Annelids in Modern Biology

Contributors.

Part I Annelids as Model Systems in Biology.

1. Developing Models for Lophotrochozoan and Annelid Biology (Kenneth M. Halanych and Elizabeth Borda).

1.1 Introduction.

1.2 Phylogenetic Considerations.

1.3 Genetic and Developmental Tools.

1.4 Annelid Model Organisms.

1.5 Other Potential Annelid Models.

2. Annelid Phylogeny—Molecular Analysis with an Emphasis on Model Annelids (Christoph Bleidorn).

2.1 Introduction.

2.2 Genes.

2.3 Molecular Annelid Phylogeny.

2.4 Choosing Model Organisms.

2.5 Branch Lengths.

2.6 Problems in Inferring Annelid Phylogeny.

2.7 Conclusions.

3. Cryptic Speciation in Clitellate Model Organisms (Christer Erséus and Daniel Gustafsson).

3.1 Introduction.

3.2 Sources and Kinds of Variation.

3.3 Examples of Clitellate Model Organisms.

3.4 Cryptic Speciation.

3.5 Conclusions and Recommendations.

4. Annelid Life Cycle Cultures (Donald J. Reish and Bruno Pernet).

4.1 Introduction.

4.2 Criteria for the Selection of Species.

4.3 Summary of Culture Techniques.

4.4 Life Cycle Cultures of Polychaeta.

4.5 Life Cycle Cultures of Oligochaeta.

4.6 Life Cycle Cultures of Hirudinea (Leeches).

Part II Evolution and Development.

5. Annelids in Evolutionary Developmental Biology (Dian-Han Kuo).

5.1 Introduction.

5.2 Evo-Devo Today.

5.3 Evo-Devo as Comparative Biology.

5.4 Why Annelid Development Is Interesting for Metazoan Evo-Devo Biologists.

5.5 Case Study 1: Segmentation.

5.6 Case Study 2: Spiral Cleavage and Axis Specifi cation.

5.7 Tools for Analyzing Molecular Mechanisms of Development.

5.8 The Future of the Annelid Model Systems for Evo-Devo.

6. Evolution, Development and Ecology of Capitella sp. I: A Waxing Model for Polychaete Studies (Susan D. Hill and Robert M. Savage).

6.1 Introduction.

6.2 Speciation Studies.

6.3 Capitella Sp. 1 Morphology.

6.4 Replacement of Lost Segments and Reproductive Trade-Offs.

6.5 Metatrochophores, Ciliary Bands and Musculature.

6.6 Gene Expression during the Specifi cation and Differentiation of Germ Layers.

6.7 Sex among the Vermes.

6.8 Annelids and the Segmentation Debate.

6.9 A-P Polarity—Hox and ParaHox Genes.

6.10 Annelid Genomics: Draft Genome Sequence.

6.11 The Future—Where Is This Going?

7. Stem Cell Genesis and Differentiation in Leech (Shirley A. Lang and Daniel H. Shain).

7.1 Introduction.

7.2 Stem Cell Genesis and Development.

7.3 Factors Affecting Stem Cell Genesis.

7.4 Stem Cell Differentiation.

7.5 Gene Expression.

7.6 Conclusion.

Part III Neurobiology and Regeneration.

8. Cellular and Behavioral Properties of Learning in Leech and Other Annelids (Kevin M. Crisp and Brian D. Burrell).

8.1 Introduction.

8.2 Learning in the Leech Whole-Body Shortening Refl ex and Role of the S Interneuron.

8.3 Role of the S Interneuron: Modulation of Excitability.

8.4 Learning in the Leech Swim Circuit.

8.5 Using the Leech to Study Intrinsic Forms of Sensitization.

8.6 Synaptic Plasticity in Leech CNS.

8.7 Conclusions.

9. Development, Regeneration and Immune Responses of the Leech Nervous System (Michel Salzet and Eduardo Macagno).

9.1 Introduction.

9.2 Background.

9.3 Recent Work on the Development of the Nervous System.

9.4 Neuronal Regeneration and Repair.

9.5 Neuroimmune Responses.

9.6 Cellular and Humoral Immune Mechanisms: A Leech Innate Immune Response.

9.7 Conclusions and Future Directions.

10. Lumbriculus variegatus and the Need for Speed: A Model System for Rapid Escape, Regeneration and Asexual Reproduction (Mark J. Zoran and Veronica G. Martinez).

