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Tarantulas and Social Spiders

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Tarantulas and Social Spiders Tarantulas and Social Spiders: A Tale of Sex and Silk by Jonathan Bull BSc (Hons) MSc ICL Thesis Presented to the Institute of Biology of The University of Nottingham in Partial Fulfilment of the Requirements for the Degree of Doctor of Philosophy The University of Nottingham May 2012 DEDICATION To my parents… …because they both said to dedicate it to the other… I dedicate it to both ii ACKNOWLEDGEMENTS First and foremost I would like to thank my supervisor Dr Sara Goodacre for her guidance and support. I am also hugely endebted to Dr Keith Spriggs who became my mentor in the field of RNA and without whom my understanding of the field would have been but a fraction of what it is now. Particular thanks go to Professor John Brookfield, an expert in the field of biological statistics and data retrieval. Likewise with Dr Susan Liddell for her proteomics assistance, a truly remarkable individual on par with Professor Brookfield in being able to simplify even the most complex techniques and analyses. Finally, I would really like to thank Janet Beccaloni for her time and resources at the Natural History Museum, London, permitting me access to the collections therein; ten years on and still a delight. Finally, amongst the greats, Alexander ‘Sasha’ Kondrashov… a true inspiration. I would also like to express my gratitude to those who, although may not have directly contributed, should not be forgotten due to their continued assistance and considerate nature: Dr Chris Wade (five straight hours of help was not uncommon!), Sue Buxton (direct to my bench creepy crawlies), Sheila Keeble (ventures and cleans where others dare not), Alice Young (read/checked my thesis and overcame her arachnophobia!) and all those in the Centre for Biomolecular Sciences. But above all, the greatest thanks and love go to my parents for their endless love, support and of course finance through all the hard times … iii ABSTRACT Studies of spider silks indicate that they may outperform virtually all synthetic fibres in terms of strength, elasticity and toughness. To date, most silks studied come from only a select few species and likely underrepresent the immense diversity of the clades. Here, protein and mRNA sequence analyses were used to study silk from two types of spider. The first approach used ESI tandem mass spectrometry to sequence peptide fragments of a silk from a tarantula (Mygalomorphae, Theraphosidae), a hitherto neglected family. The results confirm that the common silk types found in araneomorph spiders, Spidroin 1 and Spidroin 2, are also found in mygalomorphs. A putative N- terminal domain that bears a striking similarity to the N-terminus of araneomorph pyriform silk was isolated. If correctly identified, this would be the first ever recorded N-terminal domain for a mygalomorph. The second approach taken was to construct a cDNA library from theraphosid silk glands and adjacent tissue. Sequencing identified a significant number of uniquely truncated rRNAs. These may be the result of specific 'fragile sites' within these transcripts, which would explain the discrete classes of length polymorphisms found. The cDNA library sequences also provided evidence consistent with RNA editing and furthermore identified the presence of both transcribed nuclear pseudogenes and transposable elements. These may reflect past evolutionary horizontal gene transfer events within the spider genome. Similar analysis of next generation sequencing data from the transcriptomes of three Stegodyphus spp. (Araneomorphae) reveal a range of apparent silk types with similarity to major ampullate, minor ampullate and pyriform silks. These were identified by searching for comparative sequence homologies using Microsoft Office Word. No flagelliform silk or recognisable sticky silks were identified, which is consistent with the biology of Stegodyphus species. In addition to studies of silk, previous common conceptions of dimensional morphologies were examined to see if they could adequately sex theraphosid spiders, including the species that was the subject of the silk study already described. An independent samples t-test was conducted to compare morphologies of particular leg hairs and statistical analysis demonstrated that there were significant differences between males and females (t (70) = 9.445, p < .001). This technique may be important in future evolutionary and ecological studies of theraphosids. Keywords: Silk, major ampullate spidroin, dragline, tarantula, transcriptomics, proteomics, spines, cDNA library iv TABLE OF CONTENTS DEDICATION .......................................................................................................................... ii ACKNOWLEDGEMENTS .................................................................................................... iii ABSTRACT.............................................................................................................................. iv TABLE OF CONTENTS .......................................................................................................... v LIST OF FIGURES .................................................................................................................. ix LIST OF TABLES ................................................................................................................ xiii GLOSSARY ............................................................................................................................. xv 1 INTRODUCTION ............................................................................................... 1 1.1 The origins of spider silk ............................................................................................... 1 1.2 The history of silkworm silk .......................................................................................... 2 1.3 Other insects that produce silk ..................................................................................... 3 1.4 Spider distribution and the use of silk for aerial dispersal ............................................ 5 1.5 Spider silk ..................................................................................................................... 7 1.6 Silk protein macrostructure .......................................................................................... 9 1.7 Silk evolution ............................................................................................................. 15 1.8 Protein constraints ..................................................................................................... 19 1.9 Glands ........................................................................................................................ 21 1.9.1 Cribellum and calamistrum ............................................................................... 22 1.9.2 Cylindriform glands ........................................................................................... 24 1.9.3 Pyriform glands ................................................................................................. 24 1.9.4 Ampullate glands (major and minor) ................................................................. 24 1.9.4.1 Major ampullate glands ........................................................................... 24 1.9.4.2 Minor ampullate glands ........................................................................... 25 1.9.4.3 Dragline silk ............................................................................................. 25 1.9.5 Flagelliform glands ............................................................................................ 26 1.9.6 Aciniform glands ............................................................................................... 27 1.9.7 Aggregate glands ............................................................................................... 27 1.10 Nucleotide/genetic structure of silk genes ............................................................. 30 1.11 Silk producing conditions in the spider .................................................................. 32 1.12 In vitro production of synthetic silk ....................................................................... 33 1.13 Other properties of silk .......................................................................................... 36 1.14 Industrial applications and problems with silk ....................................................... 38 1.15 Uses and potential uses of silk ............................................................................... 41 1.16 Orb-webs ............................................................................................................... 43 1.17 Tarantula silk ......................................................................................................... 46 1.18 Tarsal silk ............................................................................................................... 50 1.19 Aims ...................................................................................................................... 52 2 MATERIALS AND METHODS ..................................................................... 53 2.1 Materials .................................................................................................................... 53 2.1.1 Chemicals and reagents .................................................................................... 53 2.2 Enzymes and kits .......................................................................................................
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