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Biology of the Integument 1 Invertebrates
Edited by 1. Bereiter-Hahn . A.G. Matoltsy K. Sylvia Richards
With 298 Figures
Springer-Verlag Berlin Heidelberg New York Tokyo 1984
Prof. Dr. JURGEN BERElTER-HAHN Fachbereich Biologie der UniversiHit, Kinematische Zellforschung Postfach 111932, D-6000 Frankfurt a.M./FRG
Prof. Dr. A. GEDEON MATOLTSY Boston University School of Medicine, Department of Dermatology 80 East Concord Street, Boston, MA 02118/USA
Dr. K. SYLVIA RICHARDS Department of Biological Sciences, University of Keele Keele, Staffordshire ST5 5BG/GB
ISBN 978-3-642-51595-8 ISBN 978-3-642-51593-4 (eBook) 10.1007/978-3-642-51593-4 DOI
Library of Congress Cataloging in Publication Data; Library of Congress Cataloging in Publication Data. Main entry under title: Biology of the integument. Bibliography: p. Includes index. Contents: v. 1. Invertebrates. 1. Skin-Collected works. 2. Body covering (Anatomy)--Collected works. 1. Bereiter-Hahn, Jiirgen. II. Matoltsy, A. Gedeon. III. Richards, K. Sylvia. QP88.5.B565 1984 591.1'858 83-20351
This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks.
Under § 54 of the German Copyright Law where copies are made for other than private use, a fee is payable to "Verwertungsgesellschaft Wort" Munich.
© by Springer-Verlag Berlin Heidelberg 1984 Sof'tcover reprint of the hardcover 1st edition 1984
The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use.
Typesetting, printing and bookbinding: Universitatsdruckerei H. Stiirtz AG, Wiirzburg. 2131/3130-543210
Preface
The integument plays an important role in the survival of metazoans by separating and protecting them from a hostile environment. Its function ranges from protection against injury and infection; partlcipation in the regulation of body temperature and water balance, to respiratory activity, monitoring of the environment and production of signals related to behaviour. All these result from specific structural, biochemical and physiological properties of intra- and extracellular components of the integument. Thus its characterization can be best accomplished by a multidisciplinary approach with authors specialized in different fields of science.
This multi-author book, in two volumes, provides an up-todate survey of the literature. The first volume deals with the integument of invertebrates, the second with that of vertebrates, both organized primarily on a phylum basis. As the level of knowledge on the integument of phyla differs considerably, the information provided is correspondingly either limited or condensed. For some of the smaller groups of invertebrates little information is available, as often only a few electron micrographs are to be found in the literature; on the other hand, from the large body of knowledge existing for vertebrates, particularly for mammals, no complete overview can be provided, but publications giving access to further information have been reviewed critically. A compilation of the knowlege on the animal integument is at the same time a compilation of the gaps; it is hoped that this monograph will stimulate further research in skin biology. The articles were written primarily for researchers, but some background information is also included for interested students and teachers. No comparable text dealing with all the invertebrate phyla or even all the vertebrate classes exists in the literature.
A book on Skin biology was proposed originally by Dr. Czeschlik of Springer-Verlag. It has been an attractive and challenging task for the editors to evaluate the literature and select subjects for these volumes. We received valuable advice from many colleagues, including Professors C. Arme (Keele), F.G. Barth (Frankfurt), D.T. Downing (Iowa City), R.M. Rieger
VI Preface
(Chapel Hill) and G.E. Rogers (Adelaide), to all of whom we are indebted as also to all the authors for their contributions to this work. The cooperation of the Publishers has been greatly appreciated.
