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Table of Contents
Intro
Sea Urchins: Biology and Ecology
Copyright
Contents
Contributors
Chapter 1: Phylogeny and classification of echinoids
1. Introduction
2. Class Echinoidea Leske, 1778
2.1. Stem group Echinoidea (Paleozoic echinoids)
2.2. Subclass Perischoechinoidea McCoy, 1849 (possibly paraphyletic)
3. Crown group Echinoidea
3.1. Subclass Cidaroidea Smith, 1984
3.2. Subclass Euechinoidea Bronn, 1860
3.2.1. Infraclass Aulodonta Jackson, 1912 (= Diadematacea sensu Mongiardino Koch et al., 2018)
3.2.2. Infraclass Carinacea Kroh and Smith, 2010
3.2.3. Irregularia Latreille, 1825
3.2.4. Stem group Irregularia
3.2.5. Crown group Irregularia
3.2.6. Neognathostomata Smith, 1981
3.2.7. Atelostomata von Zittel, 1879
3.2.8. Order Holasteroida Durham and Melville, 1957
3.2.9. Order Spatangoida Agassiz, 1840
4. Conclusion
References
Further reading
Chapter 2: Sea urchin life-history strategies
1. Concepts
2. Habitats and life-history strategies of sea urchins
2.1. The deep sea
2.2. Antarctic seas
2.3. Tropical reef flats
2.4. Kelp forests
2.5. Tropical seagrass beds
3. Conclusions
Acknowledgments
References
Further reading
Chapter 3: Gametogenesis in regular sea urchins: Structural, functional, and molecular/genomic biology
1. Sea urchin gametogenesis: Structural microenvironment
2. Sea urchin gametogenesis: Annual structural variation
2.1. The dual functionality of sea urchin gonads: Gametogenesis of germinal cells and nutrient storage in somatic NPs
2.2. Nutritive phagocytes in ovaries and testes
2.3. Intergametogenesis and NP phagocytosis in S. droebachiensis
2.4. Pregametogenesis and NP renewal in S. droebachiensis
2.5. Gametogenesis and NP utilization in S. droebachiensis.
2.6. End of gametogenesis, NP exhaustion, and spawning in S. droebachiensis
3. Sea urchin gametogenesis: Annual molecular variation
3.1. Differentiation and identification of oogonia and spermatogonia
3.2. Gamete differentiation including nutrient input (ova) and meiotic division and preparation of gametes for fertilizat ...
3.3. Meiosis in oocytes and spermatocytes
3.4. Preparation of ova and spermatozoa for fertilization
3.5. Recycling NPs and regenerating the gametogenic microenvironment
4. Sea urchin gametogenesis: Environmental control
Acknowledgments
References
Further reading
Chapter 4: Biochemical and energy requirements of gonad development in regular sea urchins
