Marine Mussels

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Elizabeth Gosling. Marine Mussels

Table of Contents

List of Tables

List of Illustrations

Guide

Pages

Marine Mussels. Ecology, Physiology, Genetics and Culture

Preface

Notes

Acknowledgements

1 Phylogeny and Evolution of Marine Mussels. Introduction

Phylogeny of the Phylum Mollusca

Phylogeny and Evolution of Bivalvia

Evolution and Adaptive Radiation in Bivalvia

Phylogeny and Evolution in the Mytilida

Significant Evolutionary Developments in the Family Mytilidae

Notes

References

2 Functional Morphology. Introduction

Shell

Structure

External Characteristics

Mantle. Structure

Function

Gills

Structure

Functions

Foot

Byssus Composition

Attachment

Labial Palps

Alimentary Canal. Stomach and Digestive Gland

Gonads

Heart and Haemolymph Vessels

Excretory Organs

Nerves and Sensory Receptors

References

3 Ecology of Marine Mussels. Introduction

Global and Local Distribution Patterns

Global Distribution

Local Distribution

Factors Affecting Geographic Distribution

Temperature, Salinity and Hydrographic Factors

Factors Affecting Local Distribution

Physical Factors. Temperature

Salinity

Wave Exposure

Substrate

Disturbance

Biological Factors

Predators, Biofouling and Competitors

Climate Change and Potential and Observed Impacts on Marine Mussels

Climate Warming

Ocean Acidification

Hypoxia

Notes

References

4 Particle Processing and Nutrient Utilisation. Introduction

Filtration and Clearance Rates

Estimation of Filtration Rate

Factors Influencing Filtration Rate. Size

Particle Concentration

Temperature

Salinity

Current Velocity

Control of Filtration Rate

Energetic Costs of Filtration

Particle Processing on the Gills and Labial Palps and in the Stomach

Selective Particle Capture on the Gills

Pre‐ingestive Particle Processing on the Gills and Labial Palps

Role of Mucus

Labial Palps and Pseudofaeces Production and Transport

Particle Selection Efficiency

Postingestive Particle Processing in the Stomach

Alimentary Canal and Digestive Process

The Stomach and Extracellular Digestion

The Digestive Gland and Intracellular Digestion

The Intestine

Mussel Nutrition

Gut Content Analysis

Stable Isotope Analysis

Fatty Acid Profile Analysis

Additional Food Sources: Picoplankton, Bacteria and DOM

Absorption Efficiency

Impacts of Mussels on Marine Ecosystems

Bivalve Feeding and the Plankton Community

Biodeposition and Mussels

Notes

References

5 Reproduction, Larval Development, Dispersal and Recruitment. Introduction

Sex Identification

Doubly Uniparental Inheritance and Sex Determination

Gametogenesis

Reproductive Cycles. Methods of Assessment

Annual Cycles

Mytilus Species

Perna Species

Geukensia Species

Modiolus Species

Mytella Species

Bathymodiolus Species

Factors Controlling Reproduction

Exogenous Regulation of Gametogenesis and Spawning

Endogenous Regulation of Gametogenesis

Annual Storage Cycle

Reproductive Effort and Fecundity

Fertilisation and Gamete Compatibility

Larval Development

Factors Affecting Larval Growth

Temperature and Salinity

Food Ration

Larval Dispersal and Population Connectivity

Larval Identification and Abundance

Settlement, Metamorphosis and Post‐Larval Dispersal. Settlement

Metamorphosis and Post‐Larval Dispersal

Recruitment

Notes

References

6 Growth. Introduction

Measurement of Absolute Growth

Length‐Frequency Distributions

Growth Curves

Mark‐and‐Recapture and Growth Increment Analysis

Shell Growth Rings

The Mussel Shell as a Marine Archive

Measurement of Allometric Growth

Scope for Growth and Dynamic Energy Budget Models. Scope for Growth

Calculation of SFG

Dynamic Energy Budget

Growth Modulators

Exogenous Growth Modulators. Food

Temperature

Salinity

Tidal Level and Wave Action

Stock Density

Carbon dioxide and Oxygen Concentrations

Other Factors

Endogenous Growth Modulators. Genotype

Size

Growth Factors

Notes

References

7 Physiology of the Circulatory, Respiratory and Excretory Systems. Cardiovascular System. Heart and Haemolymph

Haemolymph Plasma and Haemocytes

Heart Rate

Factors Affecting HR

Respiratory System

Factors Affecting Oxygen Consumption. Size

Food

Temperature