J. BEREITER-HAHN
A.G. MATOLTSY
K.S. RICHARDS
Contents
I Introduction, Intercellular Junctions and Cilia
1 Introduction. K.S. RICHARDS 1
2 Intercellular Junctions. C.R. GREEN. (With 10 Figures) 5
2.1 Introduction 5 2.2 Desmosomes 6 2.3 Gap Junctions 8 2.4 Septate Junctions 10 2.5 Tricellular Junctions 12 2.6 Tight Junctions 14
3 Cilia. M.J. SANDERSON. (With 7 Figures) 17
3.1 Introduction 18 3.2 Ciliary Beat Patterns 19 3.3 Ciliary Structure 23 3.4 Mechanisms of Ciliary Motility 30 3.5 Control of Ciliary Activity 35
II The Diploblastic Level
4 Introduction to the Diploblastic Level. G.O. MACKIE 43
5 Cnidaria: Cell Types. M. HiiNDGEN. (With 5 Figures) 47
5.1 Introduction 47 5.2 Epithelial Cells 48 5.3 Mucous Cells 52 5.4 Nerve Cells 52 5.5 Sense Cells 54 5.6 Interstitial Cells 54
6 Cnidaria: Cnidae. R.N. MARISCAL. (With 8 Figures) 57
6.1 Introduction 57 6.2 Terminology 58 6.3 The Structure of Cnidae 59 6.4 The Functioning of Cnidae 62
VIII Contents
6.5 The Functions of Cnidae 64 6.6 The Chemistry of Cnidae 65 6.7 Conclusions. . . . . . 66
7 Cnidaria: Secreted Surface. J.G. TIDBALL 69
7.1 Introduction .......... 69 7.2 Settlement and Attachment . . . . 69 7.3 Food Capture and Surface Cleaning 71 7.4 Self-Recognition . . . . 73 7.5 Support and Protection . . . . . . 74 7.6 Conclusion and Questions . . . . . 76
8 Cnidaria: Permeability, Epidermal Transport and Related Phenomena. D. SCHLICHTER. (With 10 Figures) ........... 79
8.1 Introduction . . . . . . . . . . . 79 8.2 Permeability Properties of Apical Epidermal
Membranes . . . . . . . . . . . . . . 80 8.3 Other Phenomena Related to the Outer Epidermal
Surface . . . . . . . . . . . . . . . . . .. 91
9 Ctenophora. M.-L. HERNANDEZ-NICAISE. (With 7 Figures) 96
9.1 Introduction 96 9.2 Glandular Cells 97 9.3 Ciliated Cells (Other than Locomotory) 98 9.4 Single Sensory Cells 99 9.5 The Locomotory System ...... 101 9.6 Apical Organ . . . . . . . . . . . 103 9.7 The Integument of the Tentacles: the Colloblast 107 9.8 Other Cell Types ............. 109
ill Acoelomata
10 Turbellarian Platyhelminths. S. TYLER. (With 21 Figures) ..... . 112
10.1 Introduction . . . . . . 112 10.2 Morphology of Epidermal Cells 113 10.3 Cell Web . . . . . . . . . . 119 10.4 Intercellular Junctions .... 121 10.5 Ultrarhabdites and Other Secretory Products of
Epithelial Cells (" Epitheliosomes") 123 10.6 Extracellular Matrices ....... 125 10.7 Cilia . . . . . . . . . . . . . . . 126 10.8 Non-Epithelial Cells of the Body Wall 126 10.9 Epidermal Growth and Replacement 127 10.10 Phylogenetic Significance . . . . . . 128
Contents IX
11 Parasitic Platyhelminths. L.T. THREADGOLD. (With 25 Figures) .......... . 132
11.1 Introduction ............... 132 11.2 Trematoda: Larval Stages Monogenea, Digenea, and
Aspidogastrea . . . . 133 11.3 Trematoda: Adult . . 156 11.4 Cestoda: Larval Stages 163 11.5 Cestoda: Adult 178 11.6 Summary . . . . . . 184
12 Platyhelminth Host-Parasite Interface. A.D. BEFus and R.B. PODESTA . . . . . . . . . . . . . 192
12.1 Introduction ............... 192 12.2 The Structure and Function of Parasite Surfaces 193 12.3 Properties of the Surface Membrane and Interface 194 12.4 Host-Parasite Interface and Adhesiveness . . . . . 195 12.5 Survival Tactics of Parasites Confronted by Effectors
of Host Resistance . . . . . . . . . . . . . . . 196 12.6 The Host-Parasite Interface: Some New Directions 199 12.7 Conclusion . . . . . . . . . . . . . 200
