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Diversity of Silverleaf Nightshade in Australia and Implications For Diversity of silverleaf nightshade in Australia and implications for management Xiaocheng Zhu B.Sc., Fujian Agriculture and Forestry University, China M.Sc. (by research), Fujian Agriculture and Forestry University, China A thesis presented to Charles Sturt University in fulfilment of the requirements for the degree of Doctor of Philosophy Faculty of Science Charles Sturt University Wagga Wagga, NSW, Australia February 2013 TABLE OF CONTENTS Table of Contents ................................................................................ i Certificate of Authorship ................................................................... iii Acknowledgements ........................................................................... iv List of Publications ........................................................................... vi Abstract ........................................................................................... viii Chapter 1 General introduction .......................................................... 1 Chapter 2 Literature review ............................................................... 9 2.1 Importance and distribution ..................................................... 9 2.2 Management ........................................................................... 13 2.2.1 Chemical control ............................................................. 13 2.2.2 Biocontrol ........................................................................ 15 2.2.3 Other management strategies .......................................... 16 2.3 Biology and ecology .............................................................. 17 2.3.1 Description ...................................................................... 17 2.3.2 Lifecycle and genetic diversity ....................................... 22 2.3.3 Germination and emergence............................................ 26 2.3.4 Spread .............................................................................. 28 i Chapter 3 Morphological variation of silverleaf nightshade (Solanum elaeagnifolium Cav.) in south-eastern Australia .......................................... 31 Chapter 4 Evaluation of simple sequence repeat (SSR) markers from Solanum crop species for Solanum elaeagnifolium ..................................... 43 Chapter 5 Development of SSR markers for genetic analysis of silverleaf nightshade (Solanum elaeagnifolium) and related species........... 51 Chapter 6 SSR marker analysis to determine the genetic variation in Solanum elaeagnifolium in Australia ........................................................... 59 Chapter 7 Genetic variation and structure of Solanum elaeagnifolium in Australia analysed by AFLP markers ...................................................... 67 Chapter 8 Identification of silverleaf nightshade using microsatellite markers and microstructure .......................................................................... 75 Chapter 9 Time of emergence impacts the growth and reproduction of silverleaf nightshade (Solanum elaeagnifolium Cav.) ............................. 83 Chapter 10 General discussion and conclusions .............................. 91 Appendix ........................................................................................ 104 References ...................................................................................... 109 ii CERTIFICATE OF AUTHORSHIP I, Xiaocheng Zhu, hereby declare that this submission is my own work and to the best of my knowledge and belief, understand that it contains no material previously published or written by another person, nor material which to a substantial extent has been accepted for the award of any other degree or diploma at Charles Sturt University or any other educational institution, except where due acknowledgement is made in the thesis. Any contribution made to the research by colleagues with whom I have worked at Charles Sturt University or elsewhere during my candidature is fully acknowledged. I agree that this thesis be accessible for the purpose of study and research in accordance with normal conditions established by the Executive Director, Library Services, Charles Sturt University or nominee, for the care, loan and reproduction of thesis, subject to confidentiality provisions as approved by the University. Signature 8th February 2013 iii ACKNOWLEDGEMENTS I would like to thank to my supervisory team: principle supervisor: Professor Deirdre Lemerle and co-supervisors: Drs Hanwen Wu, Harsh Raman, Geoffrey E. Burrows and Rex Stanton for their support, guidance and understanding through my PhD. Dr Rex Stanton also provided some great pictures of silverleaf nightshade for this thesis. Great appreciation also