Sensory Biology of Aquatic Animals

Source-Localization Algorithms.- Frequency Analysis.- A Frequency-Response Dilemma.- Old Mistakes and New Solutions.- Physics of Lateral-Line and Inner-Ear Sense Organs.- The Macula Neglecta.- The Boundary Layer Revisited.- Summary.- 5 Underwater Sound as a Biological Stimulus.- The Nature of Sound.- Governing Equations.- Plane Waves.- Spherical Waves.- Pressure-Velocity Ratios.- The Microvar.- Adequate Stimulus.- The Nature of the Coupling Between the Otolith and Cilia.- The Response of Hair Cells Without the Otolith.- Directional Cues.- Ambient Noise.- Underwater Sound Propagation.- Reflection.- Scattering.- Refraction.- Shallow Water Propagation.- Differences Between Underwater and Atmospheric Sound.- Summary.- 6 Detection of Weak Electric Fields.- Electric Fields in Natural Waters.- The Discovery of Bioelectric Fields.- The Origin of Bioelectric Fields.- The Representation of Bioelectric Fields.- Environmental Electric Fields.- Fields Induced by the Recipient Animal.- Behavioral Responses to Weak Electric Fields.- A New Sensory Modality Revealed.- Prey Detection, Sensitivity, and Frequency Response.- Orientation to Environmental Fields.- Detection of the Earth's Magnetic Field.- Physics of Peripheral Sensory Apparatus.- Resistivity of the External Medium.- Potential Distribution Within the Animal.- Differential Operation and Common-Mode Rejection.- Physical Properties of Ampullary Canals.- Sensory Transduction and Nerve Coding.- Electroreceptor Mechanisms.- Detection of Temporal Field Characteristics.- Detection of Spatial Field Characteristics.- Sensory Mechanisms and Central Processing.- A Prey-Detection Algorithm.- Central Processing.- Summary.- II. Behavior: Survival Functions and Stimulus Acquisition.- 7 Behavioral Ecology and Sensory Biology.- Discriminating Food from Nonfood Items.- Food: Which Food Types Should be Eaten?.- How to Avoid Being Eaten.- With Whom to Mate?.- Social Relations.- Summary and Conclusions.- 8 Sensory Performance, Behavior, and Ecology of Fish.- Thresholds.- "Noise".- Directional Perception.- Light.- Sound.- Chemicals.- The Enhancement of Sensory Awareness.- Multiple Stimulation.- Summary and Conclusions.- 9 Social Communication in the Aquatic Environment.- What Is Communication?.- Specialized Communication Sensors.- Special Problems of Communication in Aquatic Environments.- Speed of Signal Propagation.- Active Space.- Signal Attenuation Due to Geometrical Spreading, Absorption, and Scattering.- Background Noise and Receptor Sensitivity.- Signal Source Localization.- Electrical and Acoustic Communication in Mormyrid Fish.- Signal Range.- Acoustic Signalling in Pollimyrus isidori.- Audition in Mormyrids.- Nature of Electrical Signalling in Mormyrids.- Localization of Electric Signals by Mormyrids.- Signaller Identification and Signal Recognition.- Summary.- 10 The Comparative Neurology of Expectation: Stimulus Acquisition and Neurobiology of Anticipated and Unanticipated Input.- Stimulus Acquisition in Electroreception.- Plastic Corollary Discharge.- Efference Copy vs. Corollary Discharge.- Spike and Wave Signs of Central Expectation and Fluctuations of Readiness.- Fluctuations in the Expectation of Input.- Strategies for New Research.- Summary.- III. Signal Extraction: Receptor Cells and Sensory Systems.- III. A. Chemoreception.- 11 The Diversity of Chemoreceptors.- Behavioral Diversity.- Functional Diversity.- Generalists and Specialists.- Dynamic or Static Endings.- Internal and External Detection.- Structural Diversity.- Organ Systems.- Exposed or Protected Detectors.- Few or Many?.- Cilia.- Biochemical Structure.- Olfaction and Taste: Is There a Distinction?.