Description Sea urchins are a major component of the world ocean. They are important ecologically and often greatly affect marine communities. They
have an excellent fossil record and consequently are of interest to paleontologists. Research has increased in recent years stimulated
first by a recognition of their ecological importance and then because of their economic importance. Scientists around the world are
actively investigating their potential for aquaculture.
This book is designed to provide a broad understanding of the biology and ecology
of sea urchins. Synthetic chapters consider biology of sea urchins as a whole to give a broad view. The topics of these chapters include
reproduction, metabolism, endocrinology, larval ecology, growth, digestion, carotenoids, disease and nutrition. Subsequent chapters consider
the ecology of individual species that are of major importance ecologically and economically. These include species from Japan, New Zealand,
Australia, Europe, North America, South America and Africa.
Audience
Marine biologists, students and researchers studying aquaculture, fisheries, and developmental biology
Contents Chapter 1. The edible sea urchins
John M. Lawrence
1. Prehistory fishing of sea urchins
2. Contemporary commercial fishing of sea urchins
3. Aquaculture of sea urchins
4. The edible sea urchins
5. Life-history strategies of sea urchins
6. Conclusion
References
Chapter 2.
Gametogenesis and reproduction of sea urchins
Charles W. Walker, Tatsuya Unuma, Michael P. Lesser
1. Introduction
2. Structure of the
gonads of the sea urchin
3. Interacting gametogenic and nutritive phagocyte cycles in the sea urchin gonad: stages, physiology and molecular
biology
3.1. Stages in gametogenesis
3.2. Nutritive phagocytes in ovaries and testes
3.3. Inter-gametogenesis and NP phagocytosis
3.4.
Pre-gametogenesis and NP renewal
3.5. Gametogenesis and NP utilization
3.6. End of gametogenesis: NP exhaustion and spawning
3.7. Environmental
control of gametogenesis
4. Conclusions
References
Chapter 3. Biochemical and energy requirements of gonad development
Adam G. Marsh,
Stephen A. Watts
1. Introduction
2. Cellular energy utilization
3. Energy metabolism during development
4. Biochemical components of
cellular metabolism
5. Gonad growth
6. Gonad energy metabolism
6.1. Protein metabolism
6.2. Carbohydrate metabolism
6.3. Anaerobic metabolism
7. Feeding and metabolism
8. Summary
References
Chapter 4. Reproductive endocrinology of sea urchins
Kristina M. Wasson, Stephen A.
Watts
1. Sea urchin gonad
2. Exogenous regulation of reproduction
2.1. Environmental factors
2.2. Endocrine disruptors
3. Endogenous
regulation of reproduction
3.1. Steroids
3.1.1. 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
References
Chapter 5. Echinoid larval ecology
Larry R. McEdward, Benjamin G. Miner
1. Introduction
2. The echinoid life cycle
3. Life cycle diversity
4. Echinoid larval diversity
5. Reproductive ecology
5.1. Egg provisioning
5.2. Fertilization ecology
6. Larval ecology
6.1. Reproductive
strategies
6.2. Feeding
6.3. Larval growth
6.4. Phenotypic plasticity
6.5. Swimming
6.6 Mortality and defense
7. Recruitment ecology
7.1. Larval transport and dispersal
7.2. Settlement
7.3. Metamorphosis and recruitment
8. Conclusions
References
Chapter 6. Growth and
survival of post-settlement sea urchins
Thomas A. Ebert
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
References
Chapter 7. Feeding, digestion and digestibility
John M. Lawrence,
Addison L. Lawrence, Stephen A. Watts
1. Ingestion
1.1. Food chemistry
1.1.1. Attractants
1.1.2. Stimulants and deterrents
1.2. Environmental
conditions
1.2.1. Hydrodynamics
1.2.2. Light
l.2.3. Temperature
1.3. Food shape
1.4. Physiological state
1.4.1. Nutritional state
1.4.2.
