Flatfish Metamorphosis

Foreword
Preface
Contents
Chapter 1: General Introduction of Flatfish Metamorphosis
	1.1 Brief Introduction of Metamorphosis in Various Fishes
		1.1.1 How Does Metamorphosis Occur in Fish?
		1.1.2 Brief Introduction of Larval Metamorphosis in Various Fishes
			1.1.2.1 Metamorphosis of Lampreys
			1.1.2.2 Metamorphosis of Elopomorphs
				Metamorphosis of Elopiforms
				Metamorphosis of Albuliforms
				Metamorphosis of Anguilliforms
			1.1.2.3 Metamorphosis of Groupers
			1.1.2.4 Metamorphosis of Gobiids
			1.1.2.5 Metamorphosis of Carapidae (Pearlfish)
			1.1.2.6 Metamorphosis of Flatfish (Pleuronectiformes)
	1.2 Introduction of the Metamorphic Events in Various Flatfishes
		1.2.1 Metamorphosis of Japanese Flounder
		1.2.2 Metamorphosis of Barfin Flounder
		1.2.3 Metamorphosis of Slime Flounder
		1.2.4 Metamorphosis of Summer Flounder
		1.2.5 Metamorphosis of Southern Flounder
		1.2.6 Metamorphosis of Winter Flounder
		1.2.7 Metamorphosis of Turbot
		1.2.8 Metamorphosis of European Plaice
		1.2.9 Metamorphosis of Atlantic Halibut
		1.2.10 Metamorphosis of Spotted Halibut
		1.2.11 Metamorphosis of Brown Sole
		1.2.12 Metamorphosis of Dover Sole
		1.2.13 Metamorphosis of Chinese Tongue Sole
	1.3 Implications of THs in Metamorphic Flatfishes
		1.3.1 Development of the Thyroid Gland and Changes in THs Level in Flatfishes
		1.3.2 Role of THs in Tissue Development during Metamorphosis
		1.3.3 Sensitivity of Metamorphic Events to THs during Larval Development
		1.3.4 Hypothalamo-Pituitary-Thyroid Axis
		1.3.5 Factors That Influence THs Level during Metamorphosis
		1.3.6 Gene Expression Patterns of Deiodinase Enzymes and THs Receptors during Metamorphosis in Flatfishes
	1.4 Genes Involved in Flatfish Metamorphosis
	1.5 MicroRNA Expression Profile During Metamorphosis in Flatfishes
	1.6 Summary
	References
Chapter 2: Developmental Relationships among Metamorphic Events
	2.1 Relationship between Cranial Deformation and Eye Migration
		2.1.1 Cranial Bones Deform Gradually During Metamorphosis in Flatfish
		2.1.2 Cranial Asymmetry in Different Eye Variants of Flatfishes
	2.2 Relationship Between Eye Migration and Body Swim-Posture Changes
		2.2.1 Development of Lateralized Behavior During Flatfish Metamorphosis
		2.2.2 Variants with Different Activities in the Artificial Senegalese Sole Population
	2.3 Relationship Between Eye Migration and Dorsal Fin Development
		2.3.1 Relationship Between Eye Migration and Anterior Extension of the Dorsal Fin
		2.3.2 Relationship Between Eye Migration and Elongation and Regression of the Dorsal Fin
	2.4 Relationship Between Eye Migration and Left/Right Asymmetrical Pigmentation
	2.5 Left/Right Asymmetrical Pigmentation Depends on Body Swim-Posture Change
	2.6 Summary
	References
Chapter 3: New Tissue Models for Explaining Eye Migration
	3.1 Diversity of Eye Location in Flatfishes
	3.2 The Hypotheses on the Eye Migration in Flatfishes
		3.2.1 The Eye Migration Thought to Be Associated with Brain
		3.2.2 The Eye Migration Thought to Be Associated with Dorsal Fin
		3.2.3 The Eye Migration Thought to Be the Result from the Resorption of the Part of the Cranium
		3.2.4 The Eye Migration Thought Be Caused by a Twisting of the Skull
		3.2.5 The Eye Migration Thought to Be Pulled by a Ligament
		3.2.6 Hypothesis About the Migrating Eye Forcing a Passage Through the Head
	3.3 Proliferating Cells in Suborbital Area Drive Eye Migration During Metamorphosis in Flatfish
		3.3.1 Normal Metamorphic Stages Defined in Three Flatfish Species
		3.3.2 Left/Right Asymmetrical Distribution of the Proliferating Cells in Suborbital Region of Left/Right Eye Before Initial Ey...
		3.3.3 Eye Migration Retarded by the Inhibitor of Cell Division Microinjected into the Suborbital Area of Blind Side
		3.3.4 Distortion of Frontal Bones Caused by Eye Movement
		3.3.5 Our Model Proposed Based on our Finding in Cell Proliferation
	3.4 Investigation on Eye Shapes During Flatfish Metamorphosis and Adult Flatfish
		3.4.1 Change of Eye Shape During Flatfish Metamorphosis
		3.4.2 The Deformation of Eye Morphology Shape During Metamorphosis Could Be Stopped by Colchicine
		3.4.3 The Difference of Eye Diameter Between Both Eyes in Different Flatfishes
		3.4.4 The Eyes Ever Been Pushed During Flatfish Metamorphosis
	3.5 The Role of Cell Apoptosis for Eye Migration During Metamorphosing Flounder
	3.6 The Role of Autophagy During Eye Migration in Flatfish Metamorphosis
	3.7 Summary
	References
Chapter 4: Molecular Basis of Eye Migration During Flatfish Metamorphosis
	4.1 Thyroid Hormone Regulating Eye Migration
		4.1.1 Thyroid Hormone Might Mediate Eye Migration in Various Flatfishes
		4.1.2 Thyroid Hormone Regulating Eye Migration Direct During Metamorphosis in Japanese Flounder
	4.2 The Role of Retinoic Acid in Modulating Eye Migration Via Cross-Talk with Thyroid Hormones in Japanese Flounder
		4.2.1 Does Eye Migration Regulated Only by Thyroid Hormone in Flatfish
		4.2.2 Coincident Genes Expression Patterns of the Thyroid Hormone and Retinoic Acid Signaling Pathways
		4.2.3 The Eye Migration Inhibited by Retinoic Acid in Japanese Flounder
	4.3 Some Genes for Cell Proliferation Might be Involved in Eye Migration
	4.4 Environment Factors Causing Flatfish Incomplete Eye Migration During Metamorphosis
		4.4.1 Nutrition of Live Prey
		4.4.2 Vitamin A
		4.4.3 The Effect of Photoperiod on Eye Migration
		4.4.4 Temperature
		4.4.5 Environment Pollution
	4.5 Summary
	References
Chapter 5: Molecular Basis of Frontal Bones Deformation During Metamorphosis
	5.1 Deformation Process of Frontal Bones During Flatfish Metamorphosis
	5.2 Study History of the Molecular Basis of Frontal Bone Deformation
	5.3 The Relationship Between the Frontal Bone Deformation and Eye Migration
	5.4 The Roles of Cell Proliferation, Cell Apoptosis, and Cell Autophagy During the Process of Front Bone Deformation in Japane...
	5.5 The Molecular Basis of Front Bone Deformation on the Aspect of Cell