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Somatic Embryogenesis and Transformation Studies in Schlumbergera and Rhipsalidopsis

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Somatic Embryogenesis and Transformation Studies in Schlumbergera and Rhipsalidopsis Somatic Embryogenesis and Transformation Studies in Schlumbergera and Rhipsalidopsis Von der Naturwissenschaftlichen Fakultät der Universität Hannover zur Erlangung des akademischen Grades eines Doktors der Gartenbauwissenschaften -Dr. rer. hort.- genehmigte Dissertation von Ezz AL-Dein Muhammed Al-Ramamneh, M. Sc. Hort. (Jordan) geboren am 14. January 1973 in Sweileh, Jordan 2006 Referentin: Prof. Dr. Margrethe Serek Korreferentin: Prof. Dr. Sridevy Sriskandarajah Tag der Promotion: 12. Januar 2006 Zusammenfassung Ezz AL-Dein AL-Ramamneh Somatische Embryogenese und Untersuchungen zur Transformation bei Schlumbergera und Rhipsalidopsis Schlagwörter: Genetische Transformation - Osterkaktus - Transgen - Weihnachtskaktus Die In-vitro-Kultur bietet für Kakteen Möglichkeiten zur Vermehrung, somatischen Embryogenese und genetischen Transformation. In der vorliegenden Arbeit wurde somatische Embryogenese an Phyllokladien-Explantaten von Schlumbergera truncata cv. Russian Dancer induziert. Kallus, der sich an Phyllokladien-Explantaten gebildet hatte, wurde über einen Zeitraum von 16 Monaten auf MS-Medium mit Cytokininen kultiviert. Im Hinblick auf die Induktion somatischer Embryonen war diese Behandlung einer kürzeren Etablierungsphase überlegen. Somatische Embryonen differenzierten nach drei bis fünf Monaten, wenn Kallus, der in flüssigem Medium auf SH- oder MS-Basis mit 7 µM Kinetin kultiviert worden war, auf festes MS-Medium entweder mit 0,45 µM 2,4-D oder hormonfrei überführt wurde. Die Medienzusammensetzung, Wachstumsregulatoren sowie die Dauer der Kultur auf cytokininhaltigen Medien hatten einen Einfluss auf die Embryogenese. Die höchste durchschnittliche Anzahl von Embryonen in unterschiedlichen Stadien wurde erzielt, wenn die erste Kulturphase von 30 Tagen in flüssigem SH-Medium, und die zweite auf hormonfreiem MS-Medium erfolgte. Ungefähr 70 % der somatischen Embryonen keimten auf G-Medium. Anschließend wurden die auf somatische Embryonen zurückgehenden Pflanzen erfolgreich in Erde überführt. Sie zeigten genetische Stabilität im Vergleich zu den Ausgangspflanzen mit Ausnahme der Pflanzen, die sich aus somatischen Embryonen mit mehr als vier Keimblättern entwickelt hatten. Zusätzlich wurde ein sehr effizientes Regenerationssystem über Adventivsprossbildung für Schlumbergera cv. Alex und Rhipsalidopsis cv. CB5 entwickelt. Darüber hinaus wurden diese effizienten Regenerationssysteme weiterentwickelt und erfolgreich eingesetzt, um erstmals transgene Pflanzen von Rhipsalidopsis cv. CB5 zu erzeugen, die das uidA-Gen und das selektierbare Markergen nptII enthielten. Einige Faktoren, die die Transformation von Rhipsalidopsis-Kallus beeinflussen, wurden untersucht. Transformierte Rhipsalidopsis-Kallusse wurden nach längerer Kultur auf Medien mit 600 mg/l Kanamycin erhalten. Eine Vor-Inkubation von Agrobacterium tumefaciens in SIM- Medium mit Acetosyringon steigerte die Häufigkeit, mit der transgene Kallusse erzielt wurden. Desweiteren war ein zusätzlicher Waschschritt der Kallus-Explantate mit Cefotaxim nach der Ko-Kultur notwendig, um Agrobacterium tumefaciens zu unterdrücken. Der Verzicht auf Kanamycin im Medium für den letzten Kulturschritt sowie die Kultur der transformierten Kallusse unter Ernährungsstress führte zur Bildung transgener Adventivsprosse. Mit diesem Ansatz wurde eine Transformationsrate von 22,7 % erreicht. Die in dieser Arbeit erzielten Ergebnisse zeigen, dass der Agrobacterium-vermittelte Gentransfer ein vielversprechender Ansatz für die Erzeugung neuer Genotypen bei diesen beiden Kakteenarten ist. i Abstract Ezz AL-Dein AL-Ramamneh Somatic Embryogenesis and Transformation Studies in Schlumbergera and Rhipsalidopsis Keywords: Christmas cactus - Easter cactus - genetic transformation - transgene Tissue culture has emerged as a tool for cacti micropropagation, somatic embryogenesis and genetic manipulation. Somatic embryogenesis was induced from phylloclade explants of Schlumbergera truncata cv. Russian Dancer. Callus that developed on phylloclade explants and sub-cultured over a period of 16 months on MS medium containing cytokinins was superior for the induction of somatic embryos compared to callus grown for a shorter time in the establishment phase. Somatic embryos were induced after three to five months when callus grown in SH- or MS- based liquid media supplemented with 7.0 µM kinetin was transferred onto solid MS- based medium with either 0.45 µM 2,4-D or without hormones. Embryogenesis was affected by the type of medium, plant growth regulators and duration of callus exposure to cytokinins. The highest average numbers of embryos in the different stages were achieved using SH liquid medium for