Handbook of Microscopic Anatomy Continuation of Handbuch der mikroskopischen Anatomie des Menschen

Founded by Wilhelm von MoUendorff Continued by Wolfgang Bargmann

Edited by A. Oksche and L. Vollrath

B.K.B. Berkovitz A. Boyde R.M. Frank H.J. H6hling B.J. Moxham 1. Nalbandian C.H. Tonge

TEETH

With 413 Figures

Springer-Verlag Berlin Heidelberg New York

London Paris Tokyo

Handbook of Microscopic A nato my Volume V /6: Teeth

Professor Dr. Drs. h.c. A. Oksche Institut fU r Anatomie und Zytobiologie der luS\us-Liebig-Universitiit, Aulweg 123, 0-6300 Giessen

Professor Dr. L. Vollrath Anatomisches Institut der Johannes Gutenberg-Universitat, SaarstraBe 19- 21, 0-6500 Mainz

ISBN-13: 978-3-642-83498-1 0 01: 10.1007/978-3-642-83496-7

e-ISBN-i3: 978-3-642-83496-7

Library of Congress Cataloging-in-Publication Data. Teeth I edited by A. Oksche and L. Vollra th ; contributors, B.K.B. Berkovitz ... lc t al. ). p. em. - (Handbook of microscopic anatomy; v. v/6) Includes bibliographies and indexcs. 1. Teeth - Histology. 2. Teeth - Growth. 3. Teeth -Ultrastructure. I. Oksche, A. 11. Vollrath, Lutz, 1936 - . III. Berkovitz, B.K. B. IV. Series.QM311

.T44 1988 611'.3 14 - dc1988-29489 CIP.

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List of Authors

B.K.B. BERKOVITZ, Dr., King's College London, University of London, Depart­ment of Anatomy and Human Biology, Strand, OS-London WC2R 2LS

A. BOYOE. Prof. Dr., University College London. Department of Anatomy and Embryology, Gower Street, GS-London Wet E 6BT

R.M. FRANK, Prof. Dr., Universite Louis Pasteur, Faculte de Chirurgie Den­taire, 4, Rue Kirschleger, F-67085 Strasbourg-Cedex

H.J. HOHLlNG, Prof. Dr., Institut fUr Medizinische Physik der Universitiit. Hiifferstr. 68, 0-4400 Munster

B.l . MOXHAM, Dr., University of Bristol, The Medical School, Department of Anatomy, OS-Bristol 8S8 ITO

J. NALBANDIAN, Dr. . University of Connecticllt, School of Dental Medicine. 263 Fannington Avenue, Farmington, Connecticut 06032, USA

C.H. TONGE, Emeritus Proressor or Oral Anatomy, The University or Newcastle upon Tyne, OB-Newcastle upon Tyne NEt 7RU

Preface

The editors and the publisher are pleased to present another volume in this series of monographs. The topic of teeth was last reviewed within the framework of this Handbook more than fifty years ago, in 1936, by Josef Lehner and HanDs Plenk of Vienna, who wrote a comprehensive treatise on the subject in volume V /3. The introduction of new methods (e.g., transmission and scan­ning electron microscopy, histochemistry, radioautography, element analysis) and progress in dental research have made an update necessary.

In present times, characterized by scientific specialization and very rapid progress, it is virtually impossible to find a single individual prepared to review a field of research as large as that of teeth. Consequently. several authors were asked to contribute to the present volume. Originally. the intention was to cover the field in one volume. When it became clear that the material had become too extensive for a single volume, and when some authors were forced by external factors to withdraw from the project, it was decided to publish two volumes. In 1986, the volume Periodontium, written by Hubert E. Schroeder of Zurich. appeared in this series (volume VIS, 418 pages).

It is not without irony that. in a time seemingly conducive to basic and applied research, potential authors have had to struggle to prevent the closure of their institutions. As a result some researchers could not contribUle to the present volume, and in other instances the contributions were delayed. The editors are grateful to the contributing authors and the publisher for their patience. We are confident that both volumes, Periodontium and Teeth, will become standard sources of scientific information, promoting both basic and applied dental research.

