Sea Urchins

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Sea Urchins: Biology and Ecology, Fourth Edition, Volume 43 expands its coverage to include the entire class of Echinoidea, making this new edition an authoritative reference of the entire class of species. This is a valuable resource that will help readers gain a deep understanding of the basic characteristics of sea urchins, the basis of the great variation that exists in sea urchins, and how sea urchins are important components of marine ecosystems. Updated coverage includes sections on reproduction, metabolism, endocrinology, larval ecology, growth, digestion, carotenoids and disease.

Includes pertinent tables and graphs within chapters to visually summarize information
Provides case studies with research applications to provide potential solutions
Includes the entire class of Echinoidea and the effect of climate change on the biology and ecology of the species

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Front Cover 2
Sea Urchins: Biology and Ecology 5
Copyright 6
Contents 7
Contributors 17
Chapter 1: Phylogeny and classification of echinoids 19
- 1. Introduction 19
- 2. Class Echinoidea Leske, 1778 19
  - 2.1. Stem group Echinoidea (Paleozoic echinoids) 19
  - 2.2. Subclass Perischoechinoidea McCoy, 1849 (possibly paraphyletic) 20
- 3. Crown group Echinoidea 20
  - 3.1. Subclass Cidaroidea Smith, 1984 21
  - 3.2. Subclass Euechinoidea Bronn, 1860 22
    - 3.2.1. Infraclass Aulodonta Jackson, 1912 (= Diadematacea sensu Mongiardino Koch et al., 2018) 23
    - 3.2.2. Infraclass Carinacea Kroh and Smith, 2010 24

