenerated from scales(21.20%) and seed(5.65%). Somaclonal variation rate of clones regenerated from leaf in L.henryi was 20.73%,higher than that of clones regenerated from scales(1.10%) and seed(14.11%).Different exogenous hormone had different effects on lilies.Addition of NAA,6-BA and IBA had no effects on somaclonal variation rate. But 2,4-D and TDZ can reduce genetic stability of L.concolor which increased 0.74～1.09 times in somaclonal variation and L.regale which increased 0.28~0.42 times respectively.There were less significant effets on L.henryi.Somaclonal variation rate of regenerated plants from callus of L.concolor, L.regale and L.henryi was 42.10%,11.98%and 8.56%which was higher than that of regenerated plants from adventitious buds.The somaclonal variation rate of L.concolor after 5 generation times(36.71%) as well as L.regale after 8 generation times(15.22%) reaches the peak respectively,while L.henryi was increasing slowly in the 2～8 subcultures and rose sharply to 4.88%after 15 subcultures.Therefore, when generic resources preservation required to maintain exactly generic information is the purpose,it is better to select seed as the explant,NAA,6-BA or IBA as the exogenous hormone,adventitious buds as the regeneration way,and bulbs diameter of Lconcolor and L.regale is more than while L.henryi less than 1 cm as the state of preservation.When mutation breeding required more mutants is the objective,it is better to select leaf as the explant,2,4-D or TDZ as the exogenous hormone,callus as the regeneration way,and bulbs diameter is less than while L.henryi more than 1 cm as the state of preservation.
(2) Transplanted seedlings of L.regale were treated under high temperature.The results showed that heat-resistance index and chlorophyll content in leaf were decreased significantly under high temperature stress,while malondialdehyde content increased.SOD activity(5457.86 U/g),free proline content(16.84 mg/g) and soluble protein content(0.61 mg/g) in leaf of seedlings treated for 32 hours under 42℃reached the peak,with increasing 0.66,1.77 and 9.78 times respectively compared the control but 48 hours later,they declined.The results indicated that discontinuous stress under 42℃for 48 h had irreversible heat injury effects on seedlings.
(3) The study applied 5 screening methods on clones of L.regale,including high temperature stress (one-step or multiple -step screening),chemical stress(one-step or multiple-step of HYP) and compounded stress(HYP with high temperature).Identification and analysis on seedlings screened by 5 above-mentioned methods were carried out.The survival rate of seedlings screened by high temperature stress was 26%,which was higher than that of HYP stress(17%) and compounded stress(23%).The heat-resistant index of seedlings screened by high temperature stress was 0.17,which was higher than that of HYP stress(0.055) and compounded stress(0.07).During high temperature environment,the SOD activities of screened seedlings increased to a large extent with free proline and soluble protein content higher than the control.All in all,30 mutants were obtained from 5000 in vitro plants.Thus, with the respect of difficulty and efficiency in experiment,it is better to choose adventitious buds as regeneration ways,multiple-step high temperature under 37℃or 40℃as the screening methods,so that more genus mutants of L.regale can be obtained.
This paper provides theoretical basis for genetic resources preservation of lilies as well as lays a foundation for mutant breeding.
引文
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