Life in the Open Ocean
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Joseph J. Torres. Life in the Open Ocean
Table of Contents
List of Tables
List of Illustrations
Guide
Pages
Life in the Open Ocean. The Biology of Pelagic Species
Preface
Acknowledgments
1 Physics and the Physical Environment
The Vastness of the Open Ocean
The Properties of Water
Density
Viscosity
Reynolds Number
Drag
Temperature
The Oceans and Ocean Basins
Ocean Circulation
Surface Currents: Ocean Gyres and Geostrophic Flow
Coriolis Force
Ekman Transport
Ocean Gyres and Geostrophic Flow
Upwelling
Deep‐Ocean Circulation
Water Masses
Oxygen
Pressure
Sound
Light
Absorption and Scattering
Traditional Depth Zones in the Ocean
Concluding Thoughts
References
2 Physiological Accommodation to Environmental Challenges
Temperature
Terms
Temperature Effects on Survival: The Tolerance Polygon
Temperature Effects on Rate Processes – The Q10 Approximation
Patterns of Thermal Acclimation
Climatic Adaptation in Ectotherms
Temperature Compensation via Changes in Enzyme Concentration: The Quantitative Strategy for Short‐term Change
Compensation via Changes in Enzyme Quality – Isozymes, Allozymes, and Temperature Adaptation
What Properties of Enzymes Can Be Changed?
Lipids and Temperature
A Membrane Primer
Pressure
Early Work
Later Work
Whole Animal Work
Molecular Mechanisms of Adaptation to Pressure
Pressure and Membranes
Oxygen
Box 2.1 An Aside on Units
Severity of Oxygen Minima, “Dead Zones,” and the Intertidal
Adaptations to Oxygen Minima. The Aerobic Strategy
Salinity
Depth
Concluding Thoughts
References
3 The Cnidaria. Introduction
Classification. History
Classification Schemes
Phylum Cnidaria. Subphylum Medusozoa
Subphylum Anthozoa
Subphylum Myxozoa
The Hydromedusae. Morphology Basics
Morphological Detail and Life Histories
The Scyphomedusae. Basic Characteristics
Morphological Detail and Life Histories. General
Coronatae
Semaeostomae
Rhizostomae
The Cubomedusae
Foraging Strategies
General Considerations
The Cnidae
Venoms
Interaction with Prey
Direct Interception
Encounter Zone
Anthomedusae
Leptomedusae
Trachymedusae
Narcomedusae
Scyphomedusae
The Model
Swimming and Hunting Behavior
Water Flow and Swimming
Attraction Between Predator and Prey
Diets, Feeding Rates, and Impacts on Prey Populations
Rogue Hydroids: Predatory Polyps in the Midwater
Feeding in the Cubomedusae
Locomotion
The Mesoglea
Nerve Nets and Nervous Control of Swimming
Senses and Sensory Mechanisms
The Siphonophores
Terminology and Affinities of Siphonophore “Persons”
Whole Animal Organization
Life Histories
The Siphonophore Conundrum
Feeding. Fishing Behavior
Digestion
Diets and Selectivity
Ecological Importance
Locomotion
Buoyancy
Vertical Distribution
Diurnal Vertical Migration
Geographical Distribution
Organization and Sensory Mechanisms
Epithelial Conduction vs. Neural Conduction
The Cnidaria Formerly Known as Chondrophora. Classification
Characteristics of the Chondrophoran Medusa
Evolution Within the Chondrophora
Feeding in the Chondrophora
Locomotion
References
4 The Ctenophora. Introduction
Classification. History
Classification Schemes
Ctenophore Basics
Morphology. Cydippida
Lobata
Cestida
Beroida
Platyctenida
Ganeshida
Thalassocalycida
Foraging Strategies. General Considerations
Interaction with Prey. The Cydippids
The Lobates
The Cestids
The Beroids
The Platyctenids
The Ganeshids
The Thalassocalycids
Specialists
Diet, Feeding Rates, and Impacts on Prey Populations
Ctenophores as Invasive Species
Digestion
Nerves and Sense Organs: Coordination and Conduction
Locomotion