Acute Temperature Response

Acclimation to Temperature Change

Adaptation and Acclimation to Low Temperatures

Temperature Tolerances

Salinity

Oxygen Availability

Aerial Exposure

Other Factors

Excretory System. Excretion

Factors Influencing Excretion Rates

Oxygen–Nitrogen (O:N) Ratio

Osmoregulation

Notes

References

8 Mussels and Marine Environmental Contaminants. Introduction

Contaminants in the Marine Environment

Heavy Metals

Persistent Organic Contaminants

Polycyclic Aromatic Hydrocarbons

Surfactants

Polychlorinated Biphenyls

Pesticides

Dioxins

Emerging Organic Contaminants

Brominated Flame Retardants

Endocrine‐Disrupting Compounds

Pharmaceuticals and personal care products

Microplastic Particles

Nanoparticles

Contaminant Levels in Seawater, Sediments and Biota. Bioaccumulation

Absorption and Absorption Efficiency

Assimilation and Assimilation Efficiency

Bioconcentration Factor

Kinetic Modeling in Contaminant Bioaccumulation

Factors Affecting Bioconcentration in Marine Mussels

Salinity

Temperature

Age and Body Size

Reproductive Cycle and Sex

Location within the Intertidal Zone

Time of Year of Sample Collection

Biomonitoring

Bioindicators

Mussel Monitoring Programmes. The Mussel Watch Program

Mussel Watch Monitoring on the Californian Coast

Mussel Watch Monitoring in Coastal Waters of the Mediterranean Sea

Asia‐Pacific Mussel Watch

Biological Markers of Pollution

Biomarkers in Perna Viridis and Other Marine Mussel Species

Enzyme Reactions

Specific Proteins

Acetylcholinesterase Inhibition

Cellular Stress Biomarkers: Lysosome Membrane Stability and Permeability

Peroxisome Proliferation

Morphological Damage

Physiological Markers

Biomarkers of Genotoxicity

An Integrative Biomarker Approach

Notes

References

9 Population Genetics, Genomics and Selective Breeding. Introduction

Population Genetics

Molecular Markers and Applications

Genetic Variation and Population Structure in Mussels

Intraspecific Genetic Variation. Mytilus spp

Perna spp

Interspecific Genetic Variation. The Mytilus Edulis Species Group

Perna spp

Brachidontes and Bathymodiolus spp

Hybrid Zones

M. Edulis and M. Galloprovincialis Hybrid Zones

M. Edulis and M. Trossulus Hybrid Zones

M. Trossulus and M. Galloprovincialis Hybrid Zones

Invasive Species: Origins, Dispersal and Population Genetics

Mussel Genomics

Genome Mapping

Functional Genomics. Transcriptomics

Proteomics

Metabolomics

Quantitative Genetics

Selective Breeding

Breeding Objectives

Broodstock Establishment

Breeding Schemes

QTL Mapping and Marker‐Assisted Selection

Mussel Breeding Programmes

Notes

References

10 Mussel Culture. Introduction

Wild Seed Collection

Hatchery Culture of Seed and Juvenile Mussels

Culture of Microalgae

Replacement/Supplement Diets for Microalgae

Hatchery Culture of Mussels. Broodstock Conditioning

Spawning, Fertilisation and Cryopreservation

Embryo Development

Larval Rearing

Metamorphosis, Settlement and Nursery culture of Spat

Disease in the Hatchery

Grow‐out of Mussel Seed

Mussel Culture

Culture Methods. Longline Culture in Chile

Raft Culture in Spain

Bouchot Culture in France

Bottom Culture in The Netherlands

Pole/Stake Culture in Thailand

Offshore Mussel Culture

Mussel Culture and the Environment

An Ecosystem Approach to Bivalve Culture

Notes

References

11 Diseases, Parasites and the Immune Response. Introduction

Viruses

Bacteria

Fungi

Protistans

Apicomplexa

Ascetosporea

Haplosporida

Paramyxida

Chlorophyta

Ciliophora

Porifera

Platyhelminths

Trematodes

Cestodes

Turbellaria

Annelids

Crustaceans

Neoplasia

Defence Mechanisms

Genomics of Bivalve Immunity

Cellular Defence Components of the Immune System

Immune Recognition

Humoral Defence Mechanisms

Lysozymes and Proteases

Membrane Attack and Perforin Proteins

Antimicrobial Peptides

Cytokines

Oxidative Burst

Signal Transduction Pathways

Abiotic Effects on Mussel Immune Response

Notes

References

12 Mussels and Public Health. Introduction

Bacterial Infections

Salmonella

Vibrio

Escherichia Coli

Bacterial Assays

Viral Infections

Viral Assays

Parasites

Biotoxins

Harmful Algal Blooms

Detection of Marine Toxins

Functional Assays

Immunological Assays

