DOCSLIB.ORG

8.04 Biomineralization H

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
8.04 Biomineralization H 8.04 Biomineralization H. C. W. Skinner Yale University, New Haven, CT, USA and A. H. Jahren Johns Hopkins University, Baltimore, MD, USA 8.04.1 INTRODUCTION 118 8.04.1.1 Outline of the Chapter 118 8.04.1.2 Definitions and General Background on Biomineralization 119 8.04.2 BIOMINERALS 120 8.04.2.1 Calcium Carbonates 120 8.04.2.1.1 Calcite 120 8.04.2.1.2 Aragonite 122 8.04.2.1.3 Vaterite 124 8.04.2.2 Silica 124 8.04.2.2.1 Opal 124 8.04.2.3 Bioapatite 125 8.04.2.4 Iron Oxides and Hydroxides 127 8.04.2.4.1 Magnetite 127 8.04.3 EXAMPLES OF BIOMINERALIZATION 129 8.04.3.1 Introduction 129 8.04.3.2 Sulfur Biomineralization 129 8.04.3.2.1 Sulfur oxidizers 130 8.04.3.2.2 Sulfate reducers 130 8.04.3.2.3 Formation of elemental sulfur 130 8.04.3.2.4 Sulfate biomineralization 131 8.04.3.3 Iron Biomineralization 131 8.04.3.3.1 Roles of iron in archea and bacteria 131 8.04.3.3.2 Bacterial iron mineral formation 132 8.04.3.3.3 Magnetotactic bacteria 132 8.04.3.4 Carbonate Biomineralization 134 8.04.3.4.1 Cyanobacteria 134 8.04.3.4.2 Cnidaria (coelenterates) 136 8.04.3.4.3 Coccoliths 137 8.04.3.4.4 Foraminifera 139 8.04.3.4.5 Echinoids 140 8.04.3.4.6 Mollusks 141 8.04.3.4.7 Arthropods 144 8.04.3.5 Silica Biomineralization 145 8.04.3.5.1 Radiolarians 145 8.04.3.5.2 Diatoms 146 8.04.3.5.3 Sponges 150 8.04.3.6 Plant Biomineralization 152 8.04.3.6.1 Introduction 152 8.04.3.6.2 Plant biominerals 152 8.04.3.6.3 Phytoliths: indicators of the environment and paleoenvironment 154 8.04.3.6.4 Funghi and lichen biomineralization 158 8.04.3.7 Vertebrate Biomineralization 158 8.04.3.7.1 Introduction 158 117 118 Biomineralization 8.04.3.7.2 Bones and bone tissues 159 8.04.3.7.3 Cartilage 165 8.04.3.7.4 Antlers 167 8.04.3.7.5 Teeth 168 8.04.3.7.6 Otoliths 171 8.04.4 SUMMARY: WHY BIOMINERALIZE? 172 8.04.4.1 Physical or Macrobiomineralization Contributions 174 8.04.4.1.1 Defense and protection through biomineralization 174 8.04.4.2 Chemical or Microbiomineralization Contributions 174 ACKNOWLEDGMENTS 175 REFERENCES 175 8.04.1 INTRODUCTION will continue to do so, an essential ingredient for establishing and maintaining Earth’s environment Biomineralization is the process by which in the future as in the past (Figure 1). living forms influence the precipitation of mineral materials. The process creates heterogeneous accumulations, composites composed of biologic 8.04.1.1 Outline of the Chapter (or organic) and inorganic compounds, with inhomogeneous distributions that reflect the It is only with the advent of more sensitive, environment in which they form. The products higher-resolution techniques that we can identify are, however, disequilibrium assemblages, created the exact mineral components, and appreciate the and maintained during life by dynamic meta- precision control exercised by the life forms on bolism and which, on death, may retain some of their mineralized structures, and the potential that the original characteristics. The living forms their formation and evolution are responses to the discussed in this chapter produce carbonate, environment and climatic, or any other pervasive phosphate, oxalate, silica, iron, or sulfur-contain- geochemical changes. We first present the ing minerals illustrating the remarkable range of crystal chemistry, or mineralogy, of some of the biomineralization chemistries and mechanisms. common minerals that are encountered in Biomineralization, in the broadest use of the term, biomineralization (Section 8.04.2)andthen has played a role in Earth cycles since water provide examples from the range of life forms. appeared on the surface. The general perception We start with the Archea and Bacteria although that the onset of biomineralization coincides with we are just beginning to investigate their the appearance of fossils that left “hard parts” mineralization processes. However, these primi- amenable to analysis is marked geologically as the tive forms were probably the first to generate the dawn of the Cambrian. At least for some of us, the mechanisms that led to the accumulation of origins of life, and possibly biomineralization, elements, the precursor to the formation of go back 3.8 Gyr. The startling chemical and bio- biominerals. The details of the mechanisms and logical range encompassed by the term biominera- the ranges of possible creatures in these classes lization implies that life forms have adapted and remain to be fully defined and understood. altered geoenvironments from the beginning, and However, we consider them an essential base to Figure 1 The “tree of life” (source Raven et al., 1999, figure 13.8, p. 270). Introduction 119 any biomineralization discussions. Further, and calcite formation for the group of algae the perhaps most intriguing, many other biominera- coccolithophoridae (Section 8.04.3.4.3). Another lizing forms incorporate them as symbionts. illustration of the diversity of intracellular