Front Cover 2
Sea Urchins: Biology and Ecology 5
Copyright 6
Contents 7
Contributors 17
Chapter 1: Phylogeny and classification of echinoids 19
- 1. Introduction 19
- 2. Class Echinoidea Leske, 1778 19
  - 2.1. Stem group Echinoidea (Paleozoic echinoids) 19
  - 2.2. Subclass Perischoechinoidea McCoy, 1849 (possibly paraphyletic) 20
- 3. Crown group Echinoidea 20
  - 3.1. Subclass Cidaroidea Smith, 1984 21
  - 3.2. Subclass Euechinoidea Bronn, 1860 22
    - 3.2.1. Infraclass Aulodonta Jackson, 1912 (= Diadematacea sensu Mongiardino Koch et al., 2018) 23
    - 3.2.2. Infraclass Carinacea Kroh and Smith, 2010 24
    - 3.2.3. Irregularia Latreille, 1825 25
    - 3.2.4. Stem group Irregularia 26
    - 3.2.5. Crown group Irregularia 26
    - 3.2.6. Neognathostomata Smith, 1981 27
    - 3.2.7. Atelostomata von Zittel, 1879 28
    - 3.2.8. Order Holasteroida Durham and Melville, 1957 28
    - 3.2.9. Order Spatangoida Agassiz, 1840 29
  - 4. Conclusion 30
  - References 31
  - Further reading 35
- Chapter 2: Sea urchin life-history strategies 37
  - 1. Concepts 37
  - 2. Habitats and life-history strategies of sea urchins 40
    - 2.1. The deep sea 40
    - 2.2. Antarctic seas 41
    - 2.3. Tropical reef flats 42
    - 2.4. Kelp forests 42
    - 2.5. Tropical seagrass beds 43
  - 3. Conclusions 43
  - Acknowledgments 44
  - References 44
  - Further reading 46
- Chapter 3: Gametogenesis in regular sea urchins: Structural, functional, and molecular/genomic biology 47
  - 1. Sea urchin gametogenesis: Structural microenvironment 47
  - 2. Sea urchin gametogenesis: Annual structural variation 48
    - 2.1. The dual functionality of sea urchin gonads: Gametogenesis of germinal cells and nutrient storage in somatic NPs 48
    - 2.2. Nutritive phagocytes in ovaries and testes 48
    - 2.3. Intergametogenesis and NP phagocytosis in S. droebachiensis 50
    - 2.4. Pregametogenesis and NP renewal in S. droebachiensis 50
    - 2.5. Gametogenesis and NP utilization in S. droebachiensis 51
    - 2.6. End of gametogenesis, NP exhaustion, and spawning in S. droebachiensis 52
  - 3. Sea urchin gametogenesis: Annual molecular variation 52
    - 3.1. Differentiation and identification of oogonia and spermatogonia 53
    - 3.2. Gamete differentiation including nutrient input (ova) and meiotic division and preparation of gametes for fertilizat ... 54
    - 3.3. Meiosis in oocytes and spermatocytes 56
    - 3.4. Preparation of ova and spermatozoa for fertilization 57
    - 3.5. Recycling NPs and regenerating the gametogenic microenvironment 58
  - 4. Sea urchin gametogenesis: Environmental control 59
  - Acknowledgments 62
  - References 62
  - Further reading 68
- Chapter 4: Biochemical and energy requirements of gonad development in regular sea urchins 69
  - 1. Cellular energy utilization 69
  - 2. Energy metabolism during development 70
  - 3. Biochemical components of cellular metabolism 72
  - 4. Gonad growth 72
  - 5. Gonad energy metabolism 74
    - 5.1. Protein metabolism 74
    - 5.2. Carbohydrate metabolism 76
    - 5.3. Lipid metabolism 77
    - 5.4. Anaerobic metabolism 77
  - 6. Feeding and metabolism 78
  - 7. Summary 79
  - References 80
  - Acknowledgments 80
- Chapter 5: Endocrine regulation of regular echinoid reproduction 83
  - 2. Exogenous regulation of reproduction 83
    - 2.2. Endocrine disruptors 83
    - 2.1. Environmental factors 83
  - 1. Sea urchin gonads 83
  - 3. Endogenous regulation of reproduction 84
    - 3.1. Steroids 85
      - 3.1.1. Early studies on sex steroids in the gonads 85
      - 3.1.2. Steroid-converting enzymes in the gonads 85
      - 3.1.4. Response to exogenous administration of sex steroids 88
      - 3.1.3. Sex steroids in the gonads 88
    - 3.2. Protein and peptidergic factors 89
    - 3.3. Catecholaminergic and cholinergic factors 90
  - 4. Mechanisms of regulation 90
    - 4.1. Paracrine 90
    - 4.2. Endocrine 91
  - 5. Gene regulation in reproduction 91
  - 6. Conclusions 91
  - Acknowledgments 92
  - References 92
- Chapter 6: Larval ecology of echinoids 95
  - 1. Introduction 95
  - 2. Patterns in abundance 95
  - 3. Growth and development 96
  - 4. Mortality and predation 99
  - 5. Feeding 99
  - 6. Swimming behavior and vertical distributions 101
  - 7. Dispersal 102
  - 8. Settlement 104
  - 9. Conclusions 105
  - References 105
- Chapter 7: Growth and survival of postsettlement sea urchins 113
  - 1. Growth 113
    - 1.1. Introduction 113
    - 1.2. Skeletal composition 114
    - 1.3. Resorption 121
    - 1.4. Natural growth lines 122
    - 1.5. Tagging 125
    - 1.6. Growth models 129
  - 2. Survival 140
  - Acknowledgments 153
  - References 153
  - Further reading 163
- Chapter 8: Digestive system in regular sea urchins 165
  - 1. Introduction 165
  - 2. Structure and function of the definitive digestive system as a whole 165
    - 2.2. General histology of digestive system 165
      - 2.2.1. Fundamental tissue layers 165
      - 2.2.2. Ossicle-forming scleroblasts in the wall of the digestive system 166
      - 2.2.3. Coelomocytes wandering in the wall of the digestive system 168
    - 2.1. General anatomy 165
    - 2.3. Cell kinetics in the digestive system of regular sea urchins 168
    - 2.4. Passage of food through the digestive system 170
  - 3. Major regions of the regular sea urchin digestive system 170
    - 3.1. Teeth and buccal cavity 170
    - 3.2. Pharynx 171
    - 3.5. Siphonal grooves and siphons 173
    - 3.3. Esophagus 173
    - 3.4. Cecum and stomach 173
    - 3.7. Symbionts in the intestinal lumen 175
      - 3.7.1. Symbiotic bacteria 175
      - 3.7.3. Symbiotic metazoans 176
      - 3.7.2. Symbiotic protists 176
    - 3.6. Intestine and rectum 175
    - 4. Development of the digestive system 177
    - References 179
    - Acknowledgments 179
    - Further reading 181
  - Chapter 9: Ingestion, digestion, and digestibility of regular sea urchins 183
    - 1. Introduction 183
    - 2. Ingestion 183
      - 2.1. Attractants 183
      - 2.2. Stimulants 184
      - 2.3. Deterrents 184
      - 2.4. Environmental conditions 186
        
