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Sandra Shumway, University of Connecticut, Department of Marine Sciences, Groton, Connecticut, U.S.A. Jay Parsons, Fisheries and Oceans Canada, Aquaculture Science Branch, Ottawa, Ontario, Canada
Description Scallops are among the better known shellfish and are widely distributed throughout the world. They are of great economic importance,
support both commercial fisheries and mariculture efforts and occupy a unique niche in the marine environment. Contributions from world
leaders in scallop research and culture cover all facets of scallop biology including anatomy, taxonomy, physiology, ecology, larval
biology and neurobiology. Chapters are also devoted to diseases and parasites, genetics, population dynamics and the adductor muscle,
with extensive reference lists provided for each chapter.
Since the publication of the first edition of Scallops: Biology,
Ecology and Aquaculture in 1991, commercial interest in scallops has grown globally and this is reflected in the seventeen extensive
chapters covering both fisheries and aquaculture for all species of scallops in all countries where they are fished or cultured. The
Second Edition is the only comprehensive treatise on the biology of scallops and is the definitive reference source for advanced undergraduate
and graduate students, mariculturists, managers and researchers. It is a valuable reference for anyone interested in staying abreast
of the latest advances in scallops.
Audience
Fisheries, marine biologists, researchers studying shellfish and the ocean environment
Contents
Table of Contents
Dedication v Preface vii Preface from first edition ix List of contributors xi
Chapter 1. New Phylogenies of the Pectinidae (Mollusca: Bivalvia): Reconciling Morphological and
Molecular Approaches Thomas R. Waller
1.1 Introduction
1 1.1.1 Molecular genetic studies 1 1.1.2 Methods
and materials 7 1.2 New observations
10 1.2.1 Subfamily Camptonectinae Habe, 1977 10 1.2.2 Subfamily
Palliolinae Korobkov in Eberzin, 1960 10 1.2.2.1 Tribe Pseudentoliini,
new tribe 12 1.2.2.2 Tribe Adamussiini Habe, 1977
13 1.2.2.3 Tribe Eburneopectinini, new tribe 14 1.2.2.4
Tribe Serripectinini, new tribe 15 1.2.2.5 Tribe Palliolini
Korobkov in Eberzin, 1960 17 1.2.2.6 Tribe Mesopeplini, new
tribe 20 1.2.3 Origins of the Decatopecten and Pecten Groups 22 1.2.4 Eastern Atlantic Pecten 28 1.3 Conclusions 30 Acknowledgments 32 References
32 Appendix 43
Chapter 2. Development, Physiology,
Behaviour and Ecology of Scallop Larvae Simon M. Cragg
2.1 Introduction
45 2.2 Scallop life history characteristics 45 2.3 Larval
development 55 2.3.1 Embryogenesis
55 2.3.2 Development of the larval shell 56 2.3.3 Organogenesis 59 2.3.3.1 Development of ciliation, the prototroch and the velum
59 2.3.3.2 Larval mantle 64 2.3.3.3 Musculature
64 2.3.3.4 Digestive tract 68 2.3.3.5 Sense organs 68 2.3.3.6 Apical organ 68 2.3.3.7 Statocysts
73 2.3.3.8 Eye spots 73 2.3.3.9 Mantle ciliation
74 2.3.3.10 Foot 77 2.3.3.11 Nervous system
78 2.3.3.12 Other organs 79 2.3.4 Metamorphosis
79 2.4 Comparative anatomy 81 2.5 Functional morphology 84 2.5.1 Locomotion 84 2.5.2 Feeding
85 2.5.3 Respiration 85 2.5.4 Sensory reception
86 2.6 Physiology and effects of environmental variables 87 2.6.1
Energy reserves and feeding 87 2.6.2 Respiration
88 2.6.3 Growth 89 2.6.4 Biochemical events during metamorphosis 90 2.7 Behaviour and larval distribution 90 2.7.1
Characteristics of locomotion during larval development 90 2.7.2
Responses to stimuli 91 2.7.3 Distribution of larvae
94 2.7.4 Development and the larval environment 98 2.7.5
Tolerance of environmental variables 99 2.8 Rearing methods
101 2.8.1 Spawning and manipulation of zygotes 101 2.8.2 Feeding
102 2.8.3 Control of disease in cultures 103 2.8.4 Collection of spat
from wild stocks 104 References
105
Chapter 3. Structure and Function in Scallops Peter G. Beninger and Marcel Le Pennec
3.1 Introduction
123 3.1.1 An overview of the scallop body 123 3.2 The mantle and its
