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Durham E-Theses Durham E-Theses Biological studies on the rivulariaceae Kirkby, Susan M. How to cite: Kirkby, Susan M. (1975) Biological studies on the rivulariaceae, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/8282/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk BIOLOGICAL STUDIES ON THE RIVULARIACEAE By Susan M. Kirkby (B.Sc. Dunelm) A thesis submitted for the degree of Doctor of Philosophy in the University of Durham, England. August, 1975 4 NOV This thesis, which is entirely the result of my own work, has not been accepted for any degree, and is not being submitted concurrently in canditature for any .-other degree. ACKNOWLEDGMENTS I wish co acknowledge, with thanks, all those persons who have assisted me in the course of this project. Firstly I wish to thank Dr B.A. Whitton, my supervisor, for help and advice given throughout this work and for suggesting the project. I am grateful to all those people who have helped in my search for Rivularia. Thanks are due to Dr M.E. Bradshaw, Dr D.J. Bellamy, Dr M.K. Hughes, P.F. Williams, N.T.H. Holmes and Dr B. Bell. Thanks are due also to Miss C. Sinclair and A. Donaldson who have provided stimulating and useful discussions of the research topic, Mr W. Simon for technical assistance, Mr D.G. Middlemass for maintaining the growth tanks, Mr T. Brett, Mrs G.A. Walker and E.J.H. Lloyd for help with analysis of the water samples. 1 am grateful to Professor D. Boulter for allowing me to carry out my research in his department and to Durham University for financial support during this work. Finally I wish to thank the many people and in particular my husband and parents, whose constant encouragement has ensured the completion of this work. ABSTRACT This thesis reports a study on the variation of two genera of blue-green algae, Calothrix and Rivularia. The aspects chosen for detailed study were tapering and the hair. Before any comparative work could be undertaken it was necessary to develop ways of describing, in a quantitative manner, changes in the tapering and hair length. Tapering proved to be complex to describe in a satisfactory manner but four simple indices have been suggested. Measurements of individual trichomes have shown problems inherent in detailed studies of tapering, and the indices suggested by the author have been discussed in relation to this. Experimental studies on tapering have been concerned with elaborating the observations of previous authors that, in the presence of combined nitrogen, Calothrix and Gloeotrichia did not develop their characteristic taper. The effect of combined nitrogen on morphology was studied using the tapering indices. Attempts to culture material with hairs proved difficult; however a few factors which affect the development of the hair have been described. A computer program was developed in order to summarise relationships between the occurrence of hairs and other morphological features which have been described in the literature. For practical reasons it was often necessary to assign a name to members of the genus Rivularia; however .in doing this many problems were encountered. To overcome some of these difficulties an objective method of identification was devised. This was used in particular to identify material collected in the field. A brief ecological survey of Rivularia was undertaken in order to study possible correlations between certain aspects of water chemistry and the occurrence of members of the genus. -1- CONTENTS page Acknowledgments Abstract List of Tables 6 List of Figures 8 1 INTRODUCTION 10 1.1 Introduction to the family Rivulariaceae 10 1.2 Morphological variation in the Rivulariaceae 11 1.21 Differentiation of the filament 15 1.22 Variation along the filament 19 1.23 Morphological variation in response to environmental conditions 21 1.24 Taxonomic implications of morphological variation 22 1.25 Physiological aspects of morphological variation 24 1.26 Nitrogen fixation studies 25 1.3 Tapering and the hair 28 1.31 Tapering 29 1.32 Hair formation 30 1.33 Meristerns and development 33 1.4 Ecology of the Rivulariaceae 34 1.41 Introduction 34 1.42 Epilithic freshwater forms 34 1.43 Marine forms 35 1.44 Planktonic forms 37 -2- page 2 MATERIALS AND METHODS 41 2.1 List of abbreviations used in text 41 2.2 Algal materials 41 2.3 Apparatus 44 2.4 Chemicals 45 2.5 Media 45 2.6 Culture techniques 49 2.61 Cleaning of glassware 49 2.62 Sterilisation 50 2.63 