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Calcium Carbonate Biomineralisation in Disparate Systems - Common Mechanisms?

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Calcium Carbonate Biomineralisation in Disparate Systems - Common Mechanisms? England, Jennifer Katherine (2005) Calcium carbonate biomineralisation in disparate systems - common mechanisms? PhD thesis http://theses.gla.ac.uk/4024/ Copyright and moral rights for this thesis are retained by the author A copy can be downloaded for personal non-commercial research or study, without prior permission or charge This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the Author The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the Author When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given Glasgow Theses Service http://theses.gla.ac.uk/ [email protected] Calcium Carbonate Biomineralisation in Disparate Systems - Common Mechanisms? Jennifer Katherine England A thesis submitted for the degree of Doctor of Philosophy Division of Earth Sciences, Centre for Geosciences, University of Glasgow March 2005 © Jennifer England 2005 Abstract Biominerals are composite materials in which orgaruc components control mineral nucleation and structure. Calcium minerals account for over 50% of biominerals, with calcium carbonate being the most common type. This study considers the extent to which four calcium carbonate biomineral systems share common characteristics. Within the sample set, there is a range of ultrastructures and two types of calcium carbonate polymorph (calcite and aragonite). The mini survey includes three invertebrate systems: two members of the Phylum Brachiopoda; the articulated brachiopod Terebratulina retusa (Subphylum Rhynchonelliformea) and the inarticulated brachiopod Novocrania anomala (Subphylum Craniiformea), and a member of the. Mollusca, the bivalve Mytilus edulis. The fourth, outlying vertebrate system, is the eggshell of the domestic fowl, Gallus gallus. The minor element composition of each of the four systems is considered in the context of mineral ultrastructure. The shell of T. retusa comprises two layers; a primary layer of acicular calcite and an underlying secondary layer composed of calcite fibres. In thin section, a variation between the upper and lower portions of the secondary layer is evident. The concentration of magnesium, sulphur and strontium are significantly greater in the primary layer of the shell. Magnesium concentration also differs between the upper and lower regions of the secondary layer with higher concentration in the upper portion of the secondary layer. The shell of N anomala consists of two layers; a primary layer of acicular calcite and a secondary layer of calcite semi-nacre. N anomala has a high magnesium calcite shell. The concentration of minor elements does not differ significantly between the primary and secondary layers. Two calcium carbonate polymorphs occur in the M edulis shell with an outer calcite layer and an inner aragonite layer. Magnesium concentration is higher in the calcite layer while strontium concentrations are greater in the aragonite layer. Sodium concentration gradually decreases across the calcite layer from the outer surface to the calcite/aragonite boundary and increases in the aragonite layer. The eggshell of G. gallus contains shell membranes, mammillary caps, a palisade layer, a vertical crystal layer and an outer organic cuticle. The concentration of magnesium is high in the mammillary caps, and decreases as the mammillary caps fuse and then gradually increases through the palisade and vertical crystal layers to the outer cuticle. The concentration of phosphorus and potassium is low in the mammillary caps and gradually increases through the shell to reach maximum concentration in the cuticle. Variation in the concentration of minor elements in the shells of T. retusa and N. anomala do not relate to changes in mineral ultrastructure. Differences in shell chemistry between these two brachiopods may be related to differences in physiology. The principal control on the distribution of minor elements in M edulis is crystal structure. In G. gallus the concentration of minor elements changes as ultrastructure changes. However, ultrastructure is unlikely to be the main control on shell chemistry, as abrupt changes in shell ultrastructure contrast to gradual changes in element distribution throughout the shell. While there may be similarities in the mechanisms controlling the minor element composition between some systems e.g. the extent to which some organisms control the ionic composition of the mineralising medium, there does not appear to be a common principal mechanism that governs the chemical composition of these four biominerals. To some extent, more unity is evident in the biochemical characteristics