1. Cellular energy utilization
2. Energy metabolism during development
3. Biochemical components of cellular metabolism
4. Gonad growth
5. Gonad energy metabolism
5.1. Protein metabolism
5.2. Carbohydrate metabolism
5.3. Lipid metabolism
5.4. Anaerobic metabolism
6. Feeding and metabolism
7. Summary
Acknowledgments
References
Chapter 5: Endocrine regulation of regular echinoid reproduction
1. Sea urchin gonads
2. Exogenous regulation of reproduction
2.1. Environmental factors
2.2. Endocrine disruptors
3. Endogenous regulation of reproduction
3.1. Steroids
3.1.1. Early studies on sex steroids in the gonads
3.1.2. Steroid-converting enzymes in the gonads
3.1.3. Sex steroids in the gonads
3.1.4. Response to exogenous administration of sex steroids
3.2. Protein and peptidergic factors
3.3. Catecholaminergic and cholinergic factors
4. Mechanisms of regulation
4.1. Paracrine
4.2. Endocrine
5. Gene regulation in reproduction
6. Conclusions
Acknowledgments
References
Chapter 6: Larval ecology of echinoids
1. Introduction
2. Patterns in abundance.
3. Growth and development
4. Mortality and predation
5. Feeding
6. Swimming behavior and vertical distributions
7. Dispersal
8. Settlement
9. Conclusions
References
Chapter 7: Growth and survival of postsettlement sea urchins
1. Growth
1.1. Introduction
1.2. Skeletal composition
1.3. Resorption
1.4. Natural growth lines
1.5. Tagging
1.6. Growth models
2. Survival
Acknowledgments
References
Further reading
Chapter 8: Digestive system in regular sea urchins
1. Introduction
2. Structure and function of the definitive digestive system as a whole
2.1. General anatomy
2.2. General histology of digestive system
2.2.1. Fundamental tissue layers
2.2.2. Ossicle-forming scleroblasts in the wall of the digestive system
2.2.3. Coelomocytes wandering in the wall of the digestive system
2.3. Cell kinetics in the digestive system of regular sea urchins
2.4. Passage of food through the digestive system
3. Major regions of the regular sea urchin digestive system
3.1. Teeth and buccal cavity
3.2. Pharynx
3.3. Esophagus
3.4. Cecum and stomach
3.5. Siphonal grooves and siphons
3.6. Intestine and rectum
3.7. Symbionts in the intestinal lumen
3.7.1. Symbiotic bacteria
3.7.2. Symbiotic protists
3.7.3. Symbiotic metazoans
4. Development of the digestive system
Acknowledgments
References
Further reading
Chapter 9: Ingestion, digestion, and digestibility of regular sea urchins
1. Introduction
2. Ingestion
2.1. Attractants
2.2. Stimulants
2.3. Deterrents
2.4. Environmental conditions
2.4.1. Hydrodynamics
2.4.2. Light
2.4.3. Temperature
2.5. Food shape
2.6. Food quality
2.7. Physiological state
2.7.1. Nutritional state
2.7.2. Body size
2.7.3. Reproductive state
3. Digestion
3.1. Digestive enzymes.
3.2. Role of bacteria
3.2.1. Digestion
3.2.2. Nitrogen fixation
3.3. Gut transit time
4. Digestibility
5. Conclusions
Acknowledgments
References
Further reading
Chapter 10: Nutrition
1. Introduction
1.1. Why nutrition?
1.2. Nutrition vs dietetics
1.3. Historical and contemporary approaches
2. Basic considerations in sea urchin nutrition research
3. Feed development-Dietary nutrient requirements
3.1. Purified, semipurified, and practical ingredients
3.2. Physical feed characteristics
3.3. Toxins in feed ingredients
3.4. Feed management
3.4.1. Feed ration
3.4.2. Feeding rate
3.4.3. Feeding time
3.4.4. Feed ration at each feeding period
3.4.5. Preexposure/sensitization
3.4.6. Group vs individual culture
3.5. Outcome assessment
3.5.1. Ingestion
3.5.2. Digestibility
3.5.3. Survival and growth
3.5.4. Effect of nutrients on organs
3.5.5. Production, production efficiencies, and energetics
3.5.6. Functional or physiological metrics
3.5.7. Role of gut flora
3.5.8. Molecular advances
4. Basic nutrient profiles
4.1. Protein
4.2. Carbohydrate
4.3. Lipids
4.4. Fiber
4.5. Minerals
4.6. Vitamins
4.7. Feed additives
5. Basic applications of feed development
6. Summary
Acknowledgments
References
Further reading
Chapter 11: Carotenoids in sea urchins
1. Introduction
2. Distribution of carotenoids in sea urchins
2.1. Gonad pigments
2.2. Sex differences, eggs, and larvae
