魔芋离体形态发生机制及其繁殖技术
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摘要
魔芋(Konjac)是植物界中迄今发现唯一能大量合成葡苷聚糖(Konjac glucomannan)的高等植物，属天南星科(Araceae)魔芋属(Amrophophallus Blume)，为多年生草本植物，广泛分布于我国西南山区，具有悠久的栽培和利用历史。魔芋葡甘聚糖是魔芋地下球茎主要贮藏物质，为一种高分子量碳水化合物，因具有良好的胶溶性、凝胶性、增稠性、成膜性及与其它植物胶的优良复配性等优点而在食品、化工、医药保健、环保等领域具有广泛的应用。因此，魔芋具有极高的市场开发和利用价值。
魔芋为单叶柄支撑的单叶植物，通常利用球茎着生的根状茎进行无性繁殖，其繁殖系数极低，且多年沿用无性繁殖造成病原菌的积累、种性的退化，进而导致魔芋产量下降、品质劣化。由于缺乏稳定的良种繁育体系，种芋的数量和质量已经成为魔芋生产发展的瓶颈。自上个世纪80年代以来，许多研究者试图利用组织培养技术生产脱毒试管苗，以繁殖种芋，但试管苗因移栽成活率低且对田间管理技术要求较高，不能实现周年供应和长距离运输等缺点，无法在实际生产中广泛运用。二十世纪90年代以来，随着对植物变态器官发育机理认识的不断深入，人们在试管中已成功诱导出了块茎、球茎、鳞茎等营养繁殖器官，其中马铃薯试管块茎生产体系最为成熟，已经实现了工厂化生产，但有关魔芋试管球茎的研究国内外尚未见报道。本研究以花魔芋(A．rivieri)和白魔芋(A．albus)为材料，对魔芋试管球茎的形成及魔芋组织培养繁殖方式开展了系列研究，主要结果如下：
1．幼嫩叶柄是魔芋组织培养较好的外植体，愈伤组织形成率高，状态好。MS+1.0mg／L NAA+1.0mg／L BA适宜于愈伤组织诱导，而MS+0.5 mg／L NAA+0.5 mg／L BA有利于愈伤组织增殖。愈伤组织在继代培养过程中可形成3种不同的类型，即Ⅰ：水渍状，半透明；Ⅱ：淡黄色，结构疏松；Ⅲ：浅红或绿色，呈瘤状或球状，结构致密。在分化培养基上，Ⅰ型愈伤组织不能分化出芽，Ⅱ型可分化出不定芽，而Ⅲ型可通过拟球茎再生完整植株。这三种类型愈伤组织在特定的培养条件下可相互转化。组织学观察显示，三种类型的愈伤组织其组成细胞类型明显不同，其中Ⅲ型愈伤组织由保守分裂型细胞组成，其细胞中含有淀粉和葡甘聚糖颗粒，是一种半组织化结构。中等浓度(1.0～2.0 mg／L)的细胞分裂素(BA、KT、ZT和TDZ)与低浓度(0.1～1.0mg／L)的NAA配合均能促使Ⅲ型愈伤组织再生植株。
2．魔芋离体形态建成以器官发生途径为主，拟球茎和芽发生是主要形式。组织学观察显示，魔芋叶柄无维管组织形成层，维管束外层薄壁在培养过程首先启动脱分化，进而形成愈伤组织。魔芋形态建成以器官发生途径为主，在特定培养条件下也可形成体细胞胚，但频率极低，且绝大部分体胚因高度畸形化而不能形成再生植株。魔芋可通过不定芽器官发生途径或拟球茎器官发生途径形成植株，通过拟球茎形成的植株不需生根培养就具有完整的根系，而不定芽在发育过程中不能直接生根且植株生长势弱。不定芽和拟球茎均起源于Ⅲ型愈伤组织的次表皮细胞，这类细胞在分化过程中首先形成拟分生组织团。拟分生组织团在分化培养中既可直接形成芽原基，进而发生不定芽，可形成一种中间的球状体，这种球状体具顶芽分生组织，叶原基不明显，它可进一步发育成拟球茎，进而形成完整植株。
3．内源GA_3含量增加伴随着不定芽发生，而JA含量增加伴随着拟球茎发生，内源激素平衡是器官发生方式的关键调空因素。魔芋叶柄薄壁细胞在脱分化形成愈伤组织过程中，其内源激素(IAA、GA_3、ABA和JA)含量发生了明显的变化。内源IAA含量培养初期缓慢下降，10d后快速上升，30d后达到平稳。内源ABA含量在培养20d内呈下降趋势，之后迅速上升。内源GA_3和JA含量则在整个愈伤组织诱导过程中迅速下降。研究发现，三种内源激素(GA_3、ABA和JA)在形成芽和拟球茎的愈伤组织中含量变化明显不同。内源GA_3水平在拟球茎发生过程中变动不大，而在不定芽发生过程中快速上升，其绝对含量也比前者高；内源ABA在拟球茎发生过程中基本维持稳定，而在不定芽发生过程中则迅速下降；内源JA含量在拟球茎形成过程中呈明显上升趋势，而在不定芽发生过程中变化不明显。只有内源IAA无论是拟球茎还是不定芽形成过程其含量均呈下降趋势。内源GA_3与ABA和JA的平衡水平在两种不同器官发生途径中呈现出明显的差异，但GA_3／JA差异最为显著，在两种器官发生途径中其变化趋势完全相反。上述结果说明，魔芋离体器官发生形式与内源激素变化及平衡水平相关。
4．魔芋试管球茎(即拟球茎)的形成受培养基和培养条件的影响。中等浓度的BA(1.0～2.0 mg／L)与低浓度的NAA(0.1～0.5 mg／L)配合有利于Ⅲ型愈伤组织形成试管球茎，其中以MS+2.0 mg／L BA+0.5 mg／L NAA表现最佳。在一定范围内提高培养基中蔗糖浓度有利于白魔芋试管球茎形成，其中6％(w／v)蔗糖效果最好。而花魔芋试管球茎形成所要求的蔗糖浓度较低(4％)。光周期对试管球茎的发生影响不明显，而培养温度的影响则较大，22℃条件下，花魔芋和白魔芋试管球茎形成率、形成个数和单球茎鲜重均达最大值。试管球茎在培养过程可产生次级小球茎以进行增殖，但自然增殖率较低，通过损伤主芽方法打破主芽顶端优势，可促使次级小球茎的发生。
5．选择性培养魔芋愈伤组织有助于降低再生植株遗传变异频率。对魔芋Ⅲ型愈伤组织进行选择并于适宜培养基上继代培养，弃除状态发生改变的愈伤组织类型，获得不同培养时期的4个再生植株群体G0(初代培养产生)、G2(继代3次产生)、G4(继代5次产生)和G7(继代8次产生)。经两种分子标记技术(RAPD和ISSR)分析显示，G0群体变异频率最高(RAPD：15.6％；ISSR：18.0％)，G2、G4和G7的变异频率依次下降。四个群体中，G0再生植株的RAPD差异位点数最多，而其它三个群体的植株差异位点数较少。ISSR技术检测的差异位点数比RAPD技术检测到的要高，但在四个群体中分布差异不明显。上述结果表明，通过对Ⅲ型愈伤组织选择性培养不会导致变异频率增加，魔芋组织培养中的不稳定性主要发生于基因组中重复区域，造成植株性状改变的可能性较小。
6．试管球茎是建立魔芋繁殖体系的适宜方式，魔芋试管球茎田间生长优势比试管苗明显。将魔芋试管球茎和试管苗进行田间种植，结果发现，试管球茎和试管苗均可用作种芋繁殖。但试管苗只在特定的苗龄期时成活率高，且受移栽时温度影响。试管球茎需要一段时间低温贮藏以完成生理休眠，其植株生长势强，成活率高，体积大的试管球茎(重量大于0.5g)优势更明显。与试管苗相比，由试管球茎产生的球茎体积大、重量高，更适宜于商品种芋的生产。