the first culture and MS medium without hormones after 30 days. Approximately 70% of somatic embryos germinated on G medium. Furthermore, plants derived from somatic embryos were successfully potted in soil, and they showed, except those derived from somatic embryos with more than four cotyledons, genetic stability compared to mother plants. A highly efficient regeneration system through adventitious shoot formation was also developed in Schlumbergera cv. Alex and Rhipsalidopsis cv. CB5. Moreover, the development of these efficient regeneration systems was further exploited and success was demonstrated as the first report in obtaining Rhipsalidopsis cv. CB5 shoots transgenic for the uidA gene and the selectable marker nptII gene. Some of the factors influencing transformation of Rhipsalidopsis callus explants were evaluated. Transformed Rhipsalidopsis calli were obtained by extended culture on media containing 600 mg/l kanamycin. The pre-incubation of Agrobacterium tumefaciens in SIM medium containing acetosyringone raised the frequency of transgenic calli. Furthermore, a washing step with cefotaxime for the callus explants after co-culture was necessary to remove the excess Agrobacterium. The removal of kanamycin from the final medium together with the culture of the transformed calli under nutritional stress led to the formation of transgenic adventitious shoots. With this approach, a transformation efficiency of 22.7% was achieved. The results obtained in this study suggested that Agrobacterium-mediated transformation is a promising approach for generating new genotypes of these cacti. ii Dedication To my Family and Friends With Love iii Acknowledgements I would like to express my sincere appreciation and deep gratitude to my advisor Prof. Dr. Margrethe Serek for her guidance, encouragement, patience and constant support. I would like also to thank Dr. Sridevy Sriskandarajah for introducing me to the field of tissue culture, for her valuable advice regarding my research project and for the critical evaluation of this manuscript. Special thanks go to Dr. Lilli Sander for her help with molecular methods. My appreciation is also extended to Dr. Stefan Frello for the technical advice on various methods throughout the course of this study. I would like to thank Dr. Michael Hansen, for his help with scanning electron microscopy. Special thanks also go to Lise Girsel, for her expertise in histology. My thanks and gratitude also go to my fellow PhD students and other staff members of the floriculture section in the Royal Veterinary and Agricultural University in Copenhagen for their support and for providing a good working environment both in the laboratory and in my office. I would like also to thank the staff members of the floriculture section in the University of Hannover for their readiness to help both on the personal and scientific level. Thanks Dr. Traud Winkelmann for your suggestions and advice in various aspects of this research project. Special thanks go to Al-Balqa Applied University, Salt, Jordan for the PhD grant which enabled me to successfully complete this doctoral study. Moreover, my gratitude goes to the German Academic Exchange Service (DAAD) for the financial support of a six-month German language course. Lastly, I would like to thank my parents and my brother Naser for their encouragement and moral support throughout this study. iv Table of Contents Zusammenfassung……………………………………………………………………...i Abstract……………………………………………………………………………......ii Dedication…………………………………………………………………………….iii Acknowledgements…………………………………………………………………...iv List of Tables……………………………………………………………………...….ix List of Figures…………………………………………………………………………x List of Appendices……………………………………………………………………xi Abbreviations………………………………………………………………………...xii 1.0 General Introduction………………………………………………………………1 1.1 Overview………………………………………………………………………..1 1.2 Tissue culture of the Cactaceae family…………………………………………3 1.3 Plant transformation…………………………………………………………….5 1.4 Statement of the problem……………………………………………………….6 1.5 Objectives……………………………………………………………………….9 Overall goal……………………………………………………………………....9 Specific goals…………………………………………………………………….9 2.0 Plant regeneration via somatic embryogenesis in Schlumbergera.........................10 Abstract…………………………………………………………………………..10 2.1 Introduction…...……………………………………………………………….11 2.1.1 Overview………………………………………………………………….11 2.1.2 Morphogenesis and regeneration…………………………………………11 2.1.3 Organogenesis…………………………………………………………….12 2.1.4 Somatic embryogenesis…………………………………………………..12 2.1.4.1 Growth regulators…………………………………………………13 A) Auxins…………………………………………………………….. 13 B) Cytokinins………………………………………………………….14 C) Abscisic acid (ABA)……………………………………………….15 D) Daminozide………………………………………………………...15 2.1.4.2 Sugars……………………………………………………………...16 2.1.4.3 Medium…………………………………………………………....17 2.1.4.4 Age of plant material and culture conditions……………………...18
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