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題 名 | 蔗芒屬間雜種(Saccharum-Miscanthus)高貴化遺傳育種之研究 |
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作 者 | 陳燿煌; | 書刊名 | 國立臺灣大學農學院研究報告 |
卷 期 | 33:4 1993.12[民82.12] |
頁 次 | 頁345-350 |
分類號 | 434.714 |
關鍵詞 | 甘蔗; 芒; 蔗芒雜種; 高貴化; 過氧化酶; 酶譜; 形態; 解剖; 染色體不減數配子; 染體倍加; 複多元體; Sugarcane; Miscanthus; Saccharum-miscanthus hybrid; Nobilization; Peroxidase; Zymogram; Morphology; Anatomy; Chromosome; Chromosome doubling; Unreduced gamete; Amphipolyploidy; |
語 文 | 中文(Chinese) |
中文摘要 | 1975年起,台灣糖業研究所育種系開始一連串本省原產芒屬物種的利用 工作,利用的目的是希望將芒所具有的優良特性包括分�R力、宿根力、適應性和 抗病性引進到栽培甘蔗,一方面擴大甘蔗育種的親緣,一方面希望在高貴化的後 裔中,選拔到優秀的品系,命名做為栽培品種或提供做為雜交親本。從芒原種的 採集開始,經由雜交和一連串回交等高貴化的步驟。截至1989年止,共採集330 株系,從F1~BC5世代完成265個高貴化雜交,培育87878株實生苗,入選880個優 良營養系,並有部份後裔進入例行性的甘蔗品系比較試驗,顯現高貴化的成效。 本論文的主要目的,在於檢討蔗芒高貴化育種的過程中,各世代後裔雜種的真偽? 芒優良特性的傳遞,栽培甘蔗一般特性的累積,細胞遺傳學上染色體組成及傳 遞,並嘗試從不同的染色體組成進行高貴化的可能性。 以過氧化�t同位酵素�t譜為標誌,發現芒原種和栽培甘蔗親本各具特異的環帶, 其中S14環帶為全部膠片中染色最濃,完全不具多型性且為所有甘蔗親本皆出現 者,此環帶為甘蔗親本遺傳成份最佳的標誌。芒原種M8,9,10,11,12,13, 14等環帶恰位於甘蔗親本環帶S4和S5之間,且染色清楚,分辨容易,雖具高度多 型性且呈分離狀態,但分離的環帶從未逾越S4和S5的界限,且此區域不再有任何 甘蔗親本出現明顯的環帶,因此以此區域出現的環帶為芒原種遺傳成份的標誌。 就栽培甘蔗遺傳成份的標誌S14環帶而言,從F1~BC4所有參試高貴化後裔,全部 出現S14環帶,無一例外。就芒原種遺傳成份的標誌而言,除芒原種TM75-30和 TM75-38當然出現標誌環帶以外,F1後裔9/9(9個全部),BC1,8/10,BC2,7/9, BC3,9/9,BC4,8/10於S4和S5之間的區域出現一個以上的環帶。由以上的結果, 可知在高貴化過程連續回交時,理論上芒的染色體有完全丟失的可能,但在適當 的選拔壓力下,直到BC4世代大部份的後裔確信仍保有芒原種的遺傳成份。 以芒原種、栽培甘蔗親本及其高貴化各世代後裔為材料,檢討蔗芒高貴化過程中 形態與解剖的變化。就芒之可利用特性而言,本研究證實,芒原種確實具有旺盛 的分�R特性優越的宿很能力,良好的環境適應性和對露菌病和黑穗病的極強抗病 性。這些特性中,黑穗病抗性完全沒有傳遞給高貴化後裔的跡象,適應性和露菌 病的抗性可傳遞至BC1或BC2世代,分力�R和宿根方則可進一步傳遞到BC3或BC4 世代的部份後裔。栽拮甘蔗一般特性的累積,可到分為九種類型,一、數量性狀, 蔗芒差異顯著,F1世代起,各後裔的表現即與栽培甘蔗親本沒有差異。二、數量 性狀,蔗芒差異顯著,隨著高貴化世代的增加,性狀的表現,逐漸的趨近栽培甘 庶親本。三、數量性狀,蔗芒差異顯著,高貴化各世代後裔,調查數值先上升, 後再下降,逐漸趨近於栽培甘蔗親本,少數世代的表現甚至顯著的超過栽培甘蔗 親本。四、數量性狀,蔗芒差異顯著,而且高遺化各世代後裔的表現皆顯著的不 同於我培甘蔗親本,也就是完全沒有達到高貴化。五、數量性狀,蔗芒差異不顯 著,高貴化各世代後裔的表現,亦類似於芒原種或栽培甘蔗親本,無一定的傾向。 六、數量性狀,蔗芒差異不顯著,但是高貴化各世代後裔的表現,全部超過芒原 種與栽培甘蔗親本,達顯著差異,充分顯示屬間雜種優勢的現象。七、數量性狀, 蔗芒差異不顯著,高貴化各世代後裔的表現,一般是數值先上升,在某一、二世 代,與芒原種或栽培甘蔗親本差異達顯著後,再降低而接近栽培甘蔗親本。八、 質的性狀,芒原種,相對於高貴化世代後裔和栽培甘蔗視本,表現截然不同。九、 質的性狀,芒原種,栽培甘蔗親本和高貴化各世代後裔表現類似。 綜合體細胞染色體數目,花粉母細胞減數分裂染色體之配對和蔗莖節間組織對碘 液之反應等結果顯示,芒原種,利用工作,巧世代是經由SMM染色體組之雜種 進行高貴化育種。高貴化各世代理論上染色體之組成,芒原種為MM,栽培甘蔗 親本為SS,F1為SMM,BC1為3/2SM,BC2為7/4 S1/2M,BC3為15/8 S1/4M,BC, 為31/16 S1/8M。F1雜種SMV染色體組,MM可能由父本芒原種未減數花粉2n染色 體所傳遞。父本芒原種之末減數花粉,可能來自減數分裂過程中,四分子期的三 分子。此三分子可能由第二減數分裂二子細胞不同步分裂所造成,整個高貴化的 過程中,芒原種或栽培甘蔗親本的染色體皆有丟失的可能,可於第一減數後期因 逢機分離而丟失,其中尚牽涉二子細胞染色體數的不對等分離:方可於減數分裂 的過程中,因遲滯染色體或微核被排除於子細胞的核外而丟失。本研究並首次計 對芒原種、栽培甘蔗親本及屬間F1雜種減數分裂過程中,核仁數目的變化進行觀 察。 本研究成功的建立兩套蔗芒屬間雜種組織和細胞培養的模式系統。模式一,莖頂 分生組織誘導癒合組織後,芽體直接自此癒合組織上自動分比。模式二,華頂分 生組織誘導癒合組織,經繼代培養增值後,進行細胞懸浮培養一段時期,細胞團 再誘導芽體之分化。採用模式二的系統,利用蔗芒屬間雜種SM-79006,SM-79016 和SM-79037等為材料,以濃度50ppm和100ppm秋水仙鹼處埋懸浮培養細胞,經再 生得到868株衍生系。逢機選取來自SM-79016之28株衍生系,進行氣孔長度測量 及體細胞分裂鏡檢,發現氣孔長度介於25.1μm~45.0μm之間,染色體數目則分 佈於2n=Ca.103~220。二者的關係顯示氣孔長度在34.4μm以上者為染色體加倍之 複多元體植株,染色體未加倍的個體其氣孔長度皆小於34.4μm。秋水仙鹼對衍 生系染色體倍加有良好的效果,濃度50和100ppm時各有42.0和16.1的衍生系為複 多元體,未經秋水仙鹼處理之對照組,其染色體沒有倍加的現象。 農藝性狀調查結果顯示,染色體倍加的個體,某些與栽培甘蔗有關的性狀表現良 好,包括蔗莖內容佳、空蒲心小,蔗汁錘度高且完全不開花。但是亦同時造成某 些性狀的劣化,包括較差的生長勢,較短的莖長、較細的莖徑、較少的莖數,葉 片更短更狹窄,單株蔗產量甚低,植株甚為衰弱,生育競爭力極差,11月蔗齡時 只剩57.4的植株可存活。此蔗芒屬間雜種複多元體遺傳行為與育種價值有待進一 步研究 |