derivatives 126 3.2.1 Gross functional anatomy
127 3.2.2 Microanatomy and functions 130 3.3 Pallial organs and particle
processing 132 3.3.1 Gills
132 3.3.1.1 Gill axis and arch 135 3.3.1.2 Principal filaments and
dorsal expansion 135 3.3.1.3 Ordinary filament
137 3.3.1.4 Haemolymph circulation in the gill 141 3.3.1.5 Particle
processing on the gill 141 3.3.1.6 Particle selection at the gill
141 3.3.1.7 Particle retention lower size limit 143 3.3.1.8 Ingestion
volume regulation on the gill 143 3.3.2 Labial palps and lips
144 3.3.2.1 Labial palps 144 3.3.2.2 Particle processing on the labial
palps 144 3.3.2.3 Lips
147 3.4 Digestive system and digestion 150 3.4.1 Mouth and oesophagus
150 3.4.2 Stomach, crystalline style and gastric shield 150 3.4.3 The
digestive gland 154 3.4.4 Intestine, rectum and anus
157 3.4.5 Digestive sites and postingestive selection 159 3.5 Cardio-Vascular
system 159 3.5.1 General circulation
160 3.5.1.1 The arterial system 160 3.5.1.1.1 Anterior aorta
160 3.5.1.1.2 Posterior aorta 160 3.5.1.2 The venous system
162 3.5.2 The heart 162 3.5.2.1 The ventricle
162 3.5.2.2 The auricles and their excretory structures 163 3.5.2.3
Structure and ultrastructure of heart cells 163 3.5.2.4 Contraction
165 3.5.2.4.1 Refilling 165 3.5.2.4.2 Coordination of alternate AV
beat 166 3.5.2.4.3 Pacemaker mechanism
166 3.5.2.4.4 Regulation of pacemaker 167 3.5.3 Haemolymph
167 3.5.3.1 Plasma 168 3.5.3.2 Haemocytes
168 3.5.3.2.1 Haemocyte types 169 3.5.3.2.2 Functions of haemocytes
169 3.6 Excretory system 170 3.6.1 Pericardial (auricular) glands
170 3.6.2 Kidney 172 3.6.3 Functions of the kidney and pericardial
glands 173 3.7 Reproductive system
173 3.7.1 Sexuality: Gonochory, hermaphroditism and their variants
173 3.7.2 Origin and formation of the gonad 176 3.7.3 Anatomy, histology
and ultrastructure of the adult gonad 177 3.7.3.1 Outer epithelium
177 3.7.3.2 Perigonadal connective tissue 179 3.7.3.3 Inter-acinal
connective tissue 179 3.7.3.4 Haemolymph sinuses
179 3.7.3.5 Acini 180 3.7.3.6 Evacuating ducts
180 3.7.4 Gametogenesis 180 3.7.4.1 Oogenesis
180 3.7.4.1.1 Premeiotic stage 180 3.7.4.1.2 Previtellogenic stage
181 3.7.4.1.3 Vitellogenesis and metabolite transport to the oocyte 181 3.7.5
Oocyte atresia 185 3.7.6 Spermatogenesis, spermatozoon ultrastructure
and taxonomy 188 3.7.7 Fertilisation
190 3.8 Nervous and sensory systems 191 3.8.1 General organisation
of the nervous system and functional anatomy of principal ganglia 191 3.8.1.1
Cerebral and pedal ganglia 191 3.8.1.2 The parietovisceral ganglion and
its nerves 191 3.8.1.3 Histology and neurosecretions of the ganglia
196 3.8.1.4 The circumpallial nerve 198 3.8.2 Sensory structures
198 3.8.2.1 Visual system 198 3.8.2.2 Epithelial sensory cells and
tentacles 198 3.8.2.3 Abdominal sense organ
199 3.8.2.4 Osphradia 201 3.8.2.5 Statoreceptors
204 3.8.3 Neurotransmitters and neurohormones 204 3.9 Foot-byssal complex
205 3.9.1 External morphology and development of the foot-byssal complex
207 3.9.2 Anatomy and histology of the foot-byssal gland complex 208 3.9.2.1
The protein gland 208 3.9.2.2 The enzyme gland
208 3.9.2.3 The byssus 210 3.9.3 Functioning of the foot-byssal complex
210 Acknowledgments 210 References
211
Chapter 4. Scallop Adductor Muscles: Structure and Function Peter D. Chantler
4.1 Introduction
229 4.2 Structure of the striated adductor muscle 231 4.2.1 Fibre microanatomy 231 4.2.2 Actin and thin filament structure 233 4.2.3 Thick filament
structure 240 4.3 Structure of the smooth adductor muscle
243 4.3.1 Fibre microanatomy 243 4.3.2 Thin filament structure
245 4.3.3 The structure of paramyosin-rich thick filaments 247 4.4
Myosin 253 4.5 Function of the striated adductor
271 4.5.1 Mechanics 271 4.5.2 The interaction of myosin with actin
272 4.5.3 The crossbridge cycle 277 4.5.4 Myosin-linked regulation
282 4.6 Function of the smooth adductor 289 4.6.1 Physiology
289 4.6.2 Catch mechanism 292 4.7 Achievements and goals
296 Acknowledgments 298 References
298