        2.4.2. Light 186
        
        2.4.3. Temperature 186
        
        2.4.1. Hydrodynamics 186
      - 2.6. Food quality 187
      - 2.7. Physiological state 187
        
        2.7.1. Nutritional state 187
        
        2.7.2. Body size 188
        
        2.7.3. Reproductive state 188
      - 2.5. Food shape 187
      - 3. Digestion 188
        
        3.1. Digestive enzymes 189
        
        3.2. Role of bacteria 190
        
        3.2.1. Digestion 190
        
        3.2.2. Nitrogen fixation 191
        
        3.3. Gut transit time 191
      - 4. Digestibility 192
      - Acknowledgments 201
      - References 201
      - 5. Conclusions 201
      - Further reading 208
    - Chapter 10: Nutrition 209
      - 1. Introduction 209
        
        1.2. Nutrition vs dietetics 209
        
        1.1. Why nutrition? 209
        
        1.3. Historical and contemporary approaches 210
      - 2. Basic considerations in sea urchin nutrition research 210
      - 3. Feed development-Dietary nutrient requirements 211
        
        3.1. Purified, semipurified, and practical ingredients 211
        
        3.2. Physical feed characteristics 212
        
        3.3. Toxins in feed ingredients 212
        
        3.4. Feed management 212
        
        3.4.1. Feed ration 212
        
        3.4.3. Feeding time 213
        
        3.4.6. Group vs individual culture 213
        
        3.4.4. Feed ration at each feeding period 213
        
        3.4.2. Feeding rate 213
        
        3.4.5. Preexposure/sensitization 213
        
        3.5. Outcome assessment 214
        
        3.5.2. Digestibility 214
        
        3.5.3. Survival and growth 214
        
        3.5.1. Ingestion 214
        
        3.5.4. Effect of nutrients on organs 215
        
        3.5.5. Production, production efficiencies, and energetics 215
        
        3.5.7. Role of gut flora 216
        
        3.5.8. Molecular advances 216
        
        3.5.6. Functional or physiological metrics 216
        
        4. Basic nutrient profiles 216
        
        4.1. Protein 217
        
        4.2. Carbohydrate 218
        
        4.3. Lipids 219
        
        4.4. Fiber 220
        
        4.5. Minerals 220
        
        4.6. Vitamins 221
        
        4.7. Feed additives 221
        
        References 222
        
        6. Summary 222
        
        Acknowledgments 222
        
        5. Basic applications of feed development 222
        
        Further reading 226
      - Chapter 11: Carotenoids in sea urchins 227
        
        1. Introduction 227
        
        2. Distribution of carotenoids in sea urchins 227
        
        2.2. Sex differences, eggs, and larvae 227
        
        2.1. Gonad pigments 227
        
        2.4. Test pigments 230
        
        2.5. Changes during the reproductive cycle 230
        
        2.3. Gut pigments 230
        
        2.6. Heritability 231
        
        3. Metabolism of carotenoids in sea urchins 231
        
        3.1. Isomerization 231
        
        4. The effect of dietary carotenoids on gonad color and deposition 231
        
        4.1. Assessing color 231
        
        4.2. Carotenoid deposition 232
        
        5. The role of carotenoids in sea urchins 233
        
        5.1. Egg production and development 233
        
        5.2. Biological functions 233
        
        Acknowledgment 233
        
        References 234
      - Chapter 12: Sea urchin diseases: Effects from individuals to ecosystems 237
        