Up to now, konjac is the only higher plant of vegetable kingdom that largely synthesizes konjac glucomannan, an importantly industrial material. Konjac is perennial herbaceous species belonging to Amorphophallus Blume (Araceae) and widely distributed throughout the mountainous areas in Southwest China with a long history of cultivation and utilization. Konjac glucomannan, a high-molecular-weight carbohydrate, is main storage substance in konjac corm. Due to its excellent physico-chemical characteristics, such as glue dissolving, gelling, thickening, film forming and mixing with other plant glues, konjac glucomannan has high values and is used widely in the food, chemistry, medicine and health protecting, and environment protection.Konjac is a one-leaf plant with a single petiole and conventionally propagated through rhizomes produced by the main corms with a low propagation coefficient. In addition, continuous asexual reproduction accumulates pathogens in corms that causes character degeneration and hence a dramatic loss in yield and degradation in quality. Due to lack of stable seed-corm propagation system, the quantity and quality of seed corm have been bottlenecks hampering development of konjac cultivation in a large scale. Since 1980s, many researchers have paid attention to development of plant tissue culture system for konjac propagation via using in vitro plants. However, in vitro plants had many disadvantages, such as low survival rate after transplantation, requiring extensive care of field management and incapability of being supplied year-around and long-distance deliver. Since 1990s, people have approached gradually the mechanisms of plant storage organ formation, and induced many asexually propagated organs in vitro, such as tuber, corm, bulblet, etc. Among them, the production system of potato seed tuber is most mature and has been industrialized, but little information is available on konjac corm production in vitro. Using konjac (A. riviveri and A. albus) as experimental materials, we carried out a set of researches on in vitro corm formation and propagation methods. The results as followings:1. Yong petiole was optimum explant for konjac tissue culture due to high frequency of desirable callus induction. MS + 1.0 mg/L NAA + 1.0 mg/L BA was optimum for callus induction and MS+0.5 mg/L NAA + 0.5 mg/L BA for callus proliferation and growth. During callus subculture, three types of calli were identified: I. watered, translucent; II. yellowish, loose in structure; III. pink or greenish, nodular or globular, compact in texture. On differentiation medium, except for type I callus, type II and III are capable of differentiating buds. Culture of typeⅢcallus could produce complete plantlets via corm-like structure pathway. In specific culture conditions, the types of calli could change mutually. Histological observations showed that different types of calli composed of different types of cells, of which typeⅢcallus was semi-organized tissue composed of slowly proliferated cells that contain starch and glucomannan granules. Moderate concentrations of cytokinins (BA, KT, ZT and TDZ) (1.0～2.0 mg/L) combined with low concentrations of NAA (0.1～1.0 mg/L) were optimum for typeⅢcallus differentiation.