2.3. Gut pigments
2.4. Test pigments
2.5. Changes during the reproductive cycle
2.6. Heritability
3. Metabolism of carotenoids in sea urchins
3.1. Isomerization
4. The effect of dietary carotenoids on gonad color and deposition
4.1. Assessing color
4.2. Carotenoid deposition
5. The role of carotenoids in sea urchins.
5.1. Egg production and development
5.2. Biological functions
Acknowledgment
References
Chapter 12: Sea urchin diseases: Effects from individuals to ecosystems
1. Introduction
2. Documented diseases
3. Effects on individuals
3.1. Paramoebiasis
3.2. Vibriosis
3.3. Bald sea urchin disease
4. Effects on populations and ecosystems
References
Chapter 13: Immunology in sea urchins
1. Introduction
1.1. General concepts
1.2. Coelomocytes
1.3. Phagocytosis
1.4. Inflammatory process
1.5. Origin of the coelomocytes
1.6. Coelomic fluid and coelomocyte concentration
1.7. Coelomocytes as biomarkers
1.8. Coagulation and encapsulation
1.9. The complement system
2. Humoral factors
2.1. The sea urchin genome and advances in the immune system studies
References
Further reading
Chapter 14: Deep-sea sea urchins
1. Taxa characteristic of the deep sea
2. Unique morphologies
3. Life-history traits: Growth and reproduction
4. Nutrition, growth, and reproduction
4.1. Food sources, trophic positioning, and nutrient partitioning
4.2. Seasonal dietary shifts
4.3. Bioerosion of deep-sea corals
5. Ecology
5.1. Patterns in distribution, abundance, and habitat associations
5.2. Predation and competition in the deep sea
5.3. Symbionts
6. Evolutionary history
7. Summary
Acknowledgments
References
Chapter 15: Regular sea urchins as drivers of shallow benthic marine community structure
1. Introduction
2. Functional evolution of sea urchins and their rise in ecological importance
3. Agents of biological disturbance, strong interactors, and drivers of community structure
4. Drivers of ecosystem structure and function: Sea grass, coral reef, and kelp forest biome case studies
4.1. Sea grass ecosystems
4.2. Coral reef ecosystems.
4.2.1. Ecological impact of D. antillarum: Revelations from mass mortality.
Sea Urchins: Biology and Ecology
Copyright
Contents
Contributors
Chapter 1: Phylogeny and classification of echinoids
1. Introduction
2. Class Echinoidea Leske, 1778
2.1. Stem group Echinoidea (Paleozoic echinoids)
2.2. Subclass Perischoechinoidea McCoy, 1849 (possibly paraphyletic)
3. Crown group Echinoidea
3.1. Subclass Cidaroidea Smith, 1984
3.2. Subclass Euechinoidea Bronn, 1860
3.2.1. Infraclass Aulodonta Jackson, 1912 (= Diadematacea sensu Mongiardino Koch et al., 2018)
3.2.2. Infraclass Carinacea Kroh and Smith, 2010
3.2.3. Irregularia Latreille, 1825
3.2.4. Stem group Irregularia
3.2.5. Crown group Irregularia
3.2.6. Neognathostomata Smith, 1981
3.2.7. Atelostomata von Zittel, 1879
3.2.8. Order Holasteroida Durham and Melville, 1957
3.2.9. Order Spatangoida Agassiz, 1840
4. Conclusion
References
Further reading
Chapter 2: Sea urchin life-history strategies
1. Concepts
2. Habitats and life-history strategies of sea urchins
2.1. The deep sea
2.2. Antarctic seas
2.3. Tropical reef flats
2.4. Kelp forests
2.5. Tropical seagrass beds
3. Conclusions
Acknowledgments
References
Further reading
Chapter 3: Gametogenesis in regular sea urchins: Structural, functional, and molecular/genomic biology
1. Sea urchin gametogenesis: Structural microenvironment
2. Sea urchin gametogenesis: Annual structural variation
2.1. The dual functionality of sea urchin gonads: Gametogenesis of germinal cells and nutrient storage in somatic NPs
2.2. Nutritive phagocytes in ovaries and testes
2.3. Intergametogenesis and NP phagocytosis in S. droebachiensis
2.4. Pregametogenesis and NP renewal in S. droebachiensis
2.5. Gametogenesis and NP utilization in S. droebachiensis.
2.6. End of gametogenesis, NP exhaustion, and spawning in S. droebachiensis
3. Sea urchin gametogenesis: Annual molecular variation
3.1. Differentiation and identification of oogonia and spermatogonia
3.2. Gamete differentiation including nutrient input (ova) and meiotic division and preparation of gametes for fertilizat ...