A. OKSCHE

L. VOLLRATH

Contents

Tooth Development - General Aspects C.H. TONGE (With 6 Figures)

A. General Arrangement of the Dental Tissues and Their Embryological Derivation. . . . . . . . . . . . . . .. ....... 1

B. Formation of Odontogenic Sites in the Developing Mo uth 2 I. Interactions Between Odontogenic Epithelium and

Eclomesenchyme . 2 II. Primordia fo r the Different Classes of Teeth 4

III . Early Innervation and Blood Supply . 5 C. Structural Formations in Mouth Development. . . 6

I. Tooth Germ, General Morphological and Histological Features of the Enamel Organ and Denta l Papilla ........ 7

II. Dental Lamina. Enamel Niche and Accessory Laminae. 9 III. Labial and Vestibular Laminae. . . 9 IV. Linguoalveolar Sulcus.. .. . ......... 10 V. Dental Follicle and Ea rly Alveolus . . . . . . . . 10

D. The Development of the Crown a nd its Morphology . II I. Developmental Integuments of the Enamel Surface 12

E. Development of the Periodon tium. 12 I. Outline of Root Fonnation . 12

II. The Periodontal Ligament . 13 III. Epithelial Rests of Malassez 14

F. The Developmental Positions of the Teeth in the Jaws. 14 I. The Deciduous Teeth . 14

II. The Successional Teeth 15 III. The Pennanent Molars 15 IV. Growth Retardation . 16

G. Developmental Anomalies. 17 References . . . . . .. . . .. 17

Tissue Changes During Tooth Eruption B.K .B. BERKOVITZ and 8.J. MOXHAM (With 24 Figures)

A. Tissue Changes During Eruption I. The Pre-Eruptive Phase ... .

II. The Eruptive Phase ..... . I . The Development of the Periodontal Ligament During the

Eruptive Phase ............ .

21 22 22

22

x Coolcnts

2. Adaptation of the Alveolar Bone to Eruption. . . . . . . 35 3. AJterations in the Tissues Overlying the Erupting Tooth. 37

III. The Intra-Oral Phase 49 B. Mechanisms of Tooth Eruption. . . . . . . . . . . . 51

L The Collagen Contraction Hypothesis. 52 II. The Periodontal F ibroblast Contraction/ Motility Hypothesis . 54

III. The Vascular[rissue Fluid Pressure Hypothesis. 60 References . . . . . . . . . . . . . . . . . . . . . 63

Development of Dentine and Pulp R.M. FRANK and 1. NALBANDIAN (With 88 Figures)

A. Introduction . . .. .... . B. C. D. E. F.

The Dental Papilla . . ... The Internal Dental Epithelium and the Preameloblasts The Dental Basement Membrane ... . ... . . Epithelia-Mesenchymal Interactions .. The Odontoblast and the Odontoblastic Process.

J. Differentiation of the Odontoblasts . II. The Cell Body of the Differentiated Odontoblast.

III. The Odontoblastic Process .. . .. . IV. The Metabolism of the Odontoblast .

1. Collagen Biosynthesis . . . . . . . 2. Glycoprotein and Fibronectin Biosynthesis 3. Proteoglycan Biosynthesis . . 4. Biosynthesis of y-Carboxyglutamic Acid (GLA)-Containing

Proteins . .... . . 5. Phosphoprotein Biosynthesis . . ..... . 6. Calcium and Phosphate Metabolism 7. Degradative Activities of the Odontoblasts

G. Predentine .. . H. The Process of Mineralization.

L Matrix Vesicles . . II. Alkaline Phosphatases . .

HI. The Concept of Epitaxy. IV. The Role of Various Non-Collagenous Proteins V. The Role of Lipids .

J. The Development of Coronal Dentine .... . . I. Intertubular Dentine .

1. Mantle Dentine and the Dentinoenamel Junction. 2. CircumpulpaJ Dentine

II. Peri tubular Dentine . K. The Development of Root Dentine . L. The Development of the Pulp. References . . . . .

73 74 79 85 87 91 93 99

110 114 114 120 121

122 122 123 128

.130 134 137 137 138 139 140 141 141 141 147 151 152 155 159

Conttnts

Structure and Ultrastructure of Dentine R.M. FRANK and J. NALBANDIAN (With 95 Figures)

A. Basic Anatomy . . . . ....... . B. Physical and Chemical Characteristics C. Classification of Different Types o f Dentine. D. The Odontoblast and Their Processes.