2. In vitro morphogenesis of konjac occurred manily through organogenesis and adventitious bud and corm-like structure (CLS) were main pathways. Histological observations showed that cambium was absent in konjac petiole and that the cells outside vascular bundles of petiole firstly started dedifferentiation division to form callus. In vitro morphogenesis in A. rivieri occurred mainly through organogenesis while somatic embryogenesis took place under special culture conditions but with very low frequency. Most somatic embryos cannot convert to plants due high-frequency abnormality in these embryos. In vitro plants were obtained through adventitious bud or CLS pathways. The CLS derived plants had complete root systems without rooting culture while adventitious buds could not root during differentiation culture and grew slim. Both adventitious bud and CLS originated from subepidermal cells of typeⅢcallus that firstly form meristematic masses during further differentiation culture. The meristematic masses could directly develop into bud primordia that developed further into adventitious buds. Meanwhile, the masses could convert to interim globoids that have apical meristem and showed indistinct leaf primordia and developed into CLSs and complete plantlets during further subcultures.
3. Endogenous GA_3 content increased with bud formation whilst JA increased with corm development, indicating balance of endogenous hormones a key factor influencing organogenesis pathways. During the course of petiole parenchyma cells to form callus, the contents of endogenous hormones (IAA, GA_3, ABA and JA) changed obviously. Endogenous IAA level fell at initial culture stage and then rapidly rose 10d later and was stable after culture for 30d. Endogenous ABA content fell within first 20d and increased during further culture. Endogenous GA_3 and JA content decreased during the whole induction culture. The alternation of contents of the three endogenous hormones (GA_3, ABA and JA), either in the callus that formed CLSs or in the one that differentiated adventitious buds, showed different tendencies. The content of endogenous GA_3 hardly changed in CLS organogenesis but rapidly increased in adventitious bud organogenesis. Endogenous ABA level was almost stable during CLS organogenesis but rapidly fell in adventitious bud organogenesis pathway. Endogenous JA level rose obviously in CLS formation but showed an indistinct change in adventitious bud differentiation. Only the content of endogenous IAA decreased consistently in both the two organogenesis pathways. The balance of GA3 to ABA or JA showed different patterns in the two pathways, but the latter one showed most remarkable difference. The GA_3/JA ratio showed completely reversed tendencies in the two pathways. These results indicated that the pathways of konjac organogenesis in vitro were related to the changes of endogenous hormones and their balances.