3.3. Meiosis in oocytes and spermatocytes
3.4. Preparation of ova and spermatozoa for fertilization
3.5. Recycling NPs and regenerating the gametogenic microenvironment
4. Sea urchin gametogenesis: Environmental control
Acknowledgments
References
Further reading
Chapter 4: Biochemical and energy requirements of gonad development in regular sea urchins
1. Cellular energy utilization
2. Energy metabolism during development
3. Biochemical components of cellular metabolism
4. Gonad growth
5. Gonad energy metabolism
5.1. Protein metabolism
5.2. Carbohydrate metabolism
5.3. Lipid metabolism
5.4. Anaerobic metabolism
6. Feeding and metabolism
7. Summary
Acknowledgments
References
Chapter 5: Endocrine regulation of regular echinoid reproduction
1. Sea urchin gonads
2. Exogenous regulation of reproduction
2.1. Environmental factors
2.2. Endocrine disruptors
3. Endogenous regulation of reproduction
3.1. Steroids
3.1.1. Early studies on sex steroids in the gonads
3.1.2. Steroid-converting enzymes in the gonads
3.1.3. Sex steroids in the gonads
3.1.4. Response to exogenous administration of sex steroids
3.2. Protein and peptidergic factors
3.3. Catecholaminergic and cholinergic factors
4. Mechanisms of regulation
4.1. Paracrine
4.2. Endocrine
5. Gene regulation in reproduction
6. Conclusions
Acknowledgments
References
Chapter 6: Larval ecology of echinoids
1. Introduction
2. Patterns in abundance.
3. Growth and development
4. Mortality and predation
5. Feeding
6. Swimming behavior and vertical distributions
7. Dispersal
8. Settlement
9. Conclusions
References
Chapter 7: Growth and survival of postsettlement sea urchins
1. Growth
1.1. Introduction
1.2. Skeletal composition
1.3. Resorption
1.4. Natural growth lines
1.5. Tagging
1.6. Growth models
2. Survival
Acknowledgments
References
Further reading
Chapter 8: Digestive system in regular sea urchins
1. Introduction
2. Structure and function of the definitive digestive system as a whole
2.1. General anatomy
2.2. General histology of digestive system
2.2.1. Fundamental tissue layers
2.2.2. Ossicle-forming scleroblasts in the wall of the digestive system
2.2.3. Coelomocytes wandering in the wall of the digestive system
2.3. Cell kinetics in the digestive system of regular sea urchins
2.4. Passage of food through the digestive system
3. Major regions of the regular sea urchin digestive system
3.1. Teeth and buccal cavity
3.2. Pharynx
3.3. Esophagus
3.4. Cecum and stomach
3.5. Siphonal grooves and siphons
3.6. Intestine and rectum
3.7. Symbionts in the intestinal lumen
3.7.1. Symbiotic bacteria
3.7.2. Symbiotic protists
3.7.3. Symbiotic metazoans
4. Development of the digestive system
Acknowledgments
References
Further reading
Chapter 9: Ingestion, digestion, and digestibility of regular sea urchins
1. Introduction
2. Ingestion
2.1. Attractants
2.2. Stimulants
2.3. Deterrents
2.4. Environmental conditions
2.4.1. Hydrodynamics
2.4.2. Light
2.4.3. Temperature
2.5. Food shape
2.6. Food quality
2.7. Physiological state
2.7.1. Nutritional state
2.7.2. Body size
2.7.3. Reproductive state
3. Digestion
3.1. Digestive enzymes.
3.2. Role of bacteria
3.2.1. Digestion
3.2.2. Nitrogen fixation
3.3. Gut transit time
4. Digestibility
5. Conclusions
Acknowledgments
References
Further reading
Chapter 10: Nutrition
