The Ecology of Deep-Sea Hydrothermal Vents

Cover Page
Half-title Page
Title Page
Copyright Page
Dedication Page
Contents
Preface
Acknowledgments
1. The Non-Vent Deep Sea
	1.1 The Physical Environment in the Deep Sea
	1.2 The Deep-Sea Fauna
	1.3 Deep-Sea Diversity
	1.4 Biogeography and Population Genetics
	1.5 Biochemical and Physiological Adaptations to the Deep-Sea Environment
	1.6 Benthopelagic Coupling between Surface Productivity and the Deep Sea
	1.7 Rates of Biological Processes in the Deep Sea
	1.8 The Vent Contrast
	References
2. Geological Setting of Hydrothermal Vents
	2.1 What Are Mid-Ocean Ridges?
		2.1.1 How Spreading Rates for Ridge Axes Are Determined
		2.1.2 Spreading Rates
		2.1.3 Segmentation
		2.1.4 Magma Supply and Spreading Rate
	2.2 Back-Arc and Fore-Arc Spreading Centers
	2.3 Seamounts
	2.4 Volcanic and Tectonic Seafloor Features
		2.4.1 Crustal Structure
		2.4.2 Volcanic and Tectonic Fissures
		2.4.3 Lava Lakes, Drainback Features, and Lava Pillars
		2.4.4 Axial Boundary Faults
		2.4.5 Lava Flow Morphologies
		2.4.6 Emplacement of Lavas and the Time-Course of a Diking Event
		2.4.7 Lava Dating
	2.5 Deep-Sea Hydrothermal Fields
		2.5.1 Missing Heat and Hydrothermal Cooling at Ridge Crests
		2.5.2 Sulfide Deposits
			Morphological Variations
				Columnar Chimneys and Black Smokers
				White Smokers
				Beehives and Flanges
				Complex Sulfide Mounds
				Weathering of Seafloor Sulfides
			Dimensions and Ages of Active Hydrothermal Fields
		2.5.3 Low-Temperature Diffuse Flows
		2.5.4 Sediment-Hosted Hydrothermal Systems
		2.5.5 Ophiolites
	Appendix
	References
3. Chemical and Physical Properties of Vent Fluids
	3.1 Submarine Hydrothermal Circulation Cells: High-Temperature Reaction Zones
	3.2 Phase Separation
	3.3 Flow Rates, Transit Times, and Temperature of Formation
	3.4 End-Member Fluids
		3.4.1 Composition
			Basic Controls on Chemistry
		3.4.2 Magmatic Inputs
		3.4.3 Evolution of Vent-Fluid Chemistry
		3.4.4 Back-Arc Fluid Chemistries
	3.5 Thermal Radiation
	3.6 Axial Low-Temperature, Diffuse-Flow Chemistry
		3.6.1 Flow Rates, Temperature, and Temperature Variability
		3.6.2 Silicate
		3.6.3 Sulfide
		3.6.4 Oxygen
		3.6.5 Profiles of Oxygen, Sulfide, Silicate, and Temperature
		3.6.6 Methane, Manganese, and Iron
		3.6.7 Nitrogen and Phosphorus Compounds
	3.7 Flank Low-Temperature Fluids
	3.8 Global Fluxes and the Hydrothermal Influence on Ocean Chemistry and Currents
	References
4. Hydrothermal Plumes
	4.1 Anatomy of a Black-Smoker Plume
		4.1.1 Orifice
		4.1.2 Buoyant Plume
		4.1.3 Effluent Layer
	4.2 Megaplumes
	4.3 Spatial and Temporal Distributions of Plumes
		4.3.1 Relationship between Plume Distributions and Geophysical Parameters
	4.4 Plume-Driven Mesoscale Circulation
		4.4.1 Plume Vortices
		4.4.2 Advection and Downwelling
		4.4.3 Basin-Scale Circulation
	4.5 Diffuse-Flow Plumes
	References
5. Microbial Ecology
	5.1 Autotrophic Organisms at Vents
		5.1.1 Nomenclature
		5.1.2 Aerobic and Anaerobic Chemoautotrophy at Vents
			Methanotrophy
		5.1.3 Carbon Dioxide Fixation
		5.1.4 Mixotrophy
		5.1.5 Net Chemoautotrophic Production in Free-Living Hydrothermal-Vent Microorganisms
			Alternatives to Chemoautotrophy
				Organic Thermogenesis Hypothesis
				Detrital Thermal Alteration Hypothesis
	5.2 Ecology of Free-Living Microorganisms
		5.2.1 Microbial Habitats
		5.2.2 Hyperthermophiles and Superthermophiles
			Flange Microbial Ecology and the Archaea
			Microorganisms in Black-Smoker Fluids