Distribution
Ctenophores and Evolution
References
5 The Nemertea. Introduction
Classification. History
Of Germ Layers and Body Cavities
Hydrostatic Skeletons
Classification
Morphology
Proboscis Apparatus
The Pelagic Body Form and Locomotion
Color
Nervous System
Sense Organs
Circulatory System
Excretory System
Digestive System
Reproduction
Development
Foraging Strategies
Vertical and Geographic Distributions
References
6 The Annelida. Introduction
History
Classification
Box 6.1 Useful Terms
Phylum Annelida
Class Polychaeta
Subclass Echiura
Subclass Errantia
Class Clitellata
Subclass Oligochaeta
Subclass Hirudinea
The Pelagic Polychaetes
Polychaete Subclass Errantia
Order Phyllodocida
Polychaete Subclass Sedentaria
Order Terebellida
Morphology. General
External Anatomy
The Head Region
The Trunk or Metastomial Region
Internal Anatomy
Excretory System
Pelagic Species
The Nervous System
Sense Organs
Circulatory System
Pelagic Species
Gas Exchange
Pelagic Species
Digestive System
Reproduction
Epitoky
Synchronicity
The Pelagic Species. Tomopteris
Alciopini and Lopadorrynchidae
Development
Locomotion
Foraging Strategies
The Hunters
Diets
The Suspension Feeders
Poeobiidae, Chaetopteridae
Distributions. Geographical
Vertical
Bioluminescence
References
7 The Crustacea. Introduction
Arthropod Classification. History
Subphylum Crustacea
Subphylum Hexapoda
Subphylum Myriapoda
Subphylum Chelicerata
Panarthropoda Phyla. Phylum Onychophora
Phylum Tardigrada
Synopsis of Universal Arthropod Characteristics
The Crustacea
Characteristics
Classification
Subphylum Crustacea. Class Remipedia
Class Cephalocarida
Class Branchiopoda
Class Copepoda
Class Thecostraca
Class Tantulocarida
Class Mystacocarida
Class Branchiura
Class Pentastomida
Class Ostracoda
Class Malacostraca
Subclass Phyllocarida
Subclass Hoplocarida
Subclass Eumalacostraca
Superorder Eucarida
Order Euphausiacea
Order Amphionidacea
Order Decapoda
Superorder Peracarida
Order Mysida
Order Lophogastrida
Order Isopoda
Order Amphipoda
Order Cumacea
Order Tanaidacea
Order Mictacea
Order Spelaeogriphacea
Order Stygiomysida
Order Thermosbaenacea
Superorder Syncarida
Crustacean Systems. Integument and Molting. Integument
Molting
Molt Cycle Stages
Hormonal Control
Joints and Appendages. Joints
Appendages
Excretory System
Extra‐renal Mechanisms
How the System Works
Nitrogen Excretion
The Nervous System and Sensory Mechanisms. The Central Nervous System
Sensory Modalities
Photoreception
Mechanoreception
Chemoreception
Circulatory and Respiratory Systems
Circulation and Oxygen Transport in the Blood: Hemocyanin
Digestive System
Basic Development
The Micronektonic Crustacea
The Pelagic Eucarida. Order Euphausiacea. Introduction
History
Classification
Euphausiid Anatomy
Nervous System
Sensory Mechanisms
Bioluminescence and Photophores
Circulatory, Respiratory, and Excretory Systems
Digestive System
Development
Gonad Development and Spawning
Food and Feeding
Geographical Distribution
Vertical Distribution and Vertical Migration
Ecological Factors
Acoustics, Species Composition, Light, and Layer Movement
Adaptive Significance: The Environment
Adaptive Significance of Vertical Migration
Order Decapoda. Introduction
History
Decapod Anatomy
Nervous System
Sensory Mechanisms
Bioluminescence and Photophores
Functions of Bioluminescence
Circulatory, Respiratory, and Excretory Systems
Digestive System
Development
Reproduction
Dendrobranchs
Carideans
Vertical Distribution
Food and Feeding
Geographic Distribution
Infraorder Anomura; Superfamily Galatheoidea; Family Munididae; Genera Pleuroncodes, Munida, and Cervimunida. The Pelagic Squat Lobsters