of the organic matrix that are common to the four systems. The soluble organic matrices of the shells all contain small, acidic proteins. The amino acid composition of the four systems also displays some similarities such as a high glycine concentration. In each case, sulphated sugars are also associated with the soluble organic matrix. Closer examination of the organic components reveals that the protein profiles are different in each of the four systems. ii Acknowledgments I am extremely grateful to the Engineering and Physical Sciences Research Council for funding for this three year studentship (Grant number GRIR23107/01). I would firstly like to thank Dr. Maggie Cusack for her help, support and guidance over the past three years and Dr. Martin Lee for his advice and encouragement. I am indebted to Sandra Tierney who has not only provided me with exceptional technical knowledge but has been a great friend. A special thankyou is required for Robert McDonald for his assistance with EPMA and SEM, Kenny Roberts for his help with IT, Bill Higgison for assistance with XRD and John Gilleece for preparation of samples. Many thanks to Eddie for driving myself and Big Dave to Oban to collect brachiopods. I am also grateful to the crew of the RV Calanus for their assistance with brachiopod collection and making me welcome on board. I would also like to thank Dr Rob Martin and Dr Paul Edwards from the Department of Physics, University ofStrathclyde for their advice on CL spectroscopy. I would like to thank the late Sir Alwyn Williams FRS, FRSE for his help and guidance with interpretation of brachiopod shell structure. I am extremely grateful to all the postgraduates for keeping me sane, in particular to Sarah and Davie for being good friends over the last three years. A special thankyou goes to Liz Campbell for her friendship and enthusiasm for running 10K races and to Big Dave for being a bad influence (RC now). Thanks also to Wolfie for being a great friend and listener. Finally and most importantly I would like to thank my Mum, Dad and Bob for their love, support and encouragement. iii Declaration The material presented in this thesis summarises the results of three years of independent research carried out in the Division of Earth Sciences, University of Glasgow. The research was supervised by Dr. Maggie Cusack (University of Glasgow) and Dr. Martin Lee (University of Glasgow). This thesis is a result of my own research and any published or unpublished work of other researchers has been given full acknowledgment in the text. Jennifer K. England March 2005 iv Table of Contents Page Abstract i Acknowledgments iii Declaration iv Table of contents v List of Figures ix List of Tables xiv Chapter 1 Introduction 1.1 Introduction 1 1.2 Biominerals 1 1.2.1 Inorganic Composition 2 1.2.2 Organic Composition 3 1.2.3 Biomineralisation Mechanisms 4 1.3 Emergence and Early Evolution of Biominerals 5 1.4 Aims of the Study 6 1.5 The Four Biomineral Systems 7 1.5.1 Terebratulina retusa 7 1.5.2 Novocrania anomala 8 1.5.3 MytiJus edulis 9 1.5.4 Avian Eggshell (Gallus gallus) 10 Chapter 2 Materials and Methods 2.1 Materials 12 2.2 Sample Collection and Preparation 12 2.3 X-ray Diffraction (XRD) 13 2.4 Examination of Shell Ultrastructure 13 2.5 Thin Section Preparation 13 2.6 Minor Element Analysis 14 2.6.1 Sample Preparation 14 2.6.2 Electron Microprobe Analysis 14 2.6.3 Element Mapping 16 2.6.4 Cathodoluminescence (CL) Microscopy 17 2.6.5 Cathodoluminescence (CL) Spectroscopy 17 v 2.7 Determination of Total Organic Concentration and Distribution 17 2. 7.1 Loss On Ignition 17 2. 7.2 Scanning Electron Microscope Backscatter (BSE) Imaging 18 2.8 Extraction, purification and visualisation of shell protein 18 2.8.1 Extraction ofintercrystalline protein 18 2.8.2 Extraction ofintracrystalline protein 18 2.8.3 Purification ofintercrystalline and intracrystalline protein 19 2.8.4 Sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS PAGE) 19 2.8.5 Coomassie Brilliant Blue Staining 22 2.8.6 Silver Staining 22 2.8.7 Acridine Orange Staining 22 2.88 Isoelectric Focusing (IEF) 22 2.9 Amino Acid Analysis 23 2.9.1 Manual Hydrolysis 24 2.9.2 Amino Acid Analysis 24 2.9.3 Summary ofSteps for Amino Acid Analysis 24 2.10 N-terminal Sequencing 25 2.10.1 Transfer ofProtein to Problott Membrane 25 2.10.2 N-terminal Sequencing 25 2.11 Calcium Carbonate Crystal Growth 26 2.11.1 Kitano Protocol 26 2.11.2 Calcite Grown in the Presence ofProtein Extract 27 Chapter 3 Ultrastructure and Minor Element Concentration 3.1 Introduction 28 3.2 Previous Work 29 3.2.1 Magnesium 29 3.2.2 Strontium 31 3.2.3 Cathodoluminescence 32 3.3 Results 34 3.3.1 Terebratulina retusa Ultrastructure and Minor Elements 34 3.3.1.1 Ultrastructure 34 3.3.1.2 Minor Element Composition-Electron
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