10.1 Introduction.

10.2 Neural Regeneration in Oligochaetes.

10.3 Lumbriculus variegatus, a Model System for Regeneration and Asexual Reproduction.

10.4 Neural Morphallaxis.

10.5 Accessible Model for Life Science Education.

Part IV Environmental and Ecological Studies.

11. Polychaetes in Environmental Studies (Victoria Díaz-Castañeda and Donald J. Reish).

11.1 Introduction.

11.2 Estuarine Occurrence.

11.3 Intertidal Occurrence.

11.4 Mussel Beds.

11.5 Sea Grasses.

11.6 Sabellarid and Serpulid Reefs.

11.7 Benthic Community Structure.

11.8 Unusual Benthic Habitats.

11.9 Feeding Guilds.

11.10 Algal “Gardening” Behavior.

11.11 Polychaetes as Environmental Indicators and Remediators.

11.12 Biomonitoring.

11.13 Toxicological Tests.

11.14 Economic Importance of Polychaetes.

11.15 Conclusions.

12. Oligochaete Worms for Ecotoxicological Assessment of Soils and Sediments (Jörg Römbke and Philipp Egeler).

12.1 Introduction.

12.2 Principles of Environmental Risk Assessment.

12.3 Soil Tests with Lumbricidae.

12.4 Soil Tests with Enchytraeidae.

12.5 Sediment Tests with Lumbriculidae and Tubifi cidae.

12.6 Oligochaetes in Ecotoxicology.

12.7 Conclusions.

13. Evolution and Ecology of Ophryotrocha (Dorvilleidae, Eunicida) (Daniel J. Thornhill, Thomas G. Dahlgren, and Kenneth M. Halanych).

13.1 Introduction.

13.2 General Morphology.

13.3 Taxonomic and Phylogenetic Considerations.

13.4 Reproductive Biology.

13.5 Ecology.

13.6 Future Research.

14. Cosmopolitan Earthworms—A Global and Historical Perspective (Robert J. Blakemore).

14.1 Introduction.

14.2 Number of Earthworm Species.

14.3 Characteristics and Origins of Cosmopolitan Earthworms.

14.4 Overview of Results.

14.5 Discussion.

14.6 Regional Species Totals and Proportions of Exotics.

14.7 Earthworms, Archaeology and Human History.

14.8 Benefi ts and Risks of Earthworm Transportations.

14.9 Conclusions.

Part V Extreme Environments and Biological Novelties.

15. Hydrothermal Vent Annelids (Florence Pradillon and Françoise Gaill).

15.1 Introduction.

15.2 Alvinella pompejana: a Symbiotic System.

15.3 Temperature Adaptation.

15.4 Temperature Adaptation at a Molecular Level.

15.5 Alvinella Tubes.

15.6 Collagens.

15.7 Temperature Adaptation at a Cellular Level: the Case of Developing Embryos.

15.8 Behavioral Adaptation to a High-Temperature Environment.

15.9 Future Development of Thermal Adaptation Studies.

15.10 Perspectives.

16. Glacier Ice Worms (Paula L. Hartzell and Daniel H. Shain).

16.1 Introduction.

16.2 Natural History.

16.3 Classifi cation and Phylogenetic Relationships.

16.4 Origins.

16.5 Clades.

16.6 Physiology.

16.7 Conservation Status.

17. Sperm Ultrastructure in Assessing Phylogenetic Relationships among Clitellate Annelids (Roberto Marotta and Marco Ferraguti).

17.1 Introduction.

17.2 The Spermatozoon of Propappus volki (Michaelsen 1916).

17.3 Sperm Ultrastructure in Branchiobdellids, Acanthobdella peledina, and Hirudineans.

17.4 Sperm Ultrastructure inside Tubifi cidae.

17.5 Plesiomorphic Spermatozoon of Clitellates and Spermatological Apomorphic Trends.

17.6 Patterns of Spermatological Characters among Clitellates.

18. Clitellate Cocoons and Their Secretion (Jonelle Coleman and Daniel H. Shain).

18.1 Introduction.

18.2 Reproductive Biology.

18.3 Clitellum and CGCs.

18.4 Cocoon Production.

18.5 Brooding Behavior within Glossiphoniidae.

18.6 Cocoon Structure: Surface Topology and Ultrastructural Properties.

18.7 Evolution of Clitellate Cocoons and Their Secretion.

18.8 Biomaterials Applications.

Index.