1. Introduction
1.1. Why nutrition?
1.2. Nutrition vs dietetics
1.3. Historical and contemporary approaches
2. Basic considerations in sea urchin nutrition research
3. Feed development-Dietary nutrient requirements
3.1. Purified, semipurified, and practical ingredients
3.2. Physical feed characteristics
3.3. Toxins in feed ingredients
3.4. Feed management
3.4.1. Feed ration
3.4.2. Feeding rate
3.4.3. Feeding time
3.4.4. Feed ration at each feeding period
3.4.5. Preexposure/sensitization
3.4.6. Group vs individual culture
3.5. Outcome assessment
3.5.1. Ingestion
3.5.2. Digestibility
3.5.3. Survival and growth
3.5.4. Effect of nutrients on organs
3.5.5. Production, production efficiencies, and energetics
3.5.6. Functional or physiological metrics
3.5.7. Role of gut flora
3.5.8. Molecular advances
4. Basic nutrient profiles
4.1. Protein
4.2. Carbohydrate
4.3. Lipids
4.4. Fiber
4.5. Minerals
4.6. Vitamins
4.7. Feed additives
5. Basic applications of feed development
6. Summary
Acknowledgments
References
Further reading
Chapter 11: Carotenoids in sea urchins
1. Introduction
2. Distribution of carotenoids in sea urchins
2.1. Gonad pigments
2.2. Sex differences, eggs, and larvae
2.3. Gut pigments
2.4. Test pigments
2.5. Changes during the reproductive cycle
2.6. Heritability
3. Metabolism of carotenoids in sea urchins
3.1. Isomerization
4. The effect of dietary carotenoids on gonad color and deposition
4.1. Assessing color
4.2. Carotenoid deposition
5. The role of carotenoids in sea urchins.
5.1. Egg production and development
5.2. Biological functions
Acknowledgment
References
Chapter 12: Sea urchin diseases: Effects from individuals to ecosystems
1. Introduction
2. Documented diseases
3. Effects on individuals
3.1. Paramoebiasis
3.2. Vibriosis
3.3. Bald sea urchin disease
4. Effects on populations and ecosystems
References
Chapter 13: Immunology in sea urchins
1. Introduction
1.1. General concepts
1.2. Coelomocytes
1.3. Phagocytosis
1.4. Inflammatory process
1.5. Origin of the coelomocytes
1.6. Coelomic fluid and coelomocyte concentration
1.7. Coelomocytes as biomarkers
1.8. Coagulation and encapsulation
1.9. The complement system
2. Humoral factors
2.1. The sea urchin genome and advances in the immune system studies
References
Further reading
Chapter 14: Deep-sea sea urchins
1. Taxa characteristic of the deep sea
2. Unique morphologies
3. Life-history traits: Growth and reproduction
4. Nutrition, growth, and reproduction
4.1. Food sources, trophic positioning, and nutrient partitioning
4.2. Seasonal dietary shifts
4.3. Bioerosion of deep-sea corals
5. Ecology
5.1. Patterns in distribution, abundance, and habitat associations
5.2. Predation and competition in the deep sea
5.3. Symbionts
6. Evolutionary history
7. Summary
Acknowledgments
References
Chapter 15: Regular sea urchins as drivers of shallow benthic marine community structure
1. Introduction
2. Functional evolution of sea urchins and their rise in ecological importance
3. Agents of biological disturbance, strong interactors, and drivers of community structure
4. Drivers of ecosystem structure and function: Sea grass, coral reef, and kelp forest biome case studies
4.1. Sea grass ecosystems
4.2. Coral reef ecosystems.
4.2.1. Ecological impact of D. antillarum: Revelations from mass mortality.