Body size
1.4.3. Reproductive state
2. Digestion
2.1. Structure of the gut
2.2. Digestive enzymes
2.3. Gut transit time
2.4. Role of
microorganisms in digestion
3. Digestibility
4. Conclusions
References
Chapter 8. Carotenoids in sea urchins
Miyuki Tsushima
1. Introduction
2. Carotenoids distribution in sea urchins
3. Metabolism of carotenoids in sea urchins
4. The effect of dietary carotenoids on gonad
color in sea urchins
5. The role of carotenoids in sea urchins
5.1. Egg production and development
5.2. Biological functions
6. Conclusions
References
Chapter 9. Disease in sea urchins
K. Tajima, J.R.M.C. Silva, J.M. Lawrence
1. Introduction
2. Bacterial disease in Japanese
sea urchins
2.1. Evidence for bacterial disease
2.2. Symptoms of the disease
2.3. Isolation and description of the disease-causing bacterium
2.3.1. Summer disease
2.3.2. Spring disease
2.4. Biological responses to bacterial infection
2.5. Bacterial control
3. Immunological
response to bacterial diseases in sea urchins
3.1. General concept of the immune response
3.2. Coelomocyte types
3.3. Phagocytosis
3.4.
Inflammatory process
3.5. Origin of the coelomocytes
3.6. Coelomic fluid and coelomocyte concentration
3.7. Coagulation and encapsulation
3.8. The complement system and humoral factors
4. Conclusions
References
Color Plates
Chapter 10. Ecology of Centrostephanus
N. Andrew,
M. Byrne
1. Biogeography
2. Ecological impacts
2.1. Habitat structure
2.2. Food and feeding ecology
3. Population regulation
3.1. Recruitment
3.2. Predation and disease
3.3. Competition
3.4. Physical factors
4. Reproduction
4.1. Reproductive cycle
4.2. Habitat related patterns
4.2. Development and larval ecology
5. Growth and age
6. Conclusions
References
Chapter 11. Ecology of Diadema
T. R. McClanahan, N.
Muthiga
1. Introduction
1.1. Species of Diadema
1.2. Biogeography and large-scale distribution
1.3. Local distribution and abundance
patterns
2. Population biology and ecology
2.1. Reproductive biology and ecology
2.2. Reproductive cycles
2.3. Feeding ecology
2.4. Growth
and longevity
2.5. Pelagic larval dynamics
2.6. Benthic population dynamics
3. Community ecology and coexistence
3.1. Ecosystem effects
3.2. Competitive interactions with other sea urchins
3.3. Competitive interactions with fish
3.4. Predation and predators
4. Herbivory
and grazing effects
4.1. Herbivory
4.2. Bioerosion
5. Conclusions
References
Chapter 12. Ecology of Loxechinus albus
J. Vasquez
1. Introduction
2. Morphology
3. Distribution
4. Habitat and substrate preferences
5. Food
6. Reproductive ecology
7. Population ecology
8. Community
ecology
9. El Ni?o
10. Conclusion
References
Chapter 13. Ecology of Paracentrotus lividus
C.-. Boudouresque, M. Verlacque
1. Introduction
2. Distribution and habitat
2.1. Habitat
2.2. Densities
2.3. Short- and long-term changes in density
2.4. Co-occurring species
3. Food
and feeding
3.1. Food preferences
3.2. Consumption rate
3.3. Ecological consequences of feeding
3.4. Competition with other herbivores
4. Movement and migration
5. Mortality
5.1. Predators
5.2. Diseases and parasites
5.3. Other causes of mortality
6. Growth
7. Reproduction
7.1. Reproductive cycles
7.2. Spawning
7.3. Recruitment
8. Conclusions
References
Chapter 14. Ecology of Psammechinus miliaris
M.
Kelly, A. Hughes, E. Cook
1. Appearance
2. Distribution
3. Habitat
4. Density
5. Population structure
6. Food and trophic ecology
7.
