Cusack 2002-PhD Thesis-Diatom Pseudo-nitzschia in

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CHAPTER 1
GENERAL INTRODUCTION 1 INTRODUCTION This project is concerned specifically with Pseudo-nitzschia, a genus of the marine diatoms, class Bacillariophyceae (Haeckel 1878 as cited in Skov et al. 1999). Pseudonitzschia is cosmopolitan having a worldwide distribution (Cupp 1943; Hasle 1965, 1972, Hasle et al.1996; Villac et al. 1993; Hallegraeff 1994; Fryxell et al. 1997) and has become an attractive choice for research by many scientists in recent years, for the simple reason that some species within the genus can produce a toxin (domoic acid) responsible for Amnesic Shellfish Poisoning in humans. More than ten classes of unicellular algae contribute to marine phytoplankton. Of these, the diatoms (Bacillariophyceae) are one of the most common, representing a large component of the microalgal flora, with approximately 1,500 planktonic (drifting) diatom species in the sea (Sournia et al. 1991). Diatoms can assume either solitary or colonial forms and typical photosynthetic pigments include chlorophyll a, c, ß–carotene, fucoxanthin, diatoxanthin and diadinoxanthin. Diatoms characteristically have a rigid cell wall that is chiefly composed of silica (silicon dioxide). It is the symmetry of this heavy siliceous exoskeleton, referred to as the frustule that divides diatoms into the Centrales (Centric diatoms) and the Pennales (pennate diatoms) following Karsten (1928). Pennate diatoms are further separated into two subgroups, the araphid pennate diatoms that possess a sternum, and the raphid pennate diatoms in which both a sternum and raphe † are present. Pseudo-nitzschia is placed within the pennate diatoms (elongated rod shaped diatoms). As a diatom, the frustule of Pseudo-nitzschia consists of two morphologically composite units (opposing surfaces) called valves, each connected by adjoining structures referred to as girdle elements. The older valve with its girdle elements (epitheca) fits over the younger valve (hypotheca), comparable to the way a “pill box” fits together (Figure 1a). Under inspection Pseudo-nitzschia can present two views, the vale view or the girdle view, and its symmetry is based around a central line (Figure 1b). The three planes of symmetry of † A glossary of all terms used may be found in Appendix 1.
Page 1 and 2: THE DIATOM PSEUDO-NITZSCHIA (PERAGA
Page 3 and 4: i TABLE OF CONTENTS CONTENTS PAGE T
Page 5 and 6: iii TABLE OF CONTENTS TABLES PAGE C
Page 7 and 8: v TABLE OF CONTENTS FIGURES PAGE CH
Page 9 and 10: vii TABLE OF CONTENTS FIGURES PAGE
Page 11 and 12: ix TABLE OF CONTENTS PLATES PAGE CH
Page 13 and 14: ACKNOWLEDGMENTS xi ACKNOWLEDGMENTS
Page 15: OBJECTIVES OF THIS STUDY xiii OBJEC
Page 19 and 20: 3 INTRODUCTION Figure 2. Axis and p
Page 21 and 22: 5 INTRODUCTION Table 1. Nomenclatur
Page 23 and 24: 1.2 TAXONOMY 1.2.1 MORPHOLOGY 7 INT
Page 25 and 26: 9 INTRODUCTION Figure 3. Diagram sh
Page 27 and 28: 11 INTRODUCTION Table 3. Distinct f
Page 29 and 30: 13 INTRODUCTION Table 5. Additional
Page 31 and 32: 15 INTRODUCTION Table 7. Features o
Page 33 and 34: 17 INTRODUCTION Figure 5. Diagram s
Page 35 and 36: 19 INTRODUCTION other domains are h
Page 37 and 38: 21 INTRODUCTION using more conventi
Page 39 and 40: Table 8. Global distribution of spe
Page 41 and 42: 25 INTRODUCTION 1.4 DOMOIC ACID AND
Page 43 and 44: 27 INTRODUCTION domoic acid (DA) wa
Page 45 and 46: 29 INTRODUCTION seven Pseudo-nitzsc
Page 47 and 48: 1.4.1 DOMOIC ACID 31 INTRODUCTION D
Page 49 and 50: 33 INTRODUCTION One of the first ba
Page 51 and 52: 35 INTRODUCTION Initially, a lot of
Page 53 and 54: 37 INTRODUCTION P. multiseries was
Page 55 and 56: 39 INTRODUCTION pungens f. multiser
Page 57 and 58: 41 INTRODUCTION Dickman, M. and Gle
Page 59 and 60: 43 INTRODUCTION Hallegraeff, G.M. A
Page 61 and 62: 45 INTRODUCTION Impellizzeri, G., M
Page 63 and 64: 47 INTRODUCTION Mann, D. G. 1978. S
Page 65 and 66: 49 INTRODUCTION Reap, M.E. 1991. Ni
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51 INTRODUCTION Phytoplankton, Leaf
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53 INTRODUCTION Tryphonas, L., True
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CHAPTER 2
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ABSTRACT 56 CHAPTER 2 The presence
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58 CHAPTER 2 offshore. Occasional s
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Latitude (N) 57 56 55 54 53 52 51 E
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62 CHAPTER 2 Table 1. Morphometric
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Plate 1. Pseudo-nitzschia pungens (
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Plate 3. Pseudo-nitzschia australis
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Plate 4. Pseudo-nitzschia fraudulen
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Plate 5. Pseudo-nitzschia delicatis
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Plate 6. Pseudo-nitzschia pseudodel
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Plate 7. Pseudo-nitzschia cf. seria
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76 CHAPTER 2 Spatial and Temporal d
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Latitude (N) Latitude (N) 55.0 54.0
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80 CHAPTER 2 if the specimens exami
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82 CHAPTER 2 over P. multiseries le