			The \"Endeavour Model\"
			The Subsurface Biosphere
		5.2.3 Plume Microbiology
		5.2.4 Suspended Microbial Populations
		5.2.5 Microbial Community Composition
			Dominance of a Single Bacterial Phylotype at a Mid-Atlantic Ridge Vent
			Diversity and Community Structure in Microbial Mats, Loihi Seamount
			Sulfur-Oxidizing Heterotrophs at Vents
		5.2.6 Bacterial Blooms
		5.2.7 Microbial Mats
		5.2.8 The Link between Chemoautotrophic and Photosynthetic Processes
	5.3 A Search for In Situ Bacterial Photosynthesis
	5.4 Microbial Genesis of Hydrothermal Mineral Deposits
	5.5 Microbial Exploitation of Particulate Sulfides
	5.6 Biotechnology
	References
6. Symbiosis
	6.1. Discovery
		6.1.1 Sustenance of Gutless Tubeworms
		6.1.2 Endosymbiotic Bacteria in Vent Mollusks
		6.1.3 Episymbionts
	6.2 Methanotrophic Symbioses
		6.2.1 Dual Symbioses
		6.2.2 Methanotrophs in Sponges
	6.3 Adaptive Characteristics of Symbiosis
	6.4 Host Nutrition
		6.4.1 Digestive Enzymes
	6.5 Symbiont Phylogeny
		6.5.1 Endosymbiont Phylogeny and Host Fidelity
		6.5.2 Episymbiont Phylogeny
	6.6 Symbiont Acquisition
	References
7. Physiological Ecology
	7.1 Novel Metabolic Demands
	7.2 Riftia pachyptila
		7.2.1 Anatomy of a Tubeworm
		7.2.2 The Tubeworm Environment
		7.2.3 Adaptations for Carbon Uptake and Transport in Riftia pachyptila
			Host Respiratory Inorganic Carbon
			Environmental Sources of Inorganic Carbon and the Role of Carbonic Anhydrase
			pH Regulation
			Carbon Transport
			Inorganic Carbon Capacity
			Carbon Fixation Rates
		7.2.4 Sulfide
			Sulfide Toxicity
			Sulfide Uptake and Transport
			Coupling of Sulfide Detoxification and Energy Exploitation
		7.2.5 Oxygen
		7.2.6 Nitrogen
			Nitrate Respiration
	7.3 Seep Vestimentiferans and Methanotrophic Pogonophorans
	7.4 Vent and Seep Bivalve-Mollusk Symbioses
		7.4.1 Calyptogena magnified
		7.4.2 Bathymodiolid Mussels
			Bathymodiolus thermophilus
			Methanotrophic Mussels
		7.4.3 Other Mollusk Symbioses
	7.5 Physiological Ecology of Episymbiont-Invertebrate Associations
		7.5.1 Alvinella pompejana
	7.6 Sulfide Detoxification
	7.7 Growth Rates
	7.8 Thermal Adaptations
		7.8.1 Indices of Thermal Tolerance and Adaptation
			Thermal Tolerance in Alvinellid Species
	7.9 Heavy Metals and Petroleum Hydrocarbons
	7.10 Sensory Adaptations
		7.10.1 Novel Photoreceptors in Vent Shrimp
		7.10.2 Chemoreception
	References
8. Trophic Ecology
	8.1 The Food Web
		8.1.1 The Rose Garden Food Web
	8.2 Biological Sleuthing: Biomarker Assays
		8.2.1 Stable Isotope Techniques
			Notation
			Stable Isotope Evidence for the Role of Free-Living Microorganisms in Vent Food Webs
		8.2.2 Fatty Acids, Sterols, and Carotenoids
			Fatty-Acid Nomenclature
			Fatty-Acid Biomarkers
			Comparison of Lipid Characteristics of Tubeworms (Riftia pachyptila), Mussels (Bathymodiolus thermophilus), and Amphipods (Halice hesmonectes) on the East Pacific Rise
			Essential Fatty Acids
			Lipid-Condition Indices
			Sterols
			Carotenoids
	8.3 Integrated Approaches to Trophic Ecology
		8.3.1 Trophic Ecology of Vent Mussels, Bathymodiolus thermophilus
		8.3.2 Trophic Ecology of Vent Shrimp, Rimicaris exoculata, and an Anecdote about Who Eats Them
	8.4 Export of Chemosynthetic Production from Vents
	References
9. Reproductive Ecology
	9.1 Gametogenesis
		9.1.1 Evidence for Synchronous Gametogenesis
			Environmental Cues
			Recruited Synchrony
		9.1.2 Evidence for Asynchronous Gametogenesis
			Release of Gametes and Larvae
				Riftia pachyptila
				Bythograea sp.