Analytical Methods

Industrial Pollutants

Controls on the Production and Processing of Mussels and Other Bivalves. Regulatory Framework

Controlling Harvesting Areas

Bivalve Purification Procedures

Relaying

Depuration

Efficiency of Depuration

Monitoring and Quality Control

Monitoring of Live Mussels in Canada: Legislation and Regulation

Growing Areas: Sanitary Survey and Classification

Marine Biotoxin Monitoring and Control

Hazard Analysis and Critical Control Point and its Application

Notes

References

Species Index

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Subject Index

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Elizabeth Gosling

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Due to their dominant role in ingestion and respiration, the gills are among the main target organs in the bioaccumulation of pesticides, soluble heavy metals and hydrocarbons. Complex mixtures of heavy metals and polycyclic aromatic hydrocarbons (PAHs) cause morphological changes in the gill epithelium of Mytella falcata, leading to an increase in the number of gill mucous cells, haemocytes and cell turnover processes. These are possible mechanisms to compensate for cell injury and prevent entry of pollutants from gill filaments into the entire organism (David & Fontanetti 2005; David et al. 2008). Exposure to mercury over a 24‐day period caused an initial deterioration in neural and epithelial cells, increased interstitial cell oedema and reduced ciliation in Perna perna. However, after a metal‐free recovery period, gill filament morphology returned to near normal (Gregory et al. 2002). This is not unexpected, as metallothioneins (MTs) – metal‐binding, heat‐stable, low‐molecular‐weight proteins – play an important role in detoxifying trace metals in bivalves and are widely distributed in gill and digestive gland tissue. Consequently, mussel MT levels are increasingly being used as a biomarker of heavy metal contamination in coastal ecosystems (Khati et al. 2012; see Chapter 8).

The foot first appears when bivalve larvae are about 200 μm in length, and becomes functional in crawling and attachment at ~260 μm shell length. This is the pediveliger stage of development, which immediately precedes settlement and metamorphosis (see Figure 5.10). The ciliated foot is proportionately very large and sock shaped, and is made up of layers of circular and longitudinal muscles surrounding a capacious haemolymph space. A byssal duct opens at the ‘heel’ of the foot, and a byssal pedal groove extends forward along the ‘sole’ from this opening. The groove is embedded in secretory tissue, which produces the different byssal thread components (see later). While swimming, the foot is fully extended, and periodically the velum (larval swimming organ) is withdrawn and the larva sinks to the bottom and begins to crawl. If the substrate is unsuitable (i.e. does not stimulate the secretion of byssus), the foot is withdrawn and the larva once again swims off (Lutz & Kennish 1992). This cycle can be repeated many times over a period of a few days. In Mytilus, when a suitable substrate is found, the larva continues to crawl for some time, gradually ceases movement, protrudes the foot and quickly secretes a single byssal thread. In the newly attached mussel larva, this thread can be repeatedly broken and reformed before final settlement takes place. As the mussel grows in length, more and more attachment threads are secreted; this is not surprising, as larger individuals are subject to greater mechanical stress than smaller ones. To resist dislodgement, mussels cluster their threads in the direction of applied forces (e.g. facing ebb and flow of tide). The adhesive in mussel larvae differs from that of adults, resembling the mucous secreted by other benthic marine species at the larval stage (Petrone et al. 2008). The green crenella, Musculus discors, is unusual in that the byssus threads that are used to fix it to the substrate are woven into a nest or cage surrounding the shell, similar to a ball of twine. Eggs in mucous strings are retained within this nest, which may incorporate a variety of macroalgae (Merrill & Turner 1963).

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