英文摘要 | The utilization of Miscanthus spp. has been an explorativeapproach in sugarcane improvement since 1975 at Taiwan SugarResearch Institute (TSRI). An attempt has been made to introduceuseful characteristics such as profuse tillering, strong ratooningability, wide adaptation and disease resistance from Miscainthus spp. to sugarcane cultivars. Intense breeding researches were carried outon collection of Miscanthus germplasm, nobilization hybridizationand evaluation in each generation of nobilization. Up to 1989, a totalof 330 Miscanthus clones have been collected and 87878 seedlings of F1, BC1, BC2, BC3, BC4 and BC5 progenies were raised from 265 crosscombinations. Promising hybrid clones have been selected and are currentlyunder tests at stages T2 through T5 of the routine breedingprogram. All nobilized progenies in each generation , as well as Miscacnthus spp. and Saccharum parents were materials studied inthis thesis. We examined the pattern of peroxidase zymogram,variation of traits in morphology or anatomy, and composition ofgenetic background. We also made a explorative study in developingcolchichine-induced amphipolyploidy from cell suspension culture of Saccharuim-Miscanthus (S-M) hybrids. With pattern of peroxidase zymogram, differences in specificbands between Saccharum parents and Miscanthus spp. wereevident. Saccharum band S14 with the greatest sensitivity and nonpolymorphism was presented in .Saccharum parents only. It was thebest marker for genetic constitution of Saccharum parents. Miscanthus bands M8 ~ M14 with clear vision and segregatingpatterns were found between SaccAarum bands S4. and S5, andpresented in Miscanthus spp. only. These Miscanthus bands whichpositions were comparatively bounded by Saccharum band S4 and S5 could be used as markers for genetic constitution of Miscanthus spp. All nobilized progenies investigated in each generation, as well as Saccharum parents, showed the presence of S14 band. All 9 clones investigated in F1 and BC3, 8 of 10 clones in BC1 7 of 9 in BC2, 8 of10 in BC4, as well as Miscanthus spp. showed the presence of marker bands located between Saccharum bands S4 and S5. It is concluded that most of progenies in F1, BC1, BC2, BC3, and BC4 generation during nobilization process have both genetic constitutions of Miscanthus spp. and Saccharum parents simultaneously. Using Miscanthus spp., Saccharum parents and nobilized progenies as materials, we studied the variation of traits in morphology and anatomy during nobilization. The results showed that Miscanthus spp. actually had profuse number of