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84 CHAPTER 2 can be mixed across th
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86 CHAPTER 2 Hasle, G.R. 1993. Nome
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CHAPTER 3
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ABSTRACT 89 CHAPTER 3 The presence
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91 CHAPTER 3 In the Bay of Fundy (E
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Latitude (N) Latitude (N) 53.0 52.5
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Latidtude (N) 53.0 52.5 52.0 51.5 M
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97 CHAPTER 3 Quantitative phytoplan
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h (m) 0 50 100 150 200 250 0 50 100
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101 CHAPTER 3 1313 (12m), “P. ser
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103 CHAPTER 3 Figure 8. Distributio
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cells.mL -1 105 Φ (J.m -3 ) 0 50 1
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107 CHAPTER 3 up to a maximum of 7.
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109 CHAPTER 3 Figure 13. Vertical p
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113 CHAPTER 3 The “P. delicatissi
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Salinity Salinity Salinity 35.5 35.
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119 CHAPTER 3 Figure 22. Temperatur
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h (m) 0 50 100 150 200 0 50 100 150
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123 CHAPTER 3 layer were low, betwe
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125 CHAPTER 3 Figure 27. Distributi
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127 CHAPTER 3 Table 1. Ranges in th
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129 CHAPTER 3 Figure 30. Areas of a
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Salinity 36.0 35.5 35.0 34.5 34.0 3
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DISCUSSION 133 CHAPTER 3 Pseudo-nit
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135 CHAPTER 3 greater diversity and
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137 CHAPTER 3 spp. have a tendency
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139 CHAPTER 3 thermocline is genera
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141 CHAPTER 3 are solely based on t
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REFERENCES 143 CHAPTER 3 Bates, S.S
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145 CHAPTER 3 genus Pseudo-nitzschi
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147 CHAPTER 3 Míguez, Á., Fernán
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149 CHAPTER 3 Roden, C.M., Lennon,
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151 CHAPTER 3 Walz, P.M., Garrison,
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152 CHAPTER 4 CONFIRMATION OF DOMOI
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INTRODUCTION 154 CHAPTER 4 The penn
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156 CHAPTER 4 for the unknown and a
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158 CHAPTER 4 with nebulizer-assist
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160 CHAPTER 4 Cultures 1 - 4 (under
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162 CHAPTER 4 Plate 1. Pseudo-nitzs
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Cells•mL -1 (x 10 3 ) 1000 100 10
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DISCUSSION 166 CHAPTER 4 Diatoms of
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168 CHAPTER 4 many studies have sho
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170 CHAPTER 4 Bates, S. S., Garriso
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172 CHAPTER 4 Hasle, G. R. 1965. Ni
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174 CHAPTER 4 Roden, C. M., Lennon,
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CHAPTER 5
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ABSTRACT 177 CHAPTER 5 Ribosomal DN
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179 CHAPTER 5 fact, in New Zealand,
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181 CHAPTER 5 Gel purification of t
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183 CHAPTER 5 Maximum-Parsimony tre
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185 CHAPTER 5 Table 3. LSU rRNA-tar
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Phylogenetic analysis 187 CHAPTER 5
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189 CHAPTER 5 Plate 1. Scanning ele
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Figure 1 continued. 191 CHAPTER 5
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193 CHAPTER 5 Figure 1 continued. 1
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195 CHAPTER 5 Figure 3. Phylogeneti
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197 CHAPTER 5 Application of rRNA-t
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DISCUSSION 199 CHAPTER 5 Since trad
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201 CHAPTER 5 probes (Scholin et al
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203 CHAPTER 5 The fluorescently lab
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205 CHAPTER 5 If the probes are to
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207 CHAPTER 5 Maidak B. L., Cole J.