13 Nemertea. R. GIBSON. (With 6 Figures)
13.1 Introduction ..... 13.2 Ciliated Supporting Cells 13.3 Gland Cells . . . . 13.4 Epidermal Enzymes 13.5 Sensory Cells 13.6 Basal Cells .... 13.7 Other Epidermal Features
IV Pseudocoelomata
14 Nematoda. A.F. BIRD. (With 9 Figures)
14.1 Introduction 14.2 Cells 14.3 Cuticle 14.4 Conclusion
· 205
· 205 · 207 · 207 · 208 · 209 · 210 · 210
· 212
· 212 · 213
218 230
15 Acanthocephala. P.J. WHITFIELD. (With 2 Figures) 234
15.1 Introduction . . . . . . 234 15.2 Structural Organization . . . . . . . 235 15.3 Integumentary Nutrient Uptake . 238
16 Minor Pseudocoelomates. V. STORCH. (With 12 Figures)
16.1 Introduction 16.2 Gastrotricha
242
242 243
X Contents
16.3 Kinorhyncha · 247 16.4 Rotifera · 250 16.5 Priapulida · 254 16.6 Nematomorpha · 258 16.7 Kamptozoa (Entoprocta) 261 16.8 Gnathostomulida · 263
V Annelida
17 Epidermal Cells. U. WELSCH et al. (With 12 Figures) 269
17.1 Introduction 269 17.2 Supporting Cells 272 17.3 Basal Cells 278 17.4 Gland Cells 278 17.5 Sensory Cells · 284 17.6 Gills · 289 17.7 Clitellum · 290
18 Chaetae. P.C. SCHROEDER. (With 3 Figures) . 297
18.1 Introduction . 297 18.2 Structure . . . 299 18.3 Composition 301 18.4 Development 304
19 Cuticle. K.S. RICHARDS. (With 11 Figures) 310
19.1 Introduction 310 19.2 Ultrastructure . . . . . . 310 19.3 Histochemistry ..... 315 19.4 Biophysics and Biochemistry 315 19.5 Subcuticular Bacteria . . . 318 19.6 Function . . . . . . . . 319
20 Permeability and Epidermal Transport. J. GOMME . • . • • . . . . . 323
20.1 Introduction ........ 323 20.2 Terminology and Basic Concepts 324 20.3 Exchange of Respiratory Gases 325 20.4 Transpiration . . . . . . . . 326 20.5 Water Permeability in Aquatic Annelids 328 20.6 Monovalent Ion Transport in Freshwater Annelids 332 20.7 Ion Transport in Terrestrial Annelids? ..... 337 20.8 Monovalent Ion Transport in Brackish-Water and
Marine Annelids . . . . . . . . 338 20.9 Transport of Other Ions .... 345 20.10 Transport of Organic Compounds 346 20.11 Role of the Cuticle 355 20.12 Conclusion . . . . . . . . . . 358
Contents XI
VI Annelid-Related Phyla and Cuticle Evolution
21 Echiura and Sipuncula. V. STORCH. (With 3 Figures) 365
21.1 Echiura . 365 21.2 Sipuncula 372
22 Pogonophora. E.C. SOUTHWARD. (With 5 Figures) 376
22.1 Structure 376 22.2 Permeability 384
23 Evolution of the Cuticle in the Lower Eumetazoa. R.M. RmGER. (With 2 Figures) 389
23.1 Introduction and Definitions 389 23.2 On the Origin of True Cuticles 390 23.3 The Primitive Condition of True Cuticles 390 23.4 The Lines of Cuticle Specializations Within the
Cnidaria and the Lower Bilateria 391 23.5 The Function of Primitive Eumetazoan Cuticles 396
VII Mollusca
24 Epidermal Cells. A. BUBEL. (With 5 Figures) · 400 24.1 Introduction · 400 24.2 Epidermal Cell Types · 401 24.3 Wound Repair .420 24.4 Byssus · 421 24.5 Operculum · 431 24.6 Siphon Sheath · 432 24.7 Mucus · 433 24.8 Radula · 439 24.9 Beaks .440
25 Shell. N. WATABE. (With 9 Figures) · 448 25.1 Introduction · 448 25.2 Shell Structure .449 25.3 Chemical Constituents of Shells · 457 25.4 Cellular Aspects of Shell Formation · 464 25.5 Hinge System .472 25.6 Operculum · 475
26 Permeability and Epidermal Transport. M.G. STEWART. (With 2 Figures) · 486
26.1 Introduction · 486 26.2 Permeability in Marine Molluscs · 487 26.3 Permeability in Freshwater Molluscs · 496 26.4 Permeability in Terrestrial Molluscs · 497 26.5 Conclusion · 498
