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Development of a Transformation Protocol and Cell Culture System for the Commercially Important Species of Red Macroalga, Gracilaria Gracilis

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Development of a Transformation Protocol and Cell Culture System for the Commercially Important Species of Red Macroalga, Gracilaria Gracilis Town The copyright of this thesis rests with the University of Cape Town. No quotation from it or information derivedCape from it is to be published without full acknowledgement of theof source. The thesis is to be used for private study or non-commercial research purposes only. University DEVELOPMENT OF A TRANSFORMATION PROTOCOL AND CELL CULTURE SYSTEM FOR THE COMMERCIALLY IMPORTANT SPECIES OF RED MACROALGA, GRACILARIA GRACILIS by Town Suzanne Margaret Huddy Cape of A thesis submitted for the degree of Doctor of Philosophy in the Department of Molecular and Cell Biology, Faculty of Science, University of Cape Town, UniversitySouth Africa. November 2011 Abstract DEVELOPMENT OF A TRANSFORMATION PROTOCOL AND CELL CULTURE SYSTEM FOR THE COMMERCIALLY IMPORTANT SPECIES OF RED MACROALGA, GRACILARIA GRACILIS by Suzanne Margaret Huddy Marine Biotechnology Unit, Department of Molecular and Cell Biology, Faculty of Science, University of Cape Town, South Africa ABSTRACT Town Gracilaria gracilis belongs to a commercially important genus of red macroalgae that is used in the production of two commercially important grades of agar. In South Africa, the Gracilaria industry used to depend solely on theCape natural G. gracilis resource growing in Saldanha Bay, a resource which has beenof unreliable for commercial collections due to numerous population collapses over the past few years. Suspended cultivation has been suggested as the only means of establishing a reliable Gracilaria industry in South Africa. This type of cultivation however, is intensive and often leads to increased disease burden and stress. A better understanding of how this commercially important seaweed responds at a genetic level to stresses faced in the aquaculture environment would not only be advantageous to theUniversity South African industry, but this knowledge is essential for selecting and/or engineering macroalgal strains that are either more tolerant or resistant to these stresses. This requires in vivo analysis of G. gracilis gene function and regulation in order to introduce new or improved genes into G. gracilis, and for this to be possible, a protocol for transformation of recombinant DNA into G. gracilis is required. In this study a transformation and tissue culture system for G. gracilis was developed. These tools provide the necessary groundwork for future genetic manipulation studies that are essential for improving our understanding of the role that various genes play in stress response and tolerance in G. gracilis. ii Abstract Microparticle bombardment of G. gracilis thalli was investigated and optimized as a means to deliver foreign DNA into the macroalga. This technique proved successful for transformation of G. gracilis thalli with the lacZ reporter gene under the influence of the Simian virus 40 (SV40) promoter. Transient expression of the lacZ reporter gene was investigated further by comparing the effectiveness of the cauliflower mosaic virus 35S (CaMV 35S) and the Cytomegalovirus (CMV) promoters. The SV40 promoter was identified as the best promoter and selected for further transformation studies in this investigation. In addition to an effective foreign DNA delivery method, stable transformation of macroalgae requires clonal seaweed culture and techniques for plant regeneration from single cells. The isolation of macroalgal protoplasts and their regeneration into whole plants has been more successful when compared to callus induction and subsequent whole plant regeneration. In this study, G. gracilis protoplasts were investigated as a suitableTown cell culture system. A protocol for protoplast isolation and purification was developed and optimized in order to ensure that large quantities of viable protoplasts could consistently be isolated from G. gracilis thalli. Cell wall re-synthesis of G. gracilisCape protoplasts, the first step towards whole plant regeneration, was shown to occur over the first 24 hrs of culture using calcoflour staining and scanning electron microscopy.of Furthermore, the effect of light intensity and incubation temperature on whole plant regeneration from G. gracilis protoplasts was investigated. Under conditions of low light intensity (5 µmol photons m-2s-1) and high incubation temperature (18-19 ºC), G. gracilis protoplasts underwent cell division that resulted in the formation of cell clumps. However, under conditions of higher light intensity (30 µmol photons m-2Universitys-1) and either high (18-19 ºC) or low (14-15 ºC) culture temperatures, whole plants were regenerated from protoplasts. These either regenerated slowly over a period of 