- Proposed Definitions.- Diversity of Stimulatory Substances.- Numbers and Location of Receptors.- Sensory Cilium.- Crustacean Chemoreceptors.- Aesthetasc Hairs.- Esophageal Chemoreceptor in Homarus.- Funnel Canals in Dactyls of Decapod Crustacea.- Hedgehog Hairs.- Dorsal Organ of Decapod Larvae.- Conclusion.- Summary.- 12 Peripheral Filters and Chemoreceptor Cells in Fishes.- Olfactory Organ and Receptor Cell Distribution.- Taste Bud Anatomy and Distribution.- Receptor Mechanisms.- Sensitivity and Specificity to Amino Acids.- "Peripheral Filters": Transduction Processes and Receptor Site Types.- Effects of Chemical Mixtures.- Summary.- 13 Organization of Chemosensory Systems Within the Brains of Bony Fishes.- Olfactory System.- Olfactory Nerve.- Olfactory Bulb.- Olfaction in Flatfish: Lopsided Smelling.- Terminal Nerve.- Gustatory System.- Gustatory Systems in Central Nervous System.- Common Chemical Sense.- Primary Sensory Nerves and Their Site of Central Termination.- Central Pathways.- Summary.- 14 Chemoreceptor Cells in Aquatic Invertebrates: Peripheral Mechanisms of Chemical Signal Processing in Decapod Crustaceans.- Stimulus Filtering.- Crustacean Chemoreceptor Cells.- General Constraints on Receptors.- The Determination of Stimulus Quality.- Narrow Tuning.- Mixture Effects.- The Determination of Stimulus Quantity.- Behavioral Thresholds.- Physiological Thresholds.- Range Fractionation.- Detection in Chemical Noise.- Interspecific Differences in Sensitivity.- Summary.- 15 Integration of Chemosensory Information in Aquatic Invertebrates.- Structural Organization.- Functional Organization.- The Discrimination of Odor Quality.- The Discrimination of Odor Quantity.- The Discrimination of Spatial and Temporal Aspects of Odors.- Summary.- III. B. Vision.- 16 Vision in Marine Invertebrates.- Invertebrate Visual Pigments.- Eye Design in Marine Invertebrates.- Invertebrate Optical Systems.- Summary.- 17 Vision and Mating Behavior in Limulus.- Circadian Rhythms in Retinal Structure and Function.- Laboratory Studies of Circadian Rhythms in Visual Sensitivity.- The Role of Vision in Limulus Mating Behavior.- Summary.- 18 Aquatic Adaptations in Fish Eyes.- Physical Constraints on Eye Design.- Light Collection.- Resolution.- The Structure of Fish Eyes.- Focus and Resolution.- Lens Quality.- The State of Focus.- Optic Field and Binocular Field.- Retinal Function During Growth.- Retinal Growth.- Visual Acuity and Retinal Growth.- Visual Sensitivity and Retinal Growth.- The Regulation of Retinal Growth.- Teleost Photoreceptors.- Structure.- Function.- Spatial Organization.- Retinomotor Movements.- Description.- Function.- Control.- Summary.- 19 Optics of Amphibious Eyes in Vertebrates.- Vision in Air and Water.- The Metamorphosis of Amphibian Ocular Optics.- Super Accommodation in Eyes of Diving Birds.- Conclusion-Questions Asked, Answers Needed.- Summary.- 20 How to Be Unseen: An Essay in Obscurity.- The Problem Defined.- Seeing Objects.- Abbreviated Natural History of Flatfish and Their Changes in Appearance.- Adaptive Behavior and Its Structural Basis.- Adaptive Parameters: A Note on Methodology.- Design Materials.- Chromatophore Types.- Morphological Markers.- Control Processes.- Nervous Pathway for Melanophore Control: Peripheral Aspects.- Nervous Pathway for Melanophore Control: Central Aspects.- Melanophore Dispersion: How Does It Happen?.- Peripheral Control of Other Chromatophores.- Camouflage as Visual Processing of Pattern and Texture.- Afterword.- Summary.- 21 The Visual System in Teleost Fishes: Morphological Patterns and Trends.- Optic Nerve.- Connections of the Retina.- Retinofugal Projections.- Retinopetal Projections.