XII
VIII Arthropoda
27 Epidermal Cells. M. LOCKE. (With 7 Figures)
27.1 Introduction ........... . 27.2 The Basic Structure of an Epidermal Cell 27.3 Epidermal Differentiation . . . . . . 27.4 The Cytoskeleton, Cell Movement and
Morphogenetic Shape Change . . . . 27.5 Cuticle Secretion and the Control of Cuticular
Contents
502
503 504 509
511
Compartments ............. 514 27.6 The IntermoultfMoult Sequence of Epidermal
Activity ................ 516 27.7 General Factors Controlling Epidermal Activity 518 27.8 Local Controlling Factors and Cell Communication 518 27.9 Summary and Conclusions ........... 520
28 Chemo-, Hygro-, and Thermoreceptors. R.A. STEINBRECHT. (With 6 Figures) 523
28.1 Introduction .......... ...... 523 28.2 General Morphology of Arthropod Exteroceptors 524 28.3 Structure and Function in Insect Olfactory Receptors 526 28.4 Structure and Function in Insect Gustatory
Receptors . . . . . . . . . . . . . . . . 534 28.5 Structure and Function in Insect Hygro- and
Thermoreceptors . . . . . . . . . 537 28.6 Sensilla in Other Arthropod Classes .... 543 28.7 Conclusions . . . . . . . . . . . . . . . 547
29 Mechanoreception. F.G. BARTH and R. BUCKHAN. (With8 Figures) . 554
29.1 Introduction 554 29.2 Hair Sensilla 555 29.3 Strain Detectors 564 29.4 Conclusion . . 577
30 Cuticle: Biochemistry. R.H. HACKMAN. (With 3 Figures) . . . 583
30.1 Introduction .... 30.2 Cuticular Components 30.3 Stabilization of Cuticles
31 Cuticle: Organization. A.C. NEVILLE.
(With 10 Figures) ... .
31.1 General Structure ... . 31.2 The Size Levels of Structure 31.3 Physical Colours . . . . .
583 584 597
. 611
611 612 623
Contents XIII
32 Cuticle: Mechanical Properties. J.E. HILLERTON. (With 2 Figures) 626
32.1 Introduction 626 32.2 Types of Mechanical Properties Important for
Cuticles 628 32.3 Relationship of Cuticular Structure to Mechanical
Properties . 631 32.4 Relationship of Composition to Mechanical
Properties . 632 32.5 Relationship of Mechanical Properties to Function 633 32.6 Mechanical Properties During Development 634 32.7 Effect of Calcification on Mechanical Properties 635
33 Cuticle: Formation, Moulting and Control. W. GNATZY and F. ROMER. (With 10 Figures) 638
33.1 Introduction 638 33.2 Formation of the Cuticle During Moult-Intermoult
Cycle. 639 33.3 Sequence of Events in the Epidermal Cells 649 33.4 Organogenesis and Moulting of Organules 651 33.5 Moulting Control 664
34 Cuticle: Ecological Significance. N.F. HADLEY. (With 2 Figures) 685
34.1 General Functions-An Overview 685 34.2 The Waterproofing Barrier 686 34.3 Cuticular Lipids and Transpiration 686 34.4 The Cuticle and Water Vapour Absorption 691
35 Cuticle and Insecticides. A.R. GILBY 694
35.1 Introduction 694 35.2 Experimental Techniques 695 35.3 Cuticular Structure 696 35.4 Aspects of Penetration by Insecticides 697
IX Arthropod-Related Phyla
36 Onychophora. V. STORCH. (With 3 Figures) 703
36.1 Introduction 703 36.2 The Cuticle 703 36.3 The Epidermis . 704 36.4 The Epidermal Glands 705 36.5 The Sensilla 705
37 Pentastomida. V. STORCH. (With 2 Figures) 709
37.1 Introduction 709 37.2 The Cuticle 709
XIV
37.3 The Gland Cells and Ionocytes 37.4 The Sensilla . . . . . . . . .
38 Tardigrada. H. GREVEN. (With 6 Figures) 38.1 Introduction ...... . 38.2 Integument . . . . . . . . 38.3 Integument and Anhydrobiosis 38.4 Encystation . . . . . . . 38.5 Integument and Systematics 38.6 Conclusion . . . . . . .