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CONCLUDING REMARKS
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210 CONCLUDING REMARKS species were
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212 CONCLUDING REMARKS eastern Prin
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214 CONCLUDING REMARKS California.
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APPENDIX 1
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216 TERMINOLOGY TERMINOLOGY IN ACCO
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218 TERMINOLOGY TERMINOLOGY IN ACCO
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APPENDIX 2
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Pseudo-nitzschia pungens (Robust: v
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Pseudo-nitzschia multiseries (Robus
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Pseudo-nitzschia seriata f. seriata
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Pseudo-nitzschia seriata f. obtusa
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229 Pseudo-nitzschia australis g) T
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231 Pseudo-nitzschia fraudulenta th
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ELECTRON MICROSCOPY (EM) ACID CLEAN
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235 Pseudo-nitzschia heimii f) A ce
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237 Pseudo-nitzschia delicatissima
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239 Pseudo-nitzschia pseudodelicati
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Appendix III 240 APPENDIX III Micro
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Figs. 1-18 LIGHT MICROGRAPHS (LM).
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Figs. 1-18 Cont'd. Figs.1-17 Cont'd
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Figs. 19-40 Cont'd. Figs. 26-30 Con
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Fig. 41-55 TRANSMISSION ELECTRON MI
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Pseudo-nitzschia australis (Figures
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Figs. 56-86 Cont'd. LIGHT MICROGRAP
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Figs. 87-105 SCANNING ELECTRON MICR
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Figs. 106-154 TRANSMISSION ELECTRON
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Figs. 155-168 LIGHT MICROGRAPHS (LM
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270 Pseudo-nitzschia fraudulenta Fi
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Figs. 169-210 Cont'd. Figs. 178-181
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Figs. 169-210 Cont'd. 274 Pseudo-ni
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Figs. 199-226 Cont'd. TRANSMISSION
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Figs. 228-236 SCANNING ELECTRON MIC
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Pseudo-nitzschia delicatissima (Fig
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Figs. 237-247 Cont'd. LIGHT MICROGR
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Figs. 335-351 Cont'd. SCANNING ELEC
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304 Pseudo-nitzschia cf. multiserie
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Pseudo-nitzschia cf. seriata (Figur
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Appendix IV 311 APPENDIX IV A list
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313 CLEANING METHODS activity. Clea
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CHEMICAL CLEANING 315 CLEANING METH
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HYDROCHLORIC ACID AND POTASSIUM PER
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319 CLEANING METHODS Several ranges
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321 CLEANING METHODS PREPARATION OF
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Appendix V 322 APPENDIX V Short des
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324 PHYLOGENETICS pyrimidines, C an
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326 PHYLOGENETICS Figure 1. Multipl
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Figure 1 continued. 328 PHYLOGENETI
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332 PHYLOGENETICS Figure 2. Multipl
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Figure 2 continued. REFERENCES 340
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APPENDIX 6
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243 Pseudo-nitzschia pungens Cell n
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245 Pseudo-nitzschia pungens No. of
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347 Pseudo-nitzschia cf. seriata f.
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349 Pseudo-nitzschia australis Cell
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351 Pseudo-nitzschia australis Cell
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353 Pseudo-nitzschia fraudulenta Ce
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355 Pseudo-nitzschia cf. subpacific
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357 Pseudo-nitzschia delicatissima
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APPENDIX 7
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360 STATION DATA Table 1. Station p
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366 STATION DATA Table 7. Values fo
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368 STATION DATA Table 9. Values fo
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370 STATION DATA Table 11. Nutrient
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376 STATION DATA Figure 1. Vertical
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378 STATION DATA Table 17. Pseudo-n
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380 STATION DATA Table 21. Pseudo-n
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