				Calyptogena soyae
	9.2 Larval Development
		9.2.1 Vestimentifera
		9.2.2 Bathymodiolid Mussels
		9.2.3 Bythograeid Crabs
		9.2.4 Alvinocarid Shrimp
	9.3 Larval Dispersal and Retention
		9.3.1 Alvinellid Dispersal Model
		9.3.2 Plume Dispersal
		9.3.3 Megaplume Dispersal
		9.3.4 Mesoscale Flows
		9.3.5 Dispersal by Non-Larval Stages
	9.4 Settlement Cues
	9.5 Recruitment
	Appendix
	References
10. Community Dynamics
	10.1 The Early Work
	10.2 Dynamic Succession at Northeast Pacific Vents
		10.2.1 High-Resolution Time-Series Studies on the Juan de Fuca Ridge
	10.3 Community Dynamics on the Mid-Atlantic Ridge
	10.4 Eruptions
		10.4.1 The 9°N Event
		10.4.2 The CoAxial Event
		10.4.3 Sweepstakes versus Predictable Sequences
	References
11. Evolution and Biogeography
	11.1 Origins of Vent Fauna
		11.1.1 Immigrants from the Surrounding Deep Sea
		11.1.2 Immigrants with Close Shallow-Water Relatives
		11.1.3 Vent Taxa Shared with Other Chemosynthetic Ecosystems
			Taxonomic Position and Origin of the Vestimentifera
		11.1.4 Vent Taxa Shared with Both Other Chemosynthetic Ecosystems and Nonchemosynthetic Habitats
		11.1.5 Specialized Taxa Found Only at Hydrothermal Vents
		11.1.6 The \"Ancient\" Taxa
			Ancient Barnacles
			Ancient Mollusks
		11.1.7 The Newman and McLean Hypothesis of Relict Vent Faunas
			Hickman\'s Counterhypothesis
		11.2 Fossil Vent Communities
		11.3 Vent Ecosystems as Refuges from Major Planetary Extinction Events
		11.4 Species Diversity
		11.5 Taxonomic Cautionary Tales
			11.5.1 Cryptic Species
			11.5.2 Phenotypic Plasticity
			11.5.3 Ontogenetic Stages
		11.6 Biogeography
			11.6.1 Pacific Biogeographic Patterns
				Missing Mussels {Bathymodiolus thermophilus)
				Centers of Diversity along Linear Arrays of Habitat
				North America as a Biogeographical Barrier
				Mariana Hydrothermal-Vent Fauna
			11.6.2 Paleotectonic Controls on the Atlantic Vent Fauna
			11.6.3 Similarities among Global Vent Biogeographic Provinces
			11.6.4 Biogeography of Fast- versus Slow-Spreading Centers
			11.6.5 Physical Oceanography and Bathymetry
				The Romanche Fracture Zone
			11.6.6 Shallow-Water Vents
		11.7 Gene Flow and Genetic Diversity
		References
12. Cognate Communities
	12.1 Atlantic Sites
		12.1.1 Florida Escarpment (Gulf of Mexico)
		12.1.2 Louisiana Slope Hydrocarbon and Brine Seeps (Gulf of Mexico)
		12.1.3 The Laurentian Fan
		12.1.4 Barbados Subduction Zone
		12.1.5 North Sea Pockmarks
		12.1.6 Skagerrak Methane Seep
		12.1.7 The Francois Vieljeux
		12.1.8 Coral Reefs
	12.2 Pacific Sites
		12.2.1 Cascadia Subduction Zone
		12.2.2 Western Pacific Subduction Zones
			Kaiko Project
			Sagami Bay
		12.2.3 Peruvian Subduction Zone
		12.2.4 Monterey Canyon
		12.2.5 Northern California Methane Hydrate Field
		12.2.6 Guaymas Basin Transform Margin Seeps
		12.2.7 Shallow-Water Hydrocarbon Seeps
		12.2.8 British Columbia Fjords
		12.2.9 Aleutian Subduction Zone
	12.3 Whale Skeletons
		12.4 Fossil Seeps
		References
13. Hydrothermal Systems and the Origin of Life
	13.1 Earth\'s Early Environment
	13.2 Evolution of Hydrothermal Systems
	13.3 Heterotrophic versus Chemosynthetic Hypotheses for the Origin of Life
	13.4 Evidence for Thermophilic, Autotrophic Ancestors
		13.4.1 Wachterhauser\'s Outline for the Origin and Evolution of Life
		13.4.2 Synthesis of Organic Compounds in Hydrothermal Systems
	13.5 Extraterrestrial Hydrothermal Systems and the Search for Life in Outer Space
References
Index