Growth rates, ageing, and energy partitioning
8. Reproduction
9. Conclusions
References
Chapter 15. Ecology of Echinometra
T. R. McClanahan,
N. Muthiga
1. Introduction
1.1. Species of Echinometra
1.2. Biogeography and large-scale distribution
1.3. Local distribution and abundance
patterns
2. Population biology and ecology
2.1. Reproductive biology and ecology
2.2. Feeding ecology
2.3. Energetics
2.4. Growth and
longevity
2.5. Pelagic larval dynamics
2.6. Benthic population dynamics
3. Community ecology and coexistence
3.1. Competitive interactions
with other herbivores
3.2. Competitive interactions with other sea urchins
3.3. Competitive interactions with fish
3.4. Predation and
predators
4. Herbivory and grazing effects
4.1. Herbivory
4.2. Erosion of calcium carbonate
5. Conclusion
References
Chapter 16. Ecology
of Evechinus chloroticus
M. Barker
1. Introduction
2. Geographic distribution
3. Habitat
4. Associated species
4.1. Kelp
4.2. Gastropods
5. Feeding
5.1. Diet
5.2. Feeding rate
6. Movement
7. Reproduction
7.1. Gametogenesis
7.2. Reproductive cycle
7.3. Reproductive output
7.4. Size at sexual maturity
7.5. Spawning
8. Larval development
9. Recruitment
10. Population biology
10.1. Growth
10.2. Mortality
10.3.
Population genetics
References
Chapter 17. Ecology of Heliocidaris erythrogramma
John K. Keesing
1. Introduction
2. Population genetics
and colour variability
3. Reproduction, development, settlement and recruitment
4. Growth and age
5. Movement and feeding
6. Influence
on benthic plants, occurrence of urchin barrens and feeding fronts
7. Predators, parasites, commensals and other ecological interactions
8. Conclusions
References
Chapter 18. Ecology of Strongylocentrotus droebachiensis
Robert Scheibling, B. Hatcher
1. Distribution and
abundance
1.1. Geographic range
1.2. Population density and spatial distribution
1.3. Physiological tolerance limits
2. Food and feeding
2.1. Food preferences and nutrition
2.2. Feeding behavior
2.3. Feeding rates
3. Growth
3.1. Determinants of growth
3.2. Growth rates
4. Reproduction
4.1. Reproductive timing
4.2. Gonad growth and gamete production
4.3. Fertilization rates
5. Settlement and recruitment
5.1. Larval development and settlement behaviour
5.2. Temporal and spatial patterns of settlement
5.3. Temporal and spatial patterns
of recruitment
5.4. Early post-settlement mortality
6. Predation
6.1. Predators
6.2. Behavioural responses to predators
6.3. Predation
as a mechanism of population regulation
7. Disease and parasitism
7.1. Microbial pathogens
7.2. Macroparasitic infections
8. Mortality
due to abiotic factors
9. Ecological role
References
Chapter 19. Ecology of Strongylocentrotus franciscanus and Strongylocentrotus purpuratus
Laura Rodgers-Bennett
1. Introduction
2. Sea urchin grazing and kelp forest ecosystems
3. Growth and survival
3.1. Growth
3.2. Survival
3.3. Aging
4. Reproduction
5. Fertilization
6. Larvae
6.1. Larval period
6.2. Blastulae and larval behavior
6.3. Larval cloning
7. Settlement
and recruitment
8. Population regulation
8.1. Competition
8.2. Predation
8.3. Disease
8.4. Physical factors and ocean warming
9. Genetics
10. Fisheries
10.1. West coast fisheries
10.2. Fishery experiments
10.3. Fishery enhancement
10.4. Gonad enhancement
11. Fishery management
12. Conservation
12.1. Metapopulation dynamics
12.2. Sea urchins as ecosystem engineers
12.3. Ecosystem management
References
Chapter
20. Ecology of Strongylocentrotus intermedius
Yukio Agatsuma
1. Introduction
2. Geographic distribution
3. Reproduction
3.1. Size at
maturity
3.2. Difference in reproductive cycle among localities
3.3. Fixation of reproductive cycle in each area
3.4. Spawning structure
4. Larval ecology
4.1. Occurrence
4.2. Distribution
4.3. Length of larval life and survival
5. Settlement and metamorphosis
6. Food and
feeding after settlement
6.1. Food habit
6.2. Food ingestion and absorption
6.3. Diurnal changes in food intake
6.4. Chemical stimulus
on feeding
7. Growth
7.1. Longevity and growth rings
7.2. Energy transformation to growth
7.3. Differences in growth among localities
7.4. Food and growth
7.5. Water temperature and growth
7.6. Gonadal growth
8. Habitat
8.1. Juvenile habitat
8.2. Habitat structure
9.