4-6 months to produce plants which resembled the parental plants, exhibiting slender, branched thalli, or regenerated rapidly over a period of 1-3 months to produce plants which remained small with thalli that were thick and unbranched. To the best of our knowledge, this study is the first to describe whole plant regeneration from protoplasts of the commercially important agarophyte G. gracilis. Having demonstrated that G. gracilis protoplasts can develop into mature plants, we investigated the possibility of transforming G. gracilis protoplasts by PEG-mediated transfection. A transformation protocol for these protoplasts was successfully developed and iii Abstract optimized. This was subsequently used to investigate the effects of 18S rDNA-targeted homologous recombination (HR) and tobacco Rb7 matrix attachment regions (MARs) on transgene expression using egfp as a test gene. This was done in an effort to identify a possible means of increasing transgene expression. A suite of vectors was designed, constructed and transfected into G. gracilis protoplasts after which EGFP levels and the presence of egfp were monitored for nine days post-transfection. The presence of tobacco Rb7 MARs and 18S rDNA regions on vector DNA resulted in significant (P<0.05) increases in relative fluorescence and therefore EGFP levels at both 3 and 4 days post-transfection. Furthermore, targeted HR was also shown to have taken place in G. gracilis protoplasts transfected with vector DNA containing 18S rDNA homologous regions. As successful transformation in plant systems requires the use of selectable markers, G. gracilis protoplasts were assessed for their sensitivity to the antibiotics kanamycin and chloramphenicol and the herbicide BASTA®. While G. gracilisTown protoplasts exhibited a high level of resistance to kanamycin and chloramphenicol, they were shown to be sensitive to BASTA®. Vectors containing the bar gene (which confers resistance to the herbicide) under the influence of the SV40 promoter/enhancerCape were constructed and transfected into protoplasts which were assessed for their ability to survive BASTA® selection. Transfected G. gracilis protoplasts exhibited significantlyof ( P<0.01) increased survival percentages in comparison to the negative control protoplasts which were not transformed with vector DNA. In addition, 4-5% of the protoplasts survived a second round of selection 21 days post- transfection. This study has establishedUniversity some key genetic tools which are essential for future genetic manipulation studies in G. gracilis. These include effective methods to deliver foreign DNA into G. gracilis thalli and protoplasts, a screening mechanism to select G. gracilis transformants from a multitude of recipients and a method to regenerate whole plants from protoplasts. This work also adds to the current scientific knowledge of general macroalgal, and more substantially and specifically to G. gracilis, transformation systems. iv Acknowledgements ACKNOWLEDGEMENTS Firstly, I would like to thank my Lord and Saviour Jesus Christ. Your continued strength, guidance and blessings have made this possible. The completion of this thesis is a true testament to your presence in my life. Thanks to my supervisor, Assoc. Prof. Vernon Coyne, for his guidance and support throughout my postgraduate studies at UCT. I have learnt much over my time in the Marine Biotechnology Unit, and I will always value my time there. To my co-supervisor, Dr Ann Meyers, your input and advice was invaluable, and I know without a doubt that this would have been a much larger obstacle without your support. Thank you!! Town To all the Lab 201 members over the years: Rob, Bron, BBC, Chris, René, Dave, Tanya, Amy, Rael, Taryn, Ros, Bridget, Caroline, Karusha, Kyle, Kim, Ayesha, Jerry, Rogan and Phil. Thank you for the friendships shared over theCape years. Work was always a little easier knowing there was a smiling face, a witty chirp, and someone nearby who understood exactly what you were going through. To my otherof friends and colleagues at MCB, especially Mariette, Lynn, Sarah and Brian. Thanks for your contribution to my experience and wonderful chats at tea-time(s). I would also like to thank MCB staff, including the SO’s, DA’s and academics for their assistance over the years.University In particular, a big thank you to Marion Bezuidenhout for her help and advice with the microparticle bombardment. A very big thanks to Irvine and Johnson Abalone Culture Division, Danger Point, Gansbaai, South Africa, for supplying me with seaweed throughout my study. Without your assistance this project would not have been impossible. Thank you also to Prof. John Bolton and the botany department for allowing us use of your facilities. I am especially grateful to the National Research Foundation and UCT for providing me with financial assistance through the course of my studies. Acknowledgements I have been blessed to have some
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