Chapter 5. Neurobiology and Behaviour of the Scallop Lon A. Wilkens
5.1 Introduction
317 5.2 The visual system 317 5.2.1 Functional anatomy of the eyes
317 5.2.2 Retinal physiology 322 5.2.3 Receptor potential biophysics
324 5.2.4 Anatomy and physiology of vision in the central nervous system
328 5.2.5 Visual behaviours 332 5.3 The locomotory system
335 5.3.1 Escape responses and swimming 335 5.3.2 Sensory and motor
functions of the mantle 341 5.3.3 Innervation and neuromuscular physiology
of the adductor muscle 345 5.3.4 Coordination of locomotory behaviour
by the central nervous system 348 Acknowledgments
352 References 352
Chapter 6. Reproductive Physiology Bruce
J. Barber and Norman J. Blake
6.1 Introduction 357 6.2 Gametogenic
cycles 358 6.2.1 Definition
358 6.2.2 Means of assessment 358 6.2.2.1 Visual observation
360 6.2.2.2 Gonad mass and index 360 6.2.2.3 Histology
367 6.2.2.4 Abundance of larvae and spat 369 6.2.3 Variations in gametogenic
cycles 370 6.2.3.1 Intra-specific variations
370 6.2.3.2 Inter-specific variations 375 6.3 Regulation of gametogenic
cycles 375 6.3.1 Gametogenesis and fecundity
376 6.3.1.1 Exogenous regulation 376 6.3.1.2 Endogenous regulation
381 6.3.2 Spawning and spawning synchrony 385 6.3.2.1 Exogenous regulation 385 6.3.2.2 Endogenous regulation 389 6.4 Energy metabolism
390 6.4.1 Tissue weights and indices 391 6.4.2 Tissue biochemical composition 392 6.4.3 Physiological indices 395 6.4.4 Radiotracer experiments
398 6.4.5 Ultrastructure and histochemistry 398 6.4.6 Mechanisms
400 6.5 Applications to aquaculture 401 6.5.1 Broodstock conditioning
401 6.5.2 Spawning 403 6.5.3 Larval growth and survival
403 6.6 Summary 404 6.6.1 Gametogenic cycles
404 6.6.2 Regulation of gametogenesis 404 6.6.3 Energy metabolism
405 6.6.4 Applications to aquaculture 406 References
406
Chapter 7. Physiology: Energy Acquisition and Utilisation Bruce A. MacDonald, V. Monica Bricelj and Sandra E. Shumway
7.1 Introduction 417 7.2 Energy acquisition
417 7.2.1 Food sources 417 7.2.2 Feeding currents and mechanisms of
particle capture 420 7.2.3 Particle retention efficiency
421 7.2.4 Feeding rates 424 7.2.5 Clearance rate in relation to food
concentration 427 7.2.6 Influence of temperature on feeding rates
430 7.2.7 Pseudofeces production, pre- and post-ingestive particle selection
432 7.2.8 Absorption efficiency 436 7.2.9 Effects of suspended sediments
on feeding and growth 439 7.2.10 Effects of flow on feeding and growth
445 7.2.11 Effects of harmful and toxic algae 447 7.3 Energy utilisation:
Metabolic expenditure 453 7.3.1 Metabolic rate and oxygen availability
455 7.3.2 Metabolic cost of reproduction 458 7.3.3 Metabolic rate in
relation to temperature and latitude 460 7.3.4 Metabolic rate in relation
to activity levels 461 7.3.5 Anaerobic metabolism
463 7.4 Energy utilisation 467 7.4.1 Excretion and byssus secretion
467 7.4.2 Growth 468 References
474
Chapter 8. Physiological Integrations and Energy Partitioning Raymond J. Thompson and Bruce A. MacDonald
8.1
Introduction 493 8.2 Energy balance, physiological integrations and the
partitioning of energy between growth and reproduction 493 8.2.1 Energy
budgets 493 8.2.2 Scope for growth
494 8.2.3 Growth efficiency and turnover ratio 494 8.2.4 Growth of
shell and somatic tissue 495 8.2.5 Reproductive effort
496 8.3 Changes related to age of individuals 498 8.3.1 Scope for growth
498 8.3.2 Growth efficiency and turnover ratio 498 8.3.3 Somatic growth
and maximum size 499 8.3.4 Somatic production and reproductive output
500 8.3.5 Reproductive effort 502 8.4 Environmental influence on production 504 8.4.1 Scope for growth and growth efficiency 504 8.4.2 Growth
of shell and somatic tissue 506 8.4.3 Reproductive effort
509 8.5 Reproductive value and cost 511 8.5.1 Residual reproductive
value 511 8.5.2 Reproductive cost
512 8.6 Population production 514 Acknowledgments
516 References 516
Chapter 9. Nutrition in Pectinids Ana
Farias and Iker Uriarte
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