Order Amphionidacea
Introduction and History
Development and Natural History
Superorder Peracarida
Orders Lophogastrida and Mysida. Introduction
History
Anatomy
Nervous System
Sensory Mechanisms
Bioluminescence
Circulatory, Respiratory, and Excretory Systems
Digestive System
Reproduction and Development
Life History of a Lophogastrid
Food and Feeding
Locomotion
Vertical Distributions
Geographic Distributions
Order Amphipoda. Introduction
History
Classification
Anatomy
Nervous System
Sensory Mechanisms
Bioluminescence
Circulatory, Respiratory, and Excretory Systems
Digestive System
Reproduction and Development
Food and Feeding
Amphipods and Gelatinous Zooplankton
Geographic Distributions
Vertical Distributions
Cameo Players
References
8 The Mollusca. Introduction
Classification. History
The Pelagic Molluscs
Phylum Mollusca. Class Caudofoveata
Class Solenogastres
Class Monoplacophora
Class Polyplacophora
Class Scaphopoda
Class Bivalvia
Class Gastropoda
Class Cephalopoda
Body Organization
The Gastropoda. Classification
Class Gastropoda. Classification Below Subclass Given for Pelagic Species Only
Gastropod Systems and Structures
The Digestive Tract
Circulation
Respiration
Excretion
Shell Formation
The Nervous System
Sensory Mechanisms
The Pelagic Gastropods: Anatomy and Habits. The Janthinid Snails
Food and Feeding
Reproduction and Development
The Heteropods
Locomotion
Food and Feeding
Feeding Chronology
Vertical Distributions and Migrations
Abundance and Geographic Distribution
Reproduction and Development
The Pteropods
Order Pteropoda
Euthecosomata. Anatomy
Locomotion
Food and Feeding
Reproduction and Development
Pteropods, pH, and the Carbonate Cycle
Gymnosomata. Anatomy and Feeding
Locomotion
Reproduction
Superfamily Hydromyloidea
Pteropod Distributions
Vertical Migrations
The Nudibranchs
Family Glaucidae
Family Fionidae
Family Phylliroidae
The Cephalopoda
Classification
Basic Anatomy of the Major Cephalopod Groups
General
The Nautilida
The Sepiida and Spirulida
The Myopsida and Oegopsida
The Octopodiformes
Cephalopod Systems. Feeding and Digestion
The Digestive Tract
Movement of Food
Circulation
Gas Exchange
Excretion
Nervous System and Sensory Mechanisms
The Brain
The Giant Fiber System
Sensory Mechanisms
Touch, Taste, and Smell
Mechanoreception: Equilibrium and Acceleration
Hearing
Vision
Color
Bioluminescence
Locomotion and Buoyancy
Buoyancy
Life Histories
Nautiloids
Coleoids
Reproduction and Development
Vertical Distribution and Migration
Geographic Distribution
References
9 The Chordata. Introduction
Deuterostomes and the Phylogenetic Toolkit
Classification
Subphylum Tunicata. Class Ascidiacea (2935)
Class Appendicularia (68)
Class Thaliacea (78)
Basic Anatomy and Life History. The Ascidians
The Pyrosomes
The Salps
The Doliolids
The Appendicularia
The Appendicularian House
Tunicate Systems. Locomotion and Buoyancy
Pyrosomes
Salps
Doliolids
Appendicularia
Nervous Systems and Sensory Mechanisms
Pyrosomes
Doliolids
Salps
Appendicularia
Gas Exchange, Circulation, and Excretion
Trophic Role
Bioluminescence
Predators, Parasites and Other Interactions
Geographic and Vertical Distributions
References
10 The Fishes. Introduction
The Deep‐Sea Groups
A Brief History of Fishes. The Jawless Fishes
The Jawed Fishes. Teleosts
Elasmobranchs
Holocephali
The Classes of Living Fishes. Class Myxini
Class Petromyzonti
Class Elasmobranchii
Class Holocephali
Class Coelacanthi
Class Dipneusti
Class Cladistii
Class Actinopterygii
Subclass Chondrostei
Subclass Holostei