Standard growth conditions 50 2.64 Inoculum material 52 2.7 Measurement of growth and morphological variation 52 2.71 Dry weight 52 2.72 Trichome dimensions and morphological variation 53 2.8 Chemical analyses 54 2.9 Acetylene reduction 55 2.10 Collection of field data 55 2.11 Computing 60 2.12 Statistics 62 3 TAPERING AND THE HAIR 71 3.1 Introduction 71 3.2 The concept of tapering 72 3.21 Indices of tapering 72 3.22 Detailed measurements of individual trichomas 91 3.3 The hair 100 3.4 Hairs in laboratory culture 118 3.5 Discussion 134 -3- page 4 PHYSIOLOGICAL ASPECTS OF TAPERING 138 4.1 Introduction 138 4.2 The effect of NH^-N on the growth and morphology of two strains of Calothrix 138 4.21 Calothrix brevissima 138 4.22 Calothrix viguieri 140 4.3 The effect of NO^-N on the growth and morphology of two strains of Calothrix 149 4.31 Calothrix viguieri 149 4.32 Calothrix scopulorum 153 4.4 Discussion 168 5 THE DEVELOPMENT OF AN OBJECTIVE SIMULTANEOUS KEY FOR THE IDENTIFICATION OF CALOTHRIX AND RIVULARIA 172 5.1 Introduction 172 5.2 Development of a standard recording procedure 173 5.21 Standard recording procedure for describing the genera Calothrix and Rivularia 175 5.22 Guide to the standard recording procedure 181 5.23 Standard recording procedure applied to field material 186 5.3 Coding the data 189 5.4 Computer analysis of the data 190 5.41 Identification 190 5.421 Keying-down known species 193 5.412 Keying-down unknown material 199 5.42 Answering data queries 204 5.421 Tapering and the hair 204 5.422 Physiognomic form and the hair 207 -4- page 5.423 Swollen base and the hair 209 5.424 Sheath colour and the hair 209 5.5 Discussion 211 6 AN ECOLOGICAL STUDY OF RIVULARIA 215 6.1 Introduction 215 6.2 Site descriptions 217 6.21 Sampling sites near Inchnadamph,Sutherland 21? 6.22 Additional.Scottish sampling sites 217 6.23 North Pennines and Upper Teesdale 219 6.24 Westmorland 219 6.25 Malham region, West Yorkshire 221 6.26 South Wales 223 6.3 Water chemistry data 223 6.31 Comparison of water chemistries from + and - Rivularia sites 231 6.4 Distribution of Rivularia in terms of substratum and water depth 232 6.5 Discussion 233 7 DISCUSSION 239 7.1 Introductory comments 239 7.2 Tapering 240 7.3 The hair 242 7.4 The effect of combined nitrogen on tapering 242 7.5 Morphological variation and the implications 245 on taxonomy 7.6 The effect of certain factors on the development 247 of the hair 7.7 Problems associated with identifying Calothrix 249 and Rivularia -5- 7.8 The ecological significance of Rivularia 251 7.9 Concluding remarks 253 256 SUMMARY 260 REFERENCES APPENDIX I Authorities for Rivulariaceae included in 286 this work APPENDIX II Information on the morphology of Calothrix 292 and Rivularia APPENDIX III Literature related to the distribution of Rivulariaceae 337 APPENDIX IV Guide to computer records of Rivularia 348 samples Field data and trichome descriptions 350 APPENDIX V Published paper: Pleurocladia lacustris A. Braun (Phaeophyta)- a new British record. -6- List of Tables page Table 1.1 Descriptions of genera of the Rivulariaceae 12 Table 1.2 Nitrogen fixation in the Rivulariaceae 26 Table 2.1 Details of cultures 43 Table 2.2 Media composition 46 Table 2.3 Optimum growth conditions of cultures 51 Table 2.4 Copy of 1 FIELD1 program 64 Table 2.5 Copy of 'IDENTIFY' program 66 Table 2.6 Copy of 'QUESTION' program 69 Table 3.1 Trichome dimensions 81 Table 3.2 Tapering values 82 Table 3.3 Tapering order 83 Table 3.4 Taperingvalues (based on Fig. 3.5) 101 Table 3.5 Tapering order (based on Fig. 3.5) 102 Table 3.6 External surface area and volume measurements 103 Table 3.7 External surface area: volume relationships 117 Table 3.8 Occurrence of hairs in cultures of Calothrix 131 Table 4.1 The effect of initial NH^-N on the subsequent growth and morphology of Calothrix viguieri 144 Table 4.2 Basal and apical widths of Calothrix viguieri cultured in + NO^-N media and - NO^-N media 154 Table 4.3 Change in tapering of Calothrix viguieri cultured in + NO-N and - N0.J-N media 155 Table 4.4 Acetylene reduction assay of Calothrix viguieri cultured in - NO^-N medium 157 Table 4.5 Acetylene reduction assay of Calothrix viguieri cultured in + NO^-N medium 158 Table 4.6 Basal and apical widths of Calothrix scopulorum cultured in + N03~N media and - NO^-N media 163 Table 4.7 Changes in tapering of Calothrix scopulorum cultured in + NO-N and - NO-N media 164 -7- page Table
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