I. The Odontoblast Cell Body. II. The Odontoblastic Process . . .

E. Predentine. . . . . . . . . . . . . . . . F. The Dentinal Tubules a nd Their Contents .

I. The Periodontoblastic Space II. The Lamina Limitans .

III. The Enamel Spindles O . Peritubular Dentine . . .. . H. Intertubular Dentine ... . J. Dentinoenamel and Cementodentinal Junctions K . Interglobular Dentine . . ..... .. . L. Granular Layer of Tomes ...... . M. Incremental Lines and Neonatal Lines N. Translucent Dentine ....... . . . O . Regular and Irregular Secondary Dentine References . . . . . . . . . . . . . . . .

Structure and Ultrastructure of the Dental Pulp R.M. FRANK and J. NALBANDIAN (With 59 Figures)

XI

173 175 177 178 182 190

.206

.207

.209 · 21 1 .212 · 212

216 .220 .224 · 227 .228 .230 · 235 · 238

A. Basic Anatomy . . . . . . 249 B. Pulpal Cells. .. ... . 251

I. The Odontoblasts . 251 II. The Fibroblasts . . 251

III. Other Pulpal CeUs . . 259 C. Extracellular Matrix of the Pulp. . 259 D. The Ce\lular Organization of the Pulp . 263

I. The Odontoblastic Layer . 263 II. The Subodontoblastic Layer . . . 263

III. The Central Pulp . . . . . . . . 264 E. Innervation and Dentine-Pulp Sensitivity . . 264

I. Histological Nerve Distribution in the Pulp-Dentine Complex . 265 II. Microscopk and Fine Structure of Pulpal and Dentinal Nerve

Fibrils. . . . . . . . . . . . . . . . . . . . 269 Ill. Dentine-Pulp Sensitivity. . . 284

1. The Odontoblast as a Receptor Cell . 284 2. The Hydrodynamic Hypothesis. . 285 3. Direct Neural Stimulation. . 286

XII CODtCllts

F. Vascular Supply. .287 . 287 .292 . 296 .299

I. Blood Vessels . II. Pulpal Lymphatics.

G. Age Changes and Pulpal Calcifications . References. . . .. . .. .

Enamel A. BoWE (With 121 Figures)

A. Introduction . . . . . 309 B. Gross Anatomy. . 310

I. Distribution . 310 II. Naked·Eye and Magnifying-Glass-Range Features. . 311

C. Microscopy and Technical Methodology for the Study of Enamel. 313 I. Light Microscopy. 313

1. Sectioning Enamel for Light Microscopy . 313 2. Examination of Ground Sections in Light Microscopy 314 3. Renected-Light Microscopy of WeB-Polished Section

Surfaces . . . . . . . . . . . .. . ... . ... 316 II. Transmission Electron Microscopy of Enamel . . 316

1. Sectioning Enamel for Transmission Electron Microscopy .. 316 2. Transmission Electron Microscopy of Replicas . 316 3. Ion-Beam Thinning . . . . . . . . . . . . 317

III. Electron Diffraction of Enamel Crystals . . 318 1. Isolation of Enamel Crystallites. . 318 2. Electron Diffraction . . . . . . 318 3. Origin of Contrasts . . 318

IV. Scanning Electron Microscopy. . 319 1. Scanning Electron Microscopy of Enamel .319

a) Secondary Electrons. .319 b) ~ack-Scattered Electrons . . 320 c) Cathodoluminescence. . . 321

2. Sample Preparation . . . . . . . 321 a) Sample Preparation Procedures for Adult Enamel. . 321 b) Preparative Procedures for lmmature Enamel . . 323

3. X-ra"y Emission Microanalysis . . 324 a) Characteristic X-ray Emission . . 324 b) Wavelength-Dispersive X-ray Analysis . 324 c) Energy-Dispersive X-ray Analysis. . 325

V. X-ray Microscopy and Microradiography. . . 325 1. Contact and Projection Microradiography . . 325 2. Scanning X-ray Microscopy. . 325 3. X-ray Diffraction. . 326