- Conclusions.- III. C. Mechanoreception.- III. C. 1. Hydrodynamic Reception.- 22 Diversity of Lateral Line Systems: Evolutionary and Functional Considerations.- Basic Anatomy of Mechanoreceptive Lateral Line System.- Variations in Canal Structure.- Variations in Neuromast Morphology.- Spatial Distribution of Receptor Organs.- Basic Distribution Pattern of Canals.- Basic Distribution of Superficial Neuromasts.- Morphological Constraints in the Evolution of Lateral Line Systems.- Convergent Trends.- Functional Implications of Morphological Variation.- Summary and Conclusions.- 23 Mechanical Factors in the Excitation of the Lateral Lines of Fishes.- Methods.- Steps in Excitation.- Step 1. The Source to Net Acceleration of the Medium Adjacent to the Lateral Line.- Step 2. From Net Motions in the Medium Close to Lateral Lines to Motions Inside Lateral Line Canals.- Step 3. From Motions of Liquid Inside Lateral Line Canals to Forces on Cupulae.- Step 4. The Force on the Cupula to the Activity in the Nerve.- Step 5. Patterns Between Neuromasts.- The Importance of Lateral Line to the Sprat.- The Behavior of Other Types of Lateral Line.- Poromitra.- Anoplogaster.- Penaeidae (Crustacea; Decapoda; Natantia).- A Discussion.- 24 Prey Identification and Prey Localization in Surface-Feeding Fish and Fishing Spiders.- Physical Principles.- Characteristics of Prey and Nonprey Wave Stimuli.- Water Surface as Stimulus Transmission Channel.- Sound Waves.- Useful Stimulus Parameters for Distance Determination.- Distance Determination in Surface-Feeding Fish.- Electrophysiological Results.- Behavioral Results.- Sound.- Some Speculative Considerations.- Stimulus Discrimination in Fishing Spiders.- Useful Stimulus Parameters for Prey and Nonprey Wave Discrimination.- Behavioral Results.- Diagrammatical Model of Results.- Summary.- 25 Surface Wave Reception in Invertebrates and Vertebrates.- A Note on the Use of the Term "Signal".- Properties of Surface Waves.- Surface Wave Sensory Structures.- Invertebrates.- Vertebrates.- Amplitude and Frequency: Range, Sensitivity, and Discrimination.- Sensory and Behavioral Mechanisms: Predators and Prey.- Prey Identification.- Prey Localization.- Locating Wave Source Distance: Accuracy and Mechanisms.- Surface Waves and Predator Avoidance.- Communication by Surface Wave Signals.- Invertebrates.- Vertebrates.- Summary.- 26 The Representation of Hydrodynamic Parameters in the CNS of the Crayfish Procambarus.- A Circuit Monitoring Water Displacements.- Receptor Level.- The Arrangement of Thresholds and Frequency Response.- The Directionality of Response in Sensory Cells and Ventral Cord Interneurons.- Inhibitory Contrast Enhancements.- Discussion.- Displacement Receptors in the Water and Air.- Stimulus Intensity Coding.- Signal Frequency Range in this Pathway.- The Directionality of Response in this Pathway.- Neuronal Contrast Enhancements in the Pathway.- Dual Innervation of Receptors and Directional Subchannels in the Pathway.- Summary.- III. C. 2. Hearing.- 27 Role of the Fish Ear in Sound Processing.- The Fish Ear!.- Otolith Organs-Structure.- Pathways for Sound to the Ear.- What Do We Know About Signal Analysis?.- Mechanisms of Inner Ear Stimulation.- Sound Localization and Enhancement of Signal-to-Noise (S/N) Ratio.- Enhancement of S/N Ratio.- Sound Source Localization: Vectorial Analysis.- Frequency Analysis.- Temporal Analysis.- Cilia Length.- Regionalization.- Direct vs. Indirect Comparison.- Additional Considerations.- 28 Peripheral Adaptations for Spatial Hearing in Fish.- Cues Used for Directional Hearing in Air.- The Problem of Directional Hearing Underwater.- Early History.- Contemporary Views.