Subclass Teleostei
Cohort Elopomorpha
Cohort Otocephala
Cohort Euteleostei
Superorder Osmeromorpha
Superorder Ateleopodimorpha
Superorder Cyclosquamata
Superorder Scopelomorpha
Superorder Lamprimorpha
Superorder Paracanthopterygii
Superorder Acanthopterygii
Fish Systems. Basic Anatomy. External Features and Terms
Skull and Skeleton
Feeding and Digestion. Food Acquisition, the Three Dominant Modes: Ramming, Sucking, and Biting. Ram Feeding
Suction Feeding
Biting
Food Sorting: The “Pharyngeal Jaws”
Digestion: The Alimentary Canal
Circulation, Respiration, and Excretion. Circulation
Elasmobranch and Teleost Hearts
Arteries, Veins, and Continuity
Gas‐Exchange in the Teleosts and Elasmobranchs
Unidirectional Flow and Countercurrent Exchange: Maximizing the Concentration Gradient
Blood and Oxygen at the Respiratory Surface. Blood Basics
Red Cell Modifications
CO2 Transport–Far Different from O2
The Bohr and Haldane Effects
Secretion of Gases into the Swimbladder of Fishes
Nitrogen Excretion
Ammonia Excretion
Urea
Osmotic and Ionic Regulation. Introduction
Osmosis and Diffusion
The gills
Esophagus
Stomach
Intestine
Locomotion
Musculature
Red and White Muscle
Drag and Swimming Costs
Swimming Costs
Maximum Swim Speeds
Endothermy
Warm‐Brained Billfishes
Swimming in Mesopelagic Fishes
Buoyancy
The Nervous System. Anatomy and Basics
The Brain
Cranial Nerves
The Spinal Cord
Sensory Mechanisms. Sensory Modalities
Photoreception
The Choroid Body
Photoreceptors: Rods and Cones
Light and Vision
Visual and Morphological Adaptations to Deep‐Sea Life
Tubular Eyes and Retinal Adaptations
Mechanoreception
Free‐Standing Neuromasts and the Lateral Line
The Inner Ear and Sound Reception. Structure
Balance, Acceleration, and Angular Momentum
Hearing
Electroreception
Chemoreception: Olfaction and Gustation
Camouflage, Bioluminescence, Photophores. Camouflage
Bioluminescence and Photophores
References
11 Communities. Introduction
The Gulf of Mexico
The Northern California Current
The Antarctic
System Comparisons
The Decapods and Mysids
The Euphausiids
The Myctophids
Non‐myctophid Fishes
The Cephalopods
Gelatinous Zooplankton and Amphipods
Concluding Observations
Physical and Biological Factors that Change. Mean Annual Temperature
Seasonal Cycling
Annual Production
Current Patterns
References
12 Energetics. Introduction
A Model Energy Budget
Digestibility of Biomolecules
Energy Value of Biomolecules
Measuring Metabolic Rate
Oxygen Consumption Rate‐Modifying Factors
Activity
Experimental Protocol
Routine Metabolic Rate
Animal Size as a Modifier of Metabolism
Life History Strategies
Metabolism and Composition of Pelagic Species
Metabolism of Euphausiids, Decapods, Mysids, and Amphipods
Proximate Composition of Pelagic Decapods, Mysids, and Euphausiids
Terminology
Trends with Depth of Occurrence
Seasonal Changes
Trends Across Systems
Metabolism of Mesopelagic Fishes
Proximate Composition of Pelagic Fishes
Trends with Depth of Occurrence
Trends Across Systems
Energy and Life History in the Midwater Fauna
Midwater Fishes
The Cephalopods
Pteropods, Nemerteans, Annelids, Salps, and Pyrosomes
The Cnidaria and Ctenophora
Conclusions
References
Appendix A Classification of the Chordata
References
Glossary
Index
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Joseph J. Torres
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where I0 is the intensity of the incident light, I x is the intensity after traveling through distance x, and α λ is the attenuation coefficient, a function of wavelength.
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