        1. Introduction 237
        
        2. Documented diseases 238
        
        3. Effects on individuals 241
        
        3.1. Paramoebiasis 241
        
        3.2. Vibriosis 241
        
        3.3. Bald sea urchin disease 242
        
        4. Effects on populations and ecosystems 242
        
        References 243
      - Chapter 13: Immunology in sea urchins 245
        
        1. Introduction 245
        
        1.1. General concepts 245
        
        1.2. Coelomocytes 246
        
        1.4. Inflammatory process 247
        
        1.3. Phagocytosis 247
        
        1.5. Origin of the coelomocytes 248
        
        1.7. Coelomocytes as biomarkers 249
        
        1.6. Coelomic fluid and coelomocyte concentration 249
        
        1.9. The complement system 250
        
        1.8. Coagulation and encapsulation 250
        
        2. Humoral factors 250
        
        2.1. The sea urchin genome and advances in the immune system studies 251
        
        References 251
        
        Further reading 254
      - Chapter 14: Deep-sea sea urchins 255
        
        1. Taxa characteristic of the deep sea 255
        
        2. Unique morphologies 256
        
        3. Life-history traits: Growth and reproduction 258
        
        4. Nutrition, growth, and reproduction 259
        
        4.1. Food sources, trophic positioning, and nutrient partitioning 259
        
        4.2. Seasonal dietary shifts 260
        
        4.3. Bioerosion of deep-sea corals 261
        
        5. Ecology 262
        
        5.1. Patterns in distribution, abundance, and habitat associations 262
        
        5.3. Symbionts 265
        
        5.2. Predation and competition in the deep sea 265
        
        6. Evolutionary history 266
        
        Acknowledgments 267
        
        References 267
        
        7. Summary 267
      - Chapter 15: Regular sea urchins as drivers of shallow benthic marine community structure 273
        
        1. Introduction 273
        
        2. Functional evolution of sea urchins and their rise in ecological importance 273
        
        3. Agents of biological disturbance, strong interactors, and drivers of community structure 275
        
        4. Drivers of ecosystem structure and function: Sea grass, coral reef, and kelp forest biome case studies 275
        
        4.1. Sea grass ecosystems 276
        
        4.2. Coral reef ecosystems 277
        
        4.2.1. Ecological impact of D. antillarum: Revelations from mass mortality 277
        
        4.2.2. Cascading processes from herbivory by D. antillarum affecting productivity, competition, and coral recruitment 278
        
        4.2.3. Predation 280
        
        4.2.5. Ecological impacts to coral reefs from other species of Diadema 281
        
        4.2.4. Alternative stable state 281
        
        4.2.6. Echinometra: Locally important tropical grazers and bioeroders 282
        
        4.3. Kelp forest ecosystems 282
        
        4.3.1. Patterns, processes, and mechanisms of kelp forest: Barrens dynamics 283
        
        4.3.2. Temporal and spatial trends of expanding sea urchin barrens 284
        
        4.3.3. Predation 285
        
        4.3.4. Trophic cascades: General ecological theory from predator-sea urchin-kelp forest interactions 286
        
        5. Convergent patterns and process: Conclusions, synthesis, and speculation 286
        
        5.1. Adaptive breakthrough, phyletic constraints: The rise and fall of sea urchins 287
        
        5.2. Consequences of direct predator impacts 288
        
        5.3. Indirect predator impacts: Trophic cascades 288
        
        5.4. Fisheries as drivers of booms and busts in sea urchin populations 289
        
        5.5. Trophic level dysfunction, climate change, and the stability of alternative community states 289
        
        5.6. The longer view of past, present, and future 291
        
        Acknowledgments 291
        
        References 291
        
        Chapter 16: Sea urchins in a high CO2 world: Impacts of climate warming and ocean acidification across life history stages 299
        
        1. Introduction 299
        
        2. Changes in sea urchin distribution 299
        
        3. Global warming 300
        
        3.1. Sea urchin development in a warming ocean 300
        
        3.2. Effects of habitat warming on the benthic life stage of sea urchins 302
        
        4. Ocean acidification-Multiple drivers 302
        
        4.1. OA impacts on the planktonic sea urchin life phase 303
        
        4.2. OA impacts on the benthic sea urchin life phase 304
        
        5. Sea urchins living at natural acidification 305
        
        6. Multistressor effects-Combined warming and acidification 308
        
        7. Potential for climate adaptation 310
        
        8. Conclusions 310
        
        References 311
        
        Acknowledgments 311

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