bio- Following these opening sections, we move mineralization is found in the freshwater green onto CaCO3 deposition associated with cyano- alga Spirogyra hatillensis T. that contains calcium phytes (photosynthetic cyanobacteria) that marks oxalate inclusions. The inclusions are not asso- the end of the Precambrian period, ,0.6 Gyr ago ciated with the central vacuole, but instead are in (Riding, 1982). At the start of the Cambrian period cytoplasmic strands (Pueschel, 2001). (,570 Myr ago), calcareous skeletons appear; With the advent of eukaryotes, subdivisions rapidly and dramatically (within 40–50 Myr) within the cell, or compartments, were created. some form of biomineralized structures appear Within these subcellular compartments, or specia- in all existing phyla. Since that time only corals, lized anatomical sites, mineralization may be some algae, and the vertebrates have developed facilitated, with the result that biomineralization new skeletons in the marine habitat (Simkiss and became more extensive and diverse. By creating Wilbur, 1989); thus, the obvious developments of lipid membranes, the eukaryotes could selectively biomineralization are concentrated into ,1% of “pump” ions and bioaccumulate them in a small Earth’s history, along with all other major volume. In plants, this subcellular compartment is diversifications of life. We present information, usually a vacuole (Matile, 1978), whose membrane and include references to works we think will be may serve both as a pre-existing surface for most helpful to geochemists, on selected inverte- nucleation and as the ultimate determinant of brate and vertebrate skeletons many of which mineral shape, as the crystal(s) grow to fill the have a vast literature available. We select a few vacuole (Simkiss and Wilbur, 1989). Most ion structures, such as teeth (in chitons and humans), pumps translocate ions against electrochemical because they are examples of different minerali- gradients (Carapoli and Scarpa, 1982) accom- zing systems whose tissue textures, and mech- plished either by attaching the ion to a carrier anisms, are unique, and because they may be, or molecule that is moving with an electrochemical have been, important to geochemical studies. In gradient, or by directly using ATP as an energy looking to future opportunities we include brief source for the translocation. Often described introductions to otoliths and antlers, because they for Ca2þ transport, similar transport mechanisms offer novel sampling sites to test geochemical have been suggested to supply the anions that variations in the present environment. control the onset of mineral deposition in cells The importance of the survival of land-based (Simkiss and Wilbur, 1989). Like cations, anions communities on plants suggested that our purview are involved in a wide variety of cell activities. must include plant biomineralization, the Primary among them is the ability of anionic materials, mechanisms, and strategies. For our complexes, e.g., carbonate and phosphate, to act as summary we ask “why biomineralize?” and offer a inorganic pH buffers. In support of the anion- few suggestions based predominantly on plant supply hypothesis, it has long been recognized that researches. The reasons for biomineralization we there is a positive relationship between photosyn- have outlined are appropriate to other mineral- thetic rate (acquisition of CO2) and rate of calcite producing life forms, and have often been biomineralization in algae. Specific studies have discussed. However, in the process of reviewing shown that when Corallina officianalis algae the evolutionary development, some novel achieve a certain level of photosynthesis, the approaches, if not answers, to this basic question relationship between calcite biomineralization are preferred. rate and CO2 acquisition is roughly linear (Pentecost, 1978). An organic matrix or pre-existing nucleation 8.04.1.2 Definitions and General Background surface is usually considered to be the determining on Biomineralization feature in many systems, especially the higher biomineralizing systems, such as the vertebrates We classify biomineralization in our examples (Section 8.04.3.7). Organic matrices within bio- either as extracellular or as intracellular (Pentecost, mineralizing plant vacuoles (Webb et al., 1995) 1985a) and include the specific cell types if provide the sites where “seed” cations bind as loose known. We follow the standard definitions of chelates (Tyler and Simkiss, 1958), and can be Borowitzka (1982) that extracellular biominerali- alternately soluble and insoluble (Wheeler et al., zation involves inorganic, often crystalline, 1981), or a combination of the two (Degens, 1976). materials forming on the outer wall of the cell, Several nucleation centers may be present within a within the cell wall, or in the immediate surround- matrix, and each may grow independently, and ing tissue areas, and is the usual type of perhaps produce similar crystallographically biomineralization.