4. In vitro konjac corm formation was influenced by medium and culture condition. The combinations of BA at moderate concentrations (1.0～2.0 mg/L) and NAA at low concentration (0.1～0.5 rag/L) favoured corm formation from type III callus. Among these combinations, MS+2.0 mg/L BA+0.5 mg/L NAA was most effective for both A. rivieri and A. albus. Increase of sucrose concentrations in a certain range (2-8%) favoured A. albus corm formation; 6% (w/v) was most efficient. As for A. rivieri, lower concentration (4%) was required for corm formation. In vitro corm formation was not affected by light periods but greatly influenced by culture temperature. Cultured in 22℃, the percentage of corm formation, mean number of corms and mean corm fresh weight reached their maximum. Wounding the apical meristem of the corm could overcome apical dominance and promote cormel propagation.
5. Selection and culture of konjac callus favoured lowering the genetic variation frequency of plants regenerated from the callus. Four groups of regenerated plants GO (derived from the first culture), G2 (derived from the third subculture), G4 (derived from the fifth subculture) and G7 (derived from the eighth subculture) were obtained by selection and culture of well-developed typeⅢcallus. RAFD and ISSR analyses showed that the genetic variation frequency of G0 was the highest (RAPD: 15.6%; ISSR: 18.1%) followed by G2, G4 and G7. Among the four groups, GO had more variation loci (RAPD) than those of the other three. The variation loci detected by ISSR were more than those detected by RAFD and distributed randomly throughout the four groups. From these results, selective culture of typeⅢcallus provided a good measure to control genetic variation of their regenerated plants. The results revealed that the genetic instability induced by konjac tissue culture mainly took place in repeat regions of konjac genome, which hardly results in morphological variation.
6. In vitro corm system is a suitable way applicable to konjac propagation. In vitro corm had obvious advantages over in vitro plants in adaptability under field conditions. The growth of both in vitro corm and plantlets was compared in the field and the results revealed that although both in vitro derived plants and corms could produce corms after transplantation the survival rate of in vitro plants was much affected by temperature during transplanting. However, in vitro corm grew vigorously and had high survival rate, particularly when the large corms (over 0.5g in weight) were used and the physiological dormancy was broken. The harvested corms produced by in vitro corms were larger in size than those produced by plantlets and hence more suitable for commercial propagation of seed corms.