tillers, strong ratooning ability, superior adaptation to different environments and highly resistance to downy mildew (Peronosclerospora sacchari) and culmicolus smut ( Ustilago scitaminea ). Among these useful traits of Miscanthus spp., reaction in F1~BC4 hybrids to colmicolus smut was strongly affected by the Saccharum parents. It seemed that the degree of resistance to downy mildew and adaptation to different environments in F1, BC1 and BC2 hybrids resembled those in Miscanthus spp. while in BC3 and BC4, the performance of these two traits was very close to Saccharum parents. Mobilized progenies of F1, BC1, BC2, BC3, and BC4 generation had much more number of tillers and stronger ability of ratooning than their Saccharum parents, although the number of tillers and the ability of ratooning decreased gradually with the advance in nobilization generations. The variation of traits in morphology and anatomy duringnobilization process could be classified into 9 categories as follows: 1. Quantitative traits were different between Saccharum parents and Miscanthus spp. significantly. The expression of traits in FI~BC4 hybrids however all resembled Saccharum parents. 2. Quantitative traits were significantly different between Saccharum parents and Miscanthus spp., The expression of traits however gradually approached to Saccharum parents when the nobilization generation advanced. 3. Quantitative traits were significantly different between Saccharum parents and Miscanthus spp.. The traits measured was increasing gradually in the early stages, and the traits in some hybrids generations showed significantly higer than Saccharuin parents. In later stages, they were decreasing and gradually approached to Saccharunm parents. 4. Quantitative traits were significantly different between Saccharum parents and .Wscautilus spp.. The differences of traits between SaccAas-urn parents and F1 ~ BC4 hybrids were all significantly different too, indicating that there was no nobilization during the back crosses. 5. Quantitative traits were not significantly different between Saccharum parents and Miscanthus spp., The expression of the traits in F1~BC4 hybrids were similar to Saccharum parents and Miscanthus spp.. No definite trend of nobilization could be found. 6. Quantitative traits were not significantly different between Saccharum parents and Miscanthus spp. The expression of these traits in F1 ~ BC4 Hybrids all showed superior to Saccfiarum parents and Miscanthus spp. and were all significantly different. 7. Quantitative traits were not significantly different between Saccharum, parents and Miscanthus spp. However the variation of traits in each generation during nobilization was similar to category 3. 8. Qualitative traits expressed in Miscanthus spp. as compared to Sacchartim parents and nobilized progenies were absolutely different. 