Load more

Recommended publications
  • Four Hundred Million Years of Silica Biomineralization in Land Plants
    Four hundred million years of silica biomineralization in land plants Elizabeth Trembath-Reicherta,1, Jonathan Paul Wilsonb, Shawn E. McGlynna,c, and Woodward W. Fischera aDivision of Geological and Planetary Sciences, California Institute of Technology, Pasadena, CA 91125; bDepartment of Biology, Haverford College, Haverford, PA 19041; and cGraduate School of Science and Engineering, Tokyo Metropolitan University, Hachioji-shi, Tokyo 192-0397, Japan Edited by Thure E. Cerling, University of Utah, Salt Lake City, UT, and approved February 20, 2015 (received for review January 7, 2015) Biomineralization plays a fundamental role in the global silicon Silica is widely used within plants for structural support and cycle. Grasses are known to mobilize significant quantities of Si in pathogen defense (19–21), but it remains a poorly understood the form of silica biominerals and dominate the terrestrial realm aspect of plant biology. Recent work on the angiosperm Oryza today, but they have relatively recent origins and only rose to sativa demonstrated that silica accumulation is facilitated by taxonomic and ecological prominence within the Cenozoic Era. transmembrane proteins expressed in root cells (21–24). Phy- This raises questions regarding when and how the biological silica logenetic analysis revealed that these silicon transport proteins cycle evolved. To address these questions, we examined silica were derived from a diverse family of modified aquaporins that abundances of extant members of early-diverging land plant include arsenite and glycerol transporters (19, 21, 25, 26). A clades, which show that silica biomineralization is widespread different member of this aquaporin family was recently identi- across terrestrial plant linages. Particularly high silica abundances fied that enables silica uptake in the horsetail Equisetum,an are observed in lycophytes and early-diverging ferns.
    [Show full text]
  • Biomineralization and Global Biogeochemical Cycles Philippe Van Cappellen Faculty of Geosciences, Utrecht University P.O
    1122 Biomineralization and Global Biogeochemical Cycles Philippe Van Cappellen Faculty of Geosciences, Utrecht University P.O. Box 80021 3508 TA Utrecht, The Netherlands INTRODUCTION Biological activity is a dominant force shaping the chemical structure and evolution of the earth surface environment. The presence of an oxygenated atmosphere- hydrosphere surrounding an otherwise highly reducing solid earth is the most striking consequence of the rise of life on earth. Biological evolution and the functioning of ecosystems, in turn, are to a large degree conditioned by geophysical and geological processes. Understanding the interactions between organisms and their abiotic environment, and the resulting coupled evolution of the biosphere and geosphere is a central theme of research in biogeology. Biogeochemists contribute to this understanding by studying the transformations and transport of chemical substrates and products of biological activity in the environment. Biogeochemical cycles provide a general framework in which geochemists organize their knowledge and interpret their data. The cycle of a given element or substance maps out the rates of transformation in, and transport fluxes between, adjoining environmental reservoirs. The temporal and spatial scales of interest dictate the selection of reservoirs and processes included in the cycle. Typically, the need for a detailed representation of biological process rates and ecosystem structure decreases as the spatial and temporal time scales considered increase. Much progress has been made in the development of global-scale models of biogeochemical cycles. Although these models are based on fairly simple representations of the biosphere and hydrosphere, they account for the large-scale changes in the composition, redox state and biological productivity of the earth surface environment that have occurred over geological time.