X Minor Coelomate Phyla
Contents
· 711 · 712
· 714 714 715 724 725 725
· 726
39 Lophophorates. A. WILLIAMS. (With 30 Figures) 728 39.1 Introduction 728 39.2 Phoronida 728 39.3 Bryozoa 729 39.4 Brachiopoda 734 39.5 Conclusions . 744
40 Chaetognatha. P. AHNELT. (With 10 Figures) 746
40.1 Introduction ..... 746 40.2 Pluristratified Epidermis 747 40.3 Monolayered Epidermis 750 40.4 Nervous System and Receptors 752 40.5 Conclusion . . . . . . . . . 753
XI Echinodermata
41 Epidermal Cells. N.D. HOLLAND. (With 1 Figure) 756 41.1 Introduction and Definitions 756 41.2 Support Cells 758 41.3 Sensory Cells 761 41.4 Secretory Cells 762 41.5 Other Cell Types 765 41.6 Cuticle 768 41.7 Cell Proliferation and Epidermal Regeneration 769 41.8 Conclusion 770
42 Permeability and Epidermal Transport. D.R. BAMFORD · 775
42.1 Introduction · 775 42.2 The Integument and Osmotic Balance .777 42.3 Absorption of Organic Molecules · 779 42.4 Mechanisms of DOM Influx · 784 42.5 Conclusions · 786
Contents xv
xn The Protochordates
43 Hemichordata. U. WELSCH. (With 7 Figures) 790
43.1 Introduction 790 43.2 Pterobranchia . . . . . . . . . . . . . 790 43.3 Enteropneusta . . . . . . . . . . . . . 793
44 Urochordata. U. WELSCH. (With 8 Figures)
44.1 Introduction 44.2 Ascidiacea 44.3 Thaliacea . 44.4 Larvacea .
45 Cephalochordata. J. BERElTER-HAHN. (With 11 Figures) ....... .
45.1 Introduction . . . . . . . . 45.2 General Organization of the Integument 45.3 Epidermis . . . . . . . . . . 45.4 Cutis and Subcutis . . . . . . 45.5 Development and Regeneration
SUbject Index .
Index to Genera
Abbreviations
LM Light microscope/microscopy EM Electron microscope/microscopy TEM Transmission electron microscope/microscopy SEM Scanning electron microscope/microscopy RER Rough/granular endoplasmic reticulum SER Smooth endoplasmic reticulum MW Molecular weight
800 800 800 810 810
817
817 817 817 823 824
827
837
List of Contributors
You will find the addresses at the beginning of the respective contribution
Ahnelt, P. 746 Bamford, D.R. 775 Barth, F.G. 554 Befus, A.D. 192 Bereiter-Hahn, J. 817 Bird, A.F. 212 Blickhan, R. 554 Bubel, A. 400 Gibson, R. 205 Gilby, A.R. 694 Gnatzy, W. 638 Gomme, J. 323 Green, C.R. 5 Greven, H. 74 Hackman, R.H. 583 Hadley, N.F. 685 Hemandez-Nicaise, M.-L. 96 Hillerton, J.E. 626 Holland, N.D. 756 Hiindgen, M. 47 Locke, M. 502 Mackie, G.O. 43
Mariscal, R.N. 57 Neville, A.C. 611 Podesta, R.B. 192 Richards, K.S. 1, 269, 310 Rieger, R.M. 389 Romer, F. 638 Sanderson, M.J. 17 Schlichter, D. 79 Schroeder, P.C. 297 Southward, E.C. 376 Steinbrecht, R.A. 523 Stewart, M.G. 486 Storch, V. 242,269, 369,
703, 709 Threadgold, L.T. 132 Tidball, J.G. 69 Tyler, S. 112 Watabe, N. 448 Welsch, U. 269, 790, 800 Whitfield, P.J. 234 Williams, A. 728