魔芋为单叶柄支撑的单叶植物，通常利用球茎着生的根状茎进行无性繁殖，其繁殖系数极低，且多年沿用无性繁殖造成病原菌的积累、种性的退化，进而导致魔芋产量下降、品质劣化。由于缺乏稳定的良种繁育体系，种芋的数量和质量已经成为魔芋生产发展的瓶颈。自上个世纪80年代以来，许多研究者试图利用组织培养技术生产脱毒试管苗，以繁殖种芋，但试管苗因移栽成活率低且对田间管理技术要求较高，不能实现周年供应和长距离运输等缺点，无法在实际生产中广泛运用。二十世纪90年代以来，随着对植物变态器官发育机理认识的不断深入，人们在试管中已成功诱导出了块茎、球茎、鳞茎等营养繁殖器官，其中马铃薯试管块茎生产体系最为成熟，已经实现了工厂化生产，但有关魔芋试管球茎的研究国内外尚未见报道。本研究以花魔芋(A．rivieri)和白魔芋(A．albus)为材料，对魔芋试管球茎的形成及魔芋组织培养繁殖方式开展了系列研究，主要结果如下：
1．幼嫩叶柄是魔芋组织培养较好的外植体，愈伤组织形成率高，状态好。MS+1.0mg／L NAA+1.0mg／L BA适宜于愈伤组织诱导，而MS+0.5 mg／L NAA+0.5 mg／L BA有利于愈伤组织增殖。愈伤组织在继代培养过程中可形成3种不同的类型，即Ⅰ：水渍状，半透明；Ⅱ：淡黄色，结构疏松；Ⅲ：浅红或绿色，呈瘤状或球状，结构致密。在分化培养基上，Ⅰ型愈伤组织不能分化出芽，Ⅱ型可分化出不定芽，而Ⅲ型可通过拟球茎再生完整植株。这三种类型愈伤组织在特定的培养条件下可相互转化。组织学观察显示，三种类型的愈伤组织其组成细胞类型明显不同，其中Ⅲ型愈伤组织由保守分裂型细胞组成，其细胞中含有淀粉和葡甘聚糖颗粒，是一种半组织化结构。中等浓度(1.0～2.0 mg／L)的细胞分裂素(BA、KT、ZT和TDZ)与低浓度(0.1～1.0mg／L)的NAA配合均能促使Ⅲ型愈伤组织再生植株。
2．魔芋离体形态建成以器官发生途径为主，拟球茎和芽发生是主要形式。组织学观察显示，魔芋叶柄无维管组织形成层，维管束外层薄壁在培养过程首先启动脱分化，进而形成愈伤组织。魔芋形态建成以器官发生途径为主，在特定培养条件下也可形成体细胞胚，但频率极低，且绝大部分体胚因高度畸形化而不能形成再生植株。魔芋可通过不定芽器官发生途径或拟球茎器官发生途径形成植株，通过拟球茎形成的植株不需生根培养就具有完整的根系，而不定芽在发育过程中不能直接生根且植株生长势弱。不定芽和拟球茎均起源于Ⅲ型愈伤组织的次表皮细胞，这类细胞在分化过程中首先形成拟分生组织团。拟分生组织团在分化培养中既可直接形成芽原基，进而发生不定芽，可形成一种中间的球状体，这种球状体具顶芽分生组织，叶原基不明显，它可进一步发育成拟球茎，进而形成完整植株。
3．内源GA_3含量增加伴随着不定芽发生，而JA含量增加伴随着拟球茎发生，内源激素平衡是器官发生方式的关键调空因素。魔芋叶柄薄壁细胞在脱分化形成愈伤组织过程中，其内源激素(IAA、GA_3、ABA和JA)含量发生了明显的变化。内源IAA含量培养初期缓慢下降，10d后快速上升，30d后达到平稳。内源ABA含量在培养20d内呈下降趋势，之后迅速上升。内源GA_3和JA含量则在整个愈伤组织诱导过程中迅速下降。研究发现，三种内源激素(GA_3、ABA和JA)在形成芽和拟球茎的愈伤组织中含量变化明显不同。内源GA_3水平在拟球茎发生过程中变动不大，而在不定芽发生过程中快速上升，其绝对含量也比前者高；内源ABA在拟球茎发生过程中基本维持稳定，而在不定芽发生过程中则迅速下降；内源JA含量在拟球茎形成过程中呈明显上升趋势，而在不定芽发生过程中变化不明显。只有内源IAA无论是拟球茎还是不定芽形成过程其含量均呈下降趋势。内源GA_3与ABA和JA的平衡水平在两种不同器官发生途径中呈现出明显的差异，但GA_3／JA差异最为显著，在两种器官发生途径中其变化趋势完全相反。上述结果说明，魔芋离体器官发生形式与内源激素变化及平衡水平相关。
4．魔芋试管球茎(即拟球茎)的形成受培养基和培养条件的影响。中等浓度的BA(1.0～2.0 mg／L)与低浓度的NAA(0.1～0.5 mg／L)配合有利于Ⅲ型愈伤组织形成试管球茎，其中以MS+2.0 mg／L BA+0.5 mg／L NAA表现最佳。在一定范围内提高培养基中蔗糖浓度有利于白魔芋试管球茎形成，其中6％(w／v)蔗糖效果最好。而花魔芋试管球茎形成所要求的蔗糖浓度较低(4％)。光周期对试管球茎的发生影响不明显，而培养温度的影响则较大，22℃条件下，花魔芋和白魔芋试管球茎形成率、形成个数和单球茎鲜重均达最大值。试管球茎在培养过程可产生次级小球茎以进行增殖，但自然增殖率较低，通过损伤主芽方法打破主芽顶端优势，可促使次级小球茎的发生。
5．选择性培养魔芋愈伤组织有助于降低再生植株遗传变异频率。对魔芋Ⅲ型愈伤组织进行选择并于适宜培养基上继代培养，弃除状态发生改变的愈伤组织类型，获得不同培养时期的4个再生植株群体G0(初代培养产生)、G2(继代3次产生)、G4(继代5次产生)和G7(继代8次产生)。经两种分子标记技术(RAPD和ISSR)分析显示，G0群体变异频率最高(RAPD：15.6％；ISSR：18.0％)，G2、G4和G7的变异频率依次下降。四个群体中，G0再生植株的RAPD差异位点数最多，而其它三个群体的植株差异位点数较少。ISSR技术检测的差异位点数比RAPD技术检测到的要高，但在四个群体中分布差异不明显。上述结果表明，通过对Ⅲ型愈伤组织选择性培养不会导致变异频率增加，魔芋组织培养中的不稳定性主要发生于基因组中重复区域，造成植株性状改变的可能性较小。
6．试管球茎是建立魔芋繁殖体系的适宜方式，魔芋试管球茎田间生长优势比试管苗明显。将魔芋试管球茎和试管苗进行田间种植，结果发现，试管球茎和试管苗均可用作种芋繁殖。但试管苗只在特定的苗龄期时成活率高，且受移栽时温度影响。试管球茎需要一段时间低温贮藏以完成生理休眠，其植株生长势强，成活率高，体积大的试管球茎(重量大于0.5g)优势更明显。与试管苗相比，由试管球茎产生的球茎体积大、重量高，更适宜于商品种芋的生产。