D. Development of the Tooth Genn Prior to Enamel Formation . . 326 1. The Tooth Genn . . . . . . . .. . . ...... . 326

II. Ameloblast Differentiation and the Pre-ameloblast. . 327 E. Enamel Formation . . . . . . . . . . . . . . . 328

I. The Fully Differentiated Ameloblast. . . 328

Contents XIII

II. Ultrastructural Detail Within Secretory-Phase Enamel. . 338 III. Discontinuities in tbe Crystal Orientation Pattern . 339 IV. Amelogenins and Other Matrix Components (Biochemistry,

Histochemistry, Hi stology and Retention During Maturation) . 341 v. Histology of Demineralised Enamel Matrix. . . . . . . . . . 342

VI. Histological and Histochemical Changes with Maturation. . 343

F. Enamel Crystals and Prisms. . . . . . . . . . . . . . 343 I. Crystal Chemistry of the Inorganic Component of Enamel 343

II. Size and Shape of Enamel Crystallites. . . . . . 345 III . Crystal Orientation and the Existence of Prisms . 347

1. The Enamel-Dentine Junction. .. ..... . 347 2. Majority-Phase Enamel Secretion. . . . . . . 348 3. Development of a Prism-Free Surface Layer. . 351 4. Modal Crystallite Orientation. . . . . . . . . . 353 5. Explanation of Crystallite Orientation in 3-D as a Function

of the Developmental Enamel Surface Morphology. .. . 353 6. Prism Patterns . . . . . . . . . . . . . 357

a) The Characteristic Profiles of the Developing Enamel Surface and the Three Packing Arrangements of Prisms. 357

b) Pattern I Enamel: Complete Prism Boundaries . 367 c) Patterns 2 and 3: Incomplete Prism Boundaries . 368 d) Pattern 2 Enamel. . . . . . . . . 368 e) Pattern 3 Enamel. . . . . . . . . . 368 f) Distribution of Principal Prism Types . 369 g) Arcades or Arches in Human Enamel Pattern 3 Prisms:

A Question of Depth or Decussation? . h) The Prism Boundary Discontinuity .. i) Seams, Pseudo-prisms and Evolution.

IV. Incrementa l Phenomena. 1. Cross-striations and/or Varicosities .... 2. The Incremental Lines or Brown Striae of Retzius 3. Neonatal Line. . . .......... . 4. Development of Cross-striations

V. General Directions of the Prisms as Seen in Low-Powered Light Microscopy 1. Prism Orientations .. 2. Prism Decussation

a) Movement of Ameloblasts . b) Extreme Decussation in the Rat Incisor c) Other Mammals ..... d) Human Enamel. e) Functional Significance .

G. The Enamel-Dentine Junction. I. The Shape of the Junction ..

II. Features Originating at the Enamel-Dentine Junction 1. Spindles. 2. Tufts ....... .

.370 · 371 · 371 · 371 .371 .376 .378 .382

· 389 · 389 · 391 · 391 .394 .394 · 396 · 401

.407

. 407

.408

.408 · 411

XIV Contents

3. Lamellae 4. Enamel Tubules .

H. Mature Enamel Surface Features I. Fissures ....

II. Prism·Free Enamel and Smooth Areas III . Pits, Perikymata and Imbrication Lines . IV. Cervical Margins .

V. Brochs VI. Surface-Overlapping Projections and Isolated Deep Pits .

VII. Regional Differences in the Enamel Surface VIII. Cement on the Enamel Surface. . .. . .....

IX. Calculus on the Enamel Surface .. J. Maturation of the Enamel and Eruption of the C rown .

1. Maturation and Maturation Ameloblasts . II. Information from Rodent Incisor Studies .

1. Main Features ...... . 2. Transitional Zones . 3. Rume-Ended Ameloblasts . 4. Smooth-Ended Ameloblasts . 5. Labelling Studies. 6. Cyclical Phenomena . .

III. Studies on Rooted Teeth . IV. Mechanisms or Maturation and Matrix Removal. V. End or Maturation - Variations in Composition .

VI. Eruption or Emergence .. VII. Post-eruptive Maturation

K. Functional Changes in Enamel 1. Wear ..

II. Erosion . . . . . . . . . . L Pathology

I. Acquired and Inherited Disorders II. Linear 'Hypoplasias .

III. Hypoplastic Grooves IV. Hypoplastic Pits.

V. Dental Fluorosis .. VI. Enamel Caries

VII. Iatrogenic Cutting and Fracturing. VIII. Acid-Etching .

Rererences . . . . . . . .