9. Qualitative traits expressed in Miscanthus spp., .Saccharum parents and nobilized progenies were all similar. The results of studies on chromosome number of somatic cells, chromosome pairing during meiosis of pollen mother cell (PMC) and respond to iodine solution in internode tissue indicated that the expected genome composition during nobilization of Saccharum parents, Miscanthus spp. and nobilized progenies was as follows: Miscanthus spp. MM, Saccharum parents SS, F1 SMM, BC1 3/2S M, BC2 7/4S 1/2M, BC3, 15/8S 1/4M, BC4, 31/16S 1/8M. MM genome presented in F1 hydrid might be the result of un-reduced pollen of Miscanthus spp.. We had found triads in the tetrad stage during PMC meiosis and the triads with 2n number of chromosomes might develop into un-reduced pollen. The possible source of triads might be due to un-synchronized division in two daughter cells during meiosis II . During nobilization process, chromosomes from Saccharum parents or Miscanthus spp. could be elimilated. There are two possible explanations with regard to the chromosome elimination.1. The elimination of chromosome resulted from random segregation in anaphase I during meiosis while unequal segregation of chromosome numbers in this stage was also observed.2. The result of lagging chromosome or micronucleus which are common found in interspecific or intergeneric hybrids. In this study, we also observed the change of nucleolus in particularstage during meiosis. We had established two model systems in tissue and cellsuspension culture for S-M hybrids. Model I: explant→callus→plantregeneration automatically. Model II: explant→ callus →callus prolification→cell suspensionculture→cell cluster prolification→plant regeneration in regenerationmedium. Using model II, a population of 868 plants were regenerated fromcolchicine-treatment of cell suspension culture of three S-M hybridsfor the study of amphipolyploidy under S-M nobilization. Cytogenetic examination of the regenerated plants showed thatthere was a wide variation of chromosome numbers from 2n=Ca.l03~220, separating into two groups. Stomatal length in the regeneratedplants ranged from 25.1 μ m to 45.0 μ m. Plants have stomata longerthan 34.4 μm. belonged to the chromosome doubled group. The groupof un-doubled chromosome number had stomata shorter than 34.4μ m.The percentage of amphipolyploid occurred in colchicine-inducedsomaclones were 42.0 at 50ppm and 16.1 at 100ppm, while none wasobtained in un-treated control. The results of studies on agronomictraits showed that plants with chromosome doubled werenon-flowering, smaller pithness and cavity, higher brix, but weakervigorousness, shorter stalk length, thinner stalk diameter, fewerstalk number, shorter and narrower leaf and lower cane yield thanplants with chromosome un-doubled. |
本系統中英文摘要資訊取自各篇刊載內容。