Up to now, konjac is the only higher plant of vegetable kingdom that largely synthesizes konjac glucomannan, an importantly industrial material. Konjac is perennial herbaceous species belonging to Amorphophallus Blume (Araceae) and widely distributed throughout the mountainous areas in Southwest China with a long history of cultivation and utilization. Konjac glucomannan, a high-molecular-weight carbohydrate, is main storage substance in konjac corm. Due to its excellent physico-chemical characteristics, such as glue dissolving, gelling, thickening, film forming and mixing with other plant glues, konjac glucomannan has high values and is used widely in the food, chemistry, medicine and health protecting, and environment protection.Konjac is a one-leaf plant with a single petiole and conventionally propagated through rhizomes produced by the main corms with a low propagation coefficient. In addition, continuous asexual reproduction accumulates pathogens in corms that causes character degeneration and hence a dramatic loss in yield and degradation in quality. Due to lack of stable seed-corm propagation system, the quantity and quality of seed corm have been bottlenecks hampering development of konjac cultivation in a large scale. Since 1980s, many researchers have paid attention to development of plant tissue culture system for konjac propagation via using in vitro plants. However, in vitro plants had many disadvantages, such as low survival rate after transplantation, requiring extensive care of field management and incapability of being supplied year-around and long-distance deliver. Since 1990s, people have approached gradually the mechanisms of plant storage organ formation, and induced many asexually propagated organs in vitro, such as tuber, corm, bulblet, etc. Among them, the production system of potato seed tuber is most mature and has been industrialized, but little information is available on konjac corm production in vitro. Using konjac (A. riviveri and A. albus) as experimental materials, we carried out a set of researches on in vitro corm formation and propagation methods. The results as followings:1. Yong petiole was optimum explant for konjac tissue culture due to high frequency of desirable callus induction. MS + 1.0 mg/L NAA + 1.0 mg/L BA was optimum for callus induction and MS+0.5 mg/L NAA + 0.5 mg/L BA for callus proliferation and growth. During callus subculture, three types of calli were identified: I. watered, translucent; II. yellowish, loose in structure; III. pink or greenish, nodular or globular, compact in texture. On differentiation medium, except for type I callus, type II and III are capable of differentiating buds. Culture of typeⅢcallus could produce complete plantlets via corm-like structure pathway. In specific culture conditions, the types of calli could change mutually. Histological observations showed that different types of calli composed of different types of cells, of which typeⅢcallus was semi-organized tissue composed of slowly proliferated cells that contain starch and glucomannan granules. Moderate concentrations of cytokinins (BA, KT, ZT and TDZ) (1.0～2.0 mg/L) combined with low concentrations of NAA (0.1～1.0 mg/L) were optimum for typeⅢcallus differentiation.