    [Show full text]
  • A Perspective on Underlying Crystal Growth Mechanisms in Biomineralization: Solution Mediated Growth Versus Nanosphere Particle Accretion
    CrystEngComm A perspective on underlying crystal growth mechanisms in biomineralization: solution mediated growth versus nanosphere particle accretion Journal: CrystEngComm Manuscript ID: CE-HIG-07-2014-001474.R1 Article Type: Highlight Date Submitted by the Author: 01-Dec-2014 Complete List of Authors: Gal, Assaf; Weizmann Institute of Science, Structural Biology Weiner, Steve; Weizmann Institute of Science, Structural Biology Addadi, Lia; Weizmann Institute of Science, Structural Biology Page 1 of 23 CrystEngComm A perspective on underlying crystal growth mechanisms in biomineralization: solution mediated growth versus nanosphere particle accretion Assaf Gal, Steve Weiner, and Lia Addadi Department of Structural Biology, Weizmann Institute of Science, Rehovot, Israel 76100 Abstract Many organisms form crystals from transient amorphous precursor phases. In the cases where the precursor phases were imaged, they consist of nanosphere particles. Interestingly, some mature biogenic crystals also have nanosphere particle morphology, but some are characterized by crystallographic faces that are smooth at the nanometer level. There are also biogenic crystals that have both crystallographic faces and nanosphere particle morphology. This highlight presents a working hypothesis, stating that some biomineralization processes involve growth by nanosphere particle accretion, where amorphous nanoparticles are incorporated as such into growing crystals and preserve their morphology upon crystallization. This process produces biogenic crystals with a nanosphere particle morphology. Other biomineralization processes proceed by ion-by-ion growth, and some cases of biological crystal growth involve both processes. We also identify several biomineralization processes which do not seem to fit this working hypothesis. It is our hope that this highlight will inspire studies that will shed more light on the underlying crystallization mechanisms in biology.
    [Show full text]
  • An Overview of Biomineralization Processes and the Problem of The
    11 An Overview of Biomineralization Processes and the Problem of the Vital Effect Steve Weiner Department of Structural Biology Weizmann Institute of Science 76100 Rehovot Israel Patricia M. Dove Department of GeoSciences Virginia Tech Blacksburg, Virginia 24061 U.S.A. “Biomineralization links soft organic tissues, which are compositionally akin to the atmosphere and oceans, with the hard materials of the solid Earth. It provides organisms with skeletons and shells while they are alive, and when they die these are deposited as sediment in environments from river plains to the deep ocean floor. It is also these hard, resistant products of life which are mainly responsible for the Earth’s fossil record. Consequently, biomineralization involves biologists, chemists, and geologists in interdisciplinary studies at one of the interfaces between Earth and life.” (Leadbeater and Riding 1986) INTRODUCTION Biomineralization refers to the processes by which organisms form minerals. The control exerted by many organisms over mineral formation distinguishes these processes from abiotic mineralization. The latter was the primary focus of earth scientists over the last century, but the emergence of biogeochemistry and the urgency of understanding the past and future evolution of the Earth are moving biological mineralization to the forefront of various fields of science, including the earth sciences. The growth in biogeochemistry has led to a number of new exciting research areas where the distinctions between the biological, chemical, and earth sciences disciplines melt away. Of the wonderful topics that are receiving renewed attention, the study of biomineral formation is perhaps the most fascinating. Truly at the interface of earth and life, biomineralization is a discipline that is certain to see major advancements as a new generation of scientists brings cross-disciplinary training and new experimental and computational methods to the most daunting problems.
    [Show full text]
  • Effect of Ph Change on Exoskeletons of Selected Saltwater Organisms Which Rely on Calcium Fixation Derya Z
    Journal of Emerging Investigators Effect of pH change on exoskeletons of selected saltwater organisms which rely on calcium fixation Derya Z. Tansel1, Ariadna Arreaza2, Berrin Tansel2 1 Coral Gables Senior High, Coral Gables, FL 2 Florida International University, Miami, FL Summary increase in H+ concentration in the last 200 years (1,2,3). The projections for rising atmospheric carbon dioxide According to atmospheric CO2 projections, ocean surface concentrations indicate that the pH levels of the ocean pH levels are estimated to decrease by 0.3-0.4 units by surface could decrease by 0.3-0.4 units by the end of the end of the 21st century. This decrease corresponds the 21st century. The objective of this research was to to an increase in the hydrogen ion concentration of about evaluate the effect of pH on the exoskeletons of six aquatic organisms commonly found in South Florida 100-150% above the levels in the late 1800s (4,5). The coastal waters. The exoskeleton samples studied were impacts of ocean acidification can be 10–50% higher from the common nutmeg (Cancellaria reticulate), near coastal areas due to proximity to anthropogenic lettered olive (Oliva sayana), stiff pen shell (Atrina rigida), sources (6). kitten’s paw (Plicatulidae), fan coral (Gorgonia ventalina), Although some species can tolerate pH changes, and common slipper shell (Crepidula fornicate). The many marine organisms and processes can be impacted, exoskeleton samples were exposed to saltwater (34% including the composition of communities and food webs salinity) at pH levels ranging from 8.3 to 6.0 for 5 days.