- Directional and Nondirectional Response of Ears.- Principles of Stimulation and Directional Coding Hypotheses.- Problems to Be Solved.- Physiological Studies of Directional Hearing.- Central Nervous System.- Other Aspects of Directional Hearing.- Cocktail Party Effect.- Reflex Orientation.- Cues for the Structure of Acoustic Environments.- Summary and Conclusions.- 29 Central Connections of the Octavolateralis System: Evolutionary Considerations.- Patterns of Organization.- Primitive Pattern of Organization.- Patterns of Organization in Bony Fish.- Electrosensory Systems.- Evolutionary History of Electroreception.- Possible Origins of Teleost Electrosensory Pathways.- The Origin of the Octavolateralis System.- Summary.- Auditory Systems.- The Evolution of Central Auditory Pathways.- Inner Ear Receptor Projections.- Higher-Order Auditory Projections.- Audition and the Fish-Amphibian Transition.- Summary.- Conclusion.- III. C. 3. Equilibrium.- 30 Morphological Diversity of Equilibrium Receptor Systems in Aquatic Invertebrates.- Gravity Receptor Systems.- Gravity Orientation Without Specialized Gravity Receptor Systems.- Gravity Receptor Systems Using Cell Organelles.- Gravity Receptor Systems Lacking Hair Cells or Cuticular Sensory Hairs.- Gravity Receptor Systems with Hair Cells.- Gravity Receptor Systems with Cuticular Sensory Hairs.- Angular Acceleration Receptor Systems.- Angular Acceleration Receptor Systems with Hair Cells.- Angular Acceleration Receptor Systems with Cuticular Sensory Hairs.- Conclusions.- Summary.- 31 Equilibrium in the Vertebrates: Signals, Senses, and Steering Underwater.- Posture and Equilibrium.- Signals and Senses.- Vestibular Apparatus.- Variation in Vestibular Structure.- Vestibular Ultrastructural Organization.- Vestibular Physiology.- Steering: Posture and Locomotor Control.- Unusual Posture and Otolith Organ Structure.- Canal Sensitivity and Head Movements.- Extra-Vestibular Modulation of Posture.- Visual-Vestibular Interaction in Posture.- Other Sensory Influences on Posture.- Future Questions.- Conclusions.- III. D. Electroreception.- 32 The Electroreceptors: Diversity in Structure and Function.- Taxonomic Distribution.- Anatomy of Ampullary Electroreceptors.- Elasmobranches.- Teleosts.- Stimulus and Receptor Frequency Sensitivity.- Functional Anatomical Considerations and Biophysics.- Polarity and Sensitivity.- Noise Cancellation.- Tuberous Receptors.- Tuberous Receptor Morphology.- Electric Organ Discharge.- Enhancing Sensitivity: Impedance Matching and Noise Reduction.- Frequency Tuning of Tuberous Receptors.- Biophysics and the Origin of Tuning.- Coding Mechanissm of Tuberous Receptors.- Summary.- 33 The Neuronal Basis of Electrosensory Perception and Its Control of a Behavioral Response in a Weakly Electric Fish.- Neuroethological Research Strategies.- Neural Mechanisms of Jamming Avoidance Response (JAR).- The Study of Central Nervous Rules of Computation by Behavioral Experiments.- The Exploration of Neuronal Implementations.- General Lessons from the Study of the JAR.- Technical Difficulties in the Study of the CNS of Vertebrates.- The Nature of Distributed Parallel Processor.- The Emergence of Feature Detection and Behavioral Stimulus Specificity.- The Relation Between Neuronal Structural and Function.- Similarities to Neuronal Organization of Directional Hearing.- Speculations About the Evolution and Ontogeny of Neuronal System.- Summary.- IV. Adaptation and Sensory Systems.- 34 Sensory and Other Neural Traits and the Adaptationist Program: Mackerels of San Marco?.- Definitions of Adaptation.- Alternate Explanations of Design.- Evaluation of Hypotheses of Design.- Summary.- Author Index.- Animal Index.