2. In vitro morphogenesis of konjac occurred manily through organogenesis and adventitious bud and corm-like structure (CLS) were main pathways. Histological observations showed that cambium was absent in konjac petiole and that the cells outside vascular bundles of petiole firstly started dedifferentiation division to form callus. In vitro morphogenesis in A. rivieri occurred mainly through organogenesis while somatic embryogenesis took place under special culture conditions but with very low frequency. Most somatic embryos cannot convert to plants due high-frequency abnormality in these embryos. In vitro plants were obtained through adventitious bud or CLS pathways. The CLS derived plants had complete root systems without rooting culture while adventitious buds could not root during differentiation culture and grew slim. Both adventitious bud and CLS originated from subepidermal cells of typeⅢcallus that firstly form meristematic masses during further differentiation culture. The meristematic masses could directly develop into bud primordia that developed further into adventitious buds. Meanwhile, the masses could convert to interim globoids that have apical meristem and showed indistinct leaf primordia and developed into CLSs and complete plantlets during further subcultures.
3. Endogenous GA_3 content increased with bud formation whilst JA increased with corm development, indicating balance of endogenous hormones a key factor influencing organogenesis pathways. During the course of petiole parenchyma cells to form callus, the contents of endogenous hormones (IAA, GA_3, ABA and JA) changed obviously. Endogenous IAA level fell at initial culture stage and then rapidly rose 10d later and was stable after culture for 30d. Endogenous ABA content fell within first 20d and increased during further culture. Endogenous GA_3 and JA content decreased during the whole induction culture. The alternation of contents of the three endogenous hormones (GA_3, ABA and JA), either in the callus that formed CLSs or in the one that differentiated adventitious buds, showed different tendencies. The content of endogenous GA_3 hardly changed in CLS organogenesis but rapidly increased in adventitious bud organogenesis. Endogenous ABA level was almost stable during CLS organogenesis but rapidly fell in adventitious bud organogenesis pathway. Endogenous JA level rose obviously in CLS formation but showed an indistinct change in adventitious bud differentiation. Only the content of endogenous IAA decreased consistently in both the two organogenesis pathways. The balance of GA3 to ABA or JA showed different patterns in the two pathways, but the latter one showed most remarkable difference. The GA_3/JA ratio showed completely reversed tendencies in the two pathways. These results indicated that the pathways of konjac organogenesis in vitro were related to the changes of endogenous hormones and their balances.
4. In vitro konjac corm formation was influenced by medium and culture condition. The combinations of BA at moderate concentrations (1.0～2.0 mg/L) and NAA at low concentration (0.1～0.5 rag/L) favoured corm formation from type III callus. Among these combinations, MS+2.0 mg/L BA+0.5 mg/L NAA was most effective for both A. rivieri and A. albus. Increase of sucrose concentrations in a certain range (2-8%) favoured A. albus corm formation; 6% (w/v) was most efficient. As for A. rivieri, lower concentration (4%) was required for corm formation. In vitro corm formation was not affected by light periods but greatly influenced by culture temperature. Cultured in 22℃, the percentage of corm formation, mean number of corms and mean corm fresh weight reached their maximum. Wounding the apical meristem of the corm could overcome apical dominance and promote cormel propagation.
5. Selection and culture of konjac callus favoured lowering the genetic variation frequency of plants regenerated from the callus. Four groups of regenerated plants GO (derived from the first culture), G2 (derived from the third subculture), G4 (derived from the fifth subculture) and G7 (derived from the eighth subculture) were obtained by selection and culture of well-developed typeⅢcallus. RAFD and ISSR analyses showed that the genetic variation frequency of G0 was the highest (RAPD: 15.6%; ISSR: 18.1%) followed by G2, G4 and G7. Among the four groups, GO had more variation loci (RAPD) than those of the other three. The variation loci detected by ISSR were more than those detected by RAFD and distributed randomly throughout the four groups. From these results, selective culture of typeⅢcallus provided a good measure to control genetic variation of their regenerated plants. The results revealed that the genetic instability induced by konjac tissue culture mainly took place in repeat regions of konjac genome, which hardly results in morphological variation.
6. In vitro corm system is a suitable way applicable to konjac propagation. In vitro corm had obvious advantages over in vitro plants in adaptability under field conditions. The growth of both in vitro corm and plantlets was compared in the field and the results revealed that although both in vitro derived plants and corms could produce corms after transplantation the survival rate of in vitro plants was much affected by temperature during transplanting. However, in vitro corm grew vigorously and had high survival rate, particularly when the large corms (over 0.5g in weight) were used and the physiological dormancy was broken. The harvested corms produced by in vitro corms were larger in size than those produced by plantlets and hence more suitable for commercial propagation of seed corms.
引文
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