    [Show full text]
  • Biomineralization and Evolutionary History Andrew H
    1 111 Biomineralization and Evolutionary History Andrew H. Knoll Department of Organismic and Evolutionary Biology Harvard University Cambridge, Massachusetts, 02138 U.S.A. INTRODUCTION The Dutch ethologist Niko Tinbergen famously distinguished between proximal and ultimate explanations in biology. Proximally, biologists seek a mechanistic understanding of how organisms function; most of this volume addresses the molecular and physiological bases of biomineralization. But while much of biology might be viewed as a particularly interesting form of chemistry, it is more than that. Biology is chemistry with a history, requiring that proximal explanations be grounded in ultimate, or evolutionary, understanding. The physiological pathways by which organisms precipitate skeletal minerals and the forms and functions of the skeletons they fashion have been shaped by natural selection through geologic time, and all have constrained continuing evolution in skeleton-forming clades. In this chapter, I outline some major patterns of skeletal evolution inferred from phylogeny and fossils (Figure 1), highlighting ways that our improving mechanistic knowledge of biomineralization can help us to understand this evolutionary record (see Leadbetter and Riding 1986; Lowenstam and Weiner 1989; Carter 1990; and Simkiss and Wilbur 1989 for earlier reviews). Figure 1. A geologic time scale for the past 1000 million years, showing the principal time divisions used in Earth science and the timing of major evolutionary events discussed in this chapter. Earlier intervals of time—the Mesoproterozoic (1600–1000 million years ago) and Paleoproterozoic (2500– 1600 million years ago) eras of the Proterozoic Eon and the Archean Eon (> 2500 million years ago)— are not shown. Time scale after Remane (2000).
    [Show full text]
  • Biomineralization & Biominerals “It's a Hard Life”
    Maria Isabel López Fierro Biomineralization Literature Review & Biominerals Chair: Marc A. Meyers “It’s a hard life” Joanna McKittrick Jan Talbot Image credit: Out Of The Blue Aquaculture Ltd 1 Outline Sponge Spicule Introduction & History Basic Biomineralization Principles Saturation, Nucleation, Growth, & Organic Matrix Image credit: http://www.choijinhyuk.com Bleached Coral Biominerals & Biomineralization Models Calcium Carbonate Shells: Nacre Silica Image credit: jroy.abenaza.com2 Sponge Spicule Hydroxyapatite Bone Why do we need to understand biomineralization? Summary & Conclusions Introduction & History of Biomineralization 3 Introduction: The Process Biomineralization is the process by which living form and influence the precipitation of minerals. 5µm No single or ‘grand’ •Crystals from the inner shell layer of the Eastern mechanism. oyster onto a metal implant. •sheets converge forming a“rosette” structure. Combination of efforts •Organic matrix appears like “glue.” from cells, producing Image credit: http://www.scienceasart.org organic and inorganic Skinner and Jahren (2003)4 molecules that combine in various structural ways to form an unique material. Introduction: Why mineralize? Evolution of Biomineralization has provided organisms with a strong building material. o Minerals are stiff and brittle (& cheap energy wise) o Organic materials are soft and pliable Functions include: v Strength & Integrity v Protection. v Mobility v Storage - Biominerals are ion reservoir for cellular functions. v Cutting and grinding v Buoyancy v Optical, magnetic and gravity sensing 5 History of biominerals First biomineralization evidence from microbial stromatolites - 3500M years ago Not controlled deposition of inorganic solids 560M years ago - organisms from different phyla evolved the ability Runnegar and Bengtson (1992) to form different minerals. To date there are 64+ 6 biominerals identified.
    [Show full text]
  • An Introduction to Biomineralization
    MATERIAL SCIENCE MEETS MEDICINE – PRESENTATION ABSTRACTS BIOMINERALIZATION An Introduction to Biomineralization John Dunlop Department of Biomaterials, Max Planck Institute of Colloids and Interfaces, Potsdam-Golm Many living organisms, ranging from bacteria to plants to animals, can control the formation of mineral both within and around their cells. These bio-minerals play important functional roles in organisms. The most obvious function is structural, with nano-sized mineral particles reinforcing our bones being a well- known example. Biominerals are also used in living systems in many other ways. To name just a few examples; they are used as transparent lenses as well as magnetic sensors and they also store important ions such as calcium used in the metabolism. All of these different functions arise through the exquisite control of the biomineralization process. Nature forms these minerals in incredibly complex shapes, with well-defined sizes and in addition can control what phase of mineral is formed (crystalline or amorphous for example). As such much research has been done to understand the process of biomineralization particularly with its application in the bioinspired design of novel materials. In medicine, and especially in the field of bone biology knowledge of how bone mineral forms is fundamental in the understanding, treatment and control of degenerative diseases such as osteoporosis. This presentation will give a general overview of the concept of biomineralization, and its application to medicine. The idea is to firstly present the evolutionary background of minerals in biology, showing a variety of examples from all domains of life. The presentation will then focus on giving a short summary of how mineralized tissues form, and then give an overview on how minerals organisation controls the tissue mechanical properties.
    [Show full text]
  • Remineralization of Demineralized Dentin Using a Dual-Analog System
    Remineralization of demineralized dentin using a dual-analog system N Saxena1 S Habelitz2 GW Marshall2 LB Gower1 1 – Materials Science and Engineering, University of Florida, Gainesville, FL, USA 2 – Preventative and Restorative Dental Sciences, University of California San Francisco, San Francisco, CA, USA Correspondence to: Laurie Gower Materials Science and Engineering University of Florida 549 Gale Lemerand Dr. Gainesville, FL 32611 [email protected] 1 Abstract Objectives Improved methods are needed to remineralize dentin caries in order to promote conservation of dentin tissued and minimize the surgical interventions that are currently required for clinical treatment. Here we test the hypothesis that bulk substrates can be effectively mineralized via a dual- analog system proposed by others, using a tripolyphosphate “templating analog” and a poly(acrylic acid) or poly(aspartic acid) “sequestration analog”, the latter of which generates the polymer-induced liquid-precursor (PILP) mineralization process studied in our lab. Material & Methods – Demineralized human dentin slices were remineralized with and without pre-treatment with tripolyphosphate. Both poly(acrylic) acid and poly(aspartic acid) were used as PILP process-directing agents or “sequestration analogs”. A control experiment with no polymer present was also conducted. Results – No mineralization was observed in any of the poly(acrylic acid) groups. In both the poly(aspartic acid) and no polymer groups, tripolyphosphate inhibited mineralization on the surfaces of the specimens and promoted mineralization within the interiors of the specimens. Pre- treatment with tripolyphosphate enhanced overall mineralization of the poly(aspartic acid) group. However, when analyzed via TEM, regions with little mineral were still present. Conclusion – Poly(acrylic acid) was unable to remineralize demineralized dentin slices, even when pre-treated with tripolyphosphate.
    [Show full text]
  • Biomineralization of Collagen-Based Materials for Hard Tissue Repair
    International Journal of Molecular Sciences Review Biomineralization of Collagen-Based Materials for Hard Tissue Repair Le Yu 1 and Mei Wei 1,2,* 1 Department of Chemical and Biomolecular Engineering, Ohio University, Athens, OH 45701, USA; [email protected] 2 Department of Mechanical Engineering, Ohio University, Athens, OH 45701, USA * Correspondence: [email protected]; Tel.: +1-740-593-1474 Abstract: Hydroxyapatite (HA) reinforced collagen fibrils serve as the basic building blocks of natural bone and dentin. Mineralization of collagen fibrils play an essential role in ensuring the structural and mechanical functionalities of hard tissues such as bone and dentin. Biomineraliza- tion of collagen can be divided into intrafibrillar and extrafibrillar mineralization in terms of HA distribution relative to collagen fibrils. Intrafibrillar mineralization is termed when HA minerals are incorporated within the gap zone of collagen fibrils, while extrafibrillar mineralization refers to the minerals that are formed on the surface of collagen fibrils. However, the mechanisms re- sulting in these two types of mineralization still remain debatable. In this review, the evolution of both classical and non-classical biomineralization theories is summarized. Different intrafibrillar mineralization mechanisms, including polymer induced liquid precursor (PILP), capillary action, electrostatic attraction, size exclusion, Gibbs-Donnan equilibrium, and interfacial energy guided theories, are discussed. Exemplary strategies to induce biomimetic intrafibrillar mineralization using non-collagenous proteins (NCPs), polymer analogs, small molecules, and fluidic shear stress are discussed, and recent applications of mineralized collagen fibers for bone regeneration and dentin repair are included. Finally, conclusions are drawn on these proposed mechanisms, and the future trend of collagen-based materials for bone regeneration and tooth repair is speculated.
    [Show full text]
  • Biomineralization, Biomimetic and Non- Classical Crystallization
    BiomineralizationBiomineralizationBiomineralization,,, biomimeticbiomimeticbiomimeticandand and nonnonnon--- classicalclassicalclassicalcrystallizationcrystallization crystallization WaysWaysWaystoto tounderstandunderstand understand thethethe synthesissynthesissynthesisandand and formationformationformation mechanismsmechanismsmechanismsofof of complexcomplexcomplex materialsmaterialsmaterials HelmutHelmut CölfenCölfen Max-Planck-Institute of Colloids & Interfaces, Colloid Chemistry, Research Campus Golm, D-14424 Potsdam [email protected] CaCOCaCOCaCO333 BiomineralsBiomineralsBiominerals Coccolith (Calcite) Nacre (Aragonite) Herdmania momus (Vaterite) Characteristics of Biominerals: Complex forms generated from inorganic systems with simple structure Characteristics of inorganic crystals: Simple geometrical form but often complicated crystal structure Default: Rhombohedral Calcite ImportantImportantImportantbiomineralsbiominerals biominerals MineralMineral FormulaFormula FunctionFunction Calciumcarbonate CaCO3 Calcite Algae / Exoskeleton Birds / Eggshell Aragonite Fishes / Gravity sensor Mussels / Exoskeleton Vaterite Sea urchins / Spikes Calciumphosphate Hydroxyapatite Ca10(PO4)6(OH)2 Vertebrates / Skeleton, teeth Octa-Calciumphosphate Ca8H2(PO4)6 Vertebrates / Precursor for bone formation Silica SiO2 Algae / Exoskeleton Iron oxide Magnetite Fe3O4 Salmon, Tuna & bacteria / Magnetic field sensor Chitons / teeth EvolutionEvolutionEvolution ofofof aaa musselmusselmussel shellshellshell SEM of broken mussel shell SEM of broken
    [Show full text]
  • Terraced Iron Formations: Biogeochemical Processes Contributing to Microbial Biomineralization and Microfossil Preservation
    geosciences Article Terraced Iron Formations: Biogeochemical Processes Contributing to Microbial Biomineralization and Microfossil Preservation Jeremiah Shuster 1,2,* , Maria Angelica Rea 1,2 , Barbara Etschmann 3, Joël Brugger 3 and Frank Reith 1,2 1 School of Biological Sciences, The University of Adelaide, Adelaide, South Australia 5005, Australia; [email protected] (M.A.R.); [email protected] (F.R.) 2 CSIRO Land and Water, Contaminant Chemistry and Ecotoxicology, PMB2 Glen Osmond, South Australia 5064, Australia 3 School of Earth, Atmosphere & Environment, Monash University, Clayton, Victoria 3800, Australia; [email protected] (B.E.); [email protected] (J.B.) * Correspondence: [email protected]; Tel.: +61-8-8313-5352 Received: 30 October 2018; Accepted: 10 December 2018; Published: 13 December 2018 Abstract: Terraced iron formations (TIFs) are laminated structures that cover square meter-size areas on the surface of weathered bench faces and tailings piles at the Mount Morgan mine, which is a non-operational open pit mine located in Queensland, Australia. Sampled TIFs were analyzed using molecular and microanalytical techniques to assess the bacterial communities that likely contributed to the development of these structures. The bacterial community from the TIFs was more diverse compared to the tailings on which the TIFs had formed. The detection of both chemolithotrophic iron-oxidizing bacteria, i.e., Acidithiobacillus ferrooxidans and Mariprofundus ferrooxydans, and iron-reducing bacteria,
    [Show full text]