Community ecology
9.1. Bio-economy
9.2. Grazing effect on algal communities
10. Population dynamics
10.1. Fluctuation in larval occurrence
10.2. Juvenile recruitment
10.3. Fluctuation in gonadal growth
10.4 Effect of fisheries on population size
10.5 Predation
11. Conclusions
References
Chapter 21. Ecology of Strongylocentrotus nudus
Yukio Agatsuma
1. Introduction
2. Geographic distribution
3. Reproduction
3.1. Reproductive cycle
3.2. Spawning cues
4. Larval ecology
4.1. Occurrence
4.2. Growth and survival
5. Metamorphosis and settlement
5.1. Algal communities
5.2. Chemical inducer
6. Food and feeding after settlement
6.1. Food
6.2. Food ingestion and absorption
6.3. Food
selectivity
6.4. Chemical defense of algae
6.5. Abiotic factors on feeding
6.6. Foraging
7. Growth
7.1. Somatic growth
7.2. Gonadal growth
8. Habitat
8.1. Juvenile habitat
8.2. Movement
9. Community ecology
10. Population dynamics
10.1. Recruitment of juveniles
10.2. Annual
fluctuations
10.3. Decrease in population size
11. Conclusions
References
Chapter 22. Ecology of Hemicentrotus pulcherrimus, Pseudocentrotous
depressus, and Anthocidaris crassispina
Yukio Agatsuma
1. Introduction
2. Geographic distribution
3. Reproduction
3.1. Reproductive cycle
3.2. Abiotic factors and maturation
3.3. Spawning cue
4. Larval ecology
5. Settlement and metamorphosis
5.1. Induction with algae
5.2.
Chemical inducer
6. Food and feeding after settlement
6.1. Food
6.2. Feeding and food selectivity
7. Growth
7.1. Longevity and growth
rings
7.2. Somatic growth
7.3. Gonadal growth
8. Habitat
9. Community ecology
10. Population dynamics
11. Conclusions
References
Chapter
23. Ecology of Lytechinus
Stephen A. Watts, James B. McClintock, John M. Lawrence
1. The genus Lytechinus
2. Habitats
2.1. Lytechinus
variegatus
2.2. Lytechinus semituberculatus and Lytechinus pictus
3. Abundance
3.1. Lytechinus variegatus
3.2. Lytechinus semituberculatus
and Lytechinus pictus
4. Factors influencing distribution and abundance
4.1. Abiotic factors
4.2. Biotic factors
5. Food and feeding
6. Growth and survival
7. Reproduction
7.1. Lytechinus variegatus
7.2. Lytechinus pictus
8. Larval ecology and recruitment
9. Population
ecology
9.1. Predation
9.2. Competition
10. Community ecology
11. Conclusion
Acknowledgments
References
v
John M. Lawrence, Yukio Agatsuma
1. The genus Tripneustes
2. Distribution
3. Habitats
4. Behavior
5. Food
6. Growth
7. Reproduction
8. Recruitment
9. Mortality
10. Community
10.1. Effects of feeding
10.2. Competition
10.3. Predation
11. Conclusions
References
Chapter 25. Sea-urchin roe cuisine
John M. Lawrence
References
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