Special Aspects or Biomineralization or Dental Tissues H.J. HOHLING (With 20 Figures)

A. Does a Correlation Exist Between the Fundamental Processes or Enamel, Dentine, and Cementum Mineralization Which Might Lead

· 412 · 412 · 414 · 414 · 414 · 415 · 41 7 .420 · 422 . 423 .424 .426 .428 .428 · 431 · 431 . 431 .434 . 434 · 437 . 439 . 441 . 442 . 443 .443 .445 . 446 . 446 .447 . 447 . 447 . 448 .448 . 448 .448 . 449 · 453 . 459 .462

to a General Calcification Theory? . . . . 475

Contents

B. Fundamental Aspects of Enamel Mineralization. 1. Mineralization in an Ordered, lonotropic Gel

II. Calcium (and Phosphate) Transport ... . . . III. Current Ideas About the Morphological Aspects of Crystal

Nucleation and Crystal Growth in Enamel ....... . 1. The Prevailing Theory on the Morphology of Crystal

Nucleation and Crystal Growth in Enamel .. . . . 2. Aspects of Our Own Ideas Concerning the Morphology of

Crystal Nucleation and Crystal Growth in Enamel . 3. Does the Enamel Crystal Develop from One Nucleus or

from Several Nuclei? .. .. ... . . . IV. Description of the Matrix Macromolecules in Enamel ....

1. General Chemical Characterization. . . . . 2. Possible Crystalline, Paracrystalline Structure of the Enamel

xv

· 477 .477 · 481

· 483

· 483

.485

. 487

. 488 · 488

Proteins . . . . . . . . . . . . . . . . . . . 491 3. Amino-Acid Sequence Analysis for Active Sites for

Ca-Phosphate Nucleation . . . . . . . . . . . . 494 V. Crystal Structure of the Developing and Mature Enamel

Crystallites. . . . . . . . . . 495 1. Existence of a Primary Ca-Carbonate Mineral . . . . 495 2. Existence of a Primary Apatite with Strong Lattice

Distortions . . 495 3. Primary Existence of an Octocalciumphosphate . . . . 496

C. Fundamental Aspects of Dentine Mineralization, and, in Part, of Cementum, Bone and Cartilage . . . . . . . . . . . . . . . 497

I. General Aspects of Enamel and Dentine Mineralization. . 497 II. Mineralization of the Matrix Vesicles . . 497

1. Development and Morphology of Matrix Vesicles . . . . 497 2. Histochemical and Chemical Characterization of the

Components of the Matrix Vesicles . . 500 3. Analysis of the Fine Structure and the Pathway of the

Matrix Vesicles by Means of Staining with Bi3+ Ions . 501 4. Mineralization of the Matrix Vesicles and Their Immediate

Neighborhood . . . . . . . . . 503 a) Mineralization in the Matrix Vesicles ............ 503 b) Extravesicular Mineralization; Radial Arrangement of

the Elongated Crystallites . 504 III. Mineralization of the Collagen Fibers. . . . . . . . . 505

1. General Aspects . . . . . . ... . . . 505 2. Relation to the ParacrystalJine ColIagen Structure 508

a) Morphological Observations and Measurements of Mineralized Collagen. . 508

b) Collagen Structure . . . . . . ......... . . . 509 c) Mineralization on the Surface of the Collagen Fibers . . 514

IV. Extracollagenous Mineralization . 514 I. General Aspects. ............. . ...... 514

XVI Contents

2. Types of Macromolecules of the NODcollagenous Regions .. 515 a) Phosphoproteins · 515 b) Proteoglycans . · 515 c) Glycoproteins .. · 516 d) y-Carboxyglutarrtic Acid-Containing Proteins and

Gla-Proteins, e.g., Osteocalcin. . . . . . . . . . . . . 517 D. Conclusion: Differences and Similarities in Mineralization in Enamel,

Dentine, and Other Hard Tissues, and a General Calcification Theory 517 References . . . . . .. ...... . . . . . . . . 520

Author Index · 525

Subject Index · 551