goes to Drs Neil Coombes, De Li Liu, Guangdi Li – NSW Department of Primary Industries and Mr Craig Poynter of Spatial Data Analysis Network (SPAN), Charles Sturt University and the National Climate Centre, Bureau of Meteorology, for their help on my data analysis. I thank Mr John Garvie of the Natural Resources Management Board, Mr Robert Thompson of the NSW Department of Primary Industries and a number of state, and local organisations across Australia for their assistance in field sampling. Thanks to Dr Rosy Raman, Ms Belinda Taylor, Dr Ata Rehman, Dr Ben Stodart, Dr John Harper, Ms Natalie Allison, Dr Bree Wilson and Mr Joe Moore for their assistance in the laboratory. Thanks to Ms Kamala Anggamuthu and Ms Kim Maree Kendell for their support and help. Thanks to my wife Mrs Meifang Liu for data input and proofreading, and also for her love and tolerance. Thanks to my parents Mr Yulin Zhu and iv Mrs Lijun Chen, and my daughters Banruo Zhu and Chenxi Zhu for their support and understanding throughout my entire study. Thanks also to Charles Sturt University for providing a scholarship and funding for my research. The EH Graham Centre and the Council of Australasian Weed Societies Inc are acknowledged for their travel support to attend academic conferences. v LIST OF PUBLICATIONS Journal papers Zhu, X. C., Wu, H. W., Raman, H., Lemerle, D., Stanton, R., & Burrows, G. E. (2012). Evaluation of simple sequence repeat (SSR) markers from Solanum crop species for Solanum elaeagnifolium. Weed Research, 52(3), 217-223. doi: 10.1111/j.1365-3180.2012.00908.x. Zhu, X., Raman, H., Wu, H., Lemerle, D., Burrows, G., & Stanton, R. (2013). Development of SSR markers for genetic analysis of silverleaf nightshade (Solanum elaeagnifolium) and related species. Plant Molecular Biology Reporter, 31(1), 248-254. doi: 10.1007/s11105-012-0473-z. Zhu, X. C., Wu, H. W., Stanton, R., Burrows, G. E., Lemerle, D., & Raman, H. (2013). Morphological variation of Solanum elaeagnifolium in south- eastern Australia. Weed Research, doi: 10.1111/wre.12032. Zhu, X. C., Wu, H. W., Raman, H., Lemerle, D., Stanton, R., & Burrows, G. E. (2013). Genetic variation and structure of Solanum elaeagnifolium in Australia analysed by amplified fragment length polymorphism markers. Weed Research, doi: 10.1111/wre.12029. Zhu, X. C., Wu, H. W., Stanton, R., Raman, H., Lemerle, D., & Burrows, G. E. (2013). Time of emergence impacts the growth and reproduction of silverleaf nightshade (Solanum elaeagnifolium Cav.). Weed Biology and Management. doi:10.1111/wbm.12015. vi Zhu, X. C., Wu, H. W., Raman, H., Lemerle, D., Stanton, R., & Burrows, G. E. (2013). SSR marker analysis to determine the genetic variation in Solanum elaeagnifolium in Australia. Plant Protection Quarterly (accepted). Conference proceedings Zhu, X. C., Burrows, G., Wu, H., Raman, H., Stanton, R., & Lemerle, D. (2011). Identification of silverleaf nightshade using microsatellite markers and microstructure. Proceedings of the 23rd Asian-Pacific Weed Science Society Conference, Cairns, pp. 604-609. Zhu, X. C., Wu, H., Stanton, R., Raman, H., Lemerle, D., & Burrows, G. (2012). The impact of emergence time on silverleaf nightshade (Solanum elaeagnifolium) development and growth. Proceedings of the 18th Australasian Weeds Conference, Melbourne, pp. 329-332. Co-authored journal paper Burrows G.E., White R.G., Harper J.D.I., Heady R.D., Stanton R.A., Zhu X., Wu H., Lemerle D. (2013) Entry of fluorescent tracers into leaf trichomes of silverleaf nightshade (Solanum elaeagnifolium Cav.). American Journal of Botany. (requested revision). vii ABSTRACT Silverleaf nightshade (Solanum elaeagnifolium) or SLN is a Weed of National Significance in Australia, mainly infesting the southern cereal cropping zone and with the potential to infest 400 million hectares. SLN management relies on herbicides, but efficacy is influenced by factors such as plant morphology, genetic background, growth stage and environmental conditions. This study investigated the extent and cause of morphological and genetic variation, the distribution of different phenotypes and genotypes, and how the modes of reproduction contribute to the adaptability of SLN in Australia. SLN is a herbaceous, perennial weed that reproduces both sexually and asexually. In Australia, plants germinate from the soil seedbank and rootbank from September (spring) to April (autumn), fruits normally form in January, and the aerial growth senesces in May. High morphological variation was found between 642 SLN individuals from south-eastern Australia. Leaf length, width and area ranged from 1.44 to 10.6 cm, 0.39 to 4.09 cm, and 0.41 to 25.8 cm2, respectively. High trichome densities were found on both leaf surfaces (67 and 132 trichomes/mm2 on the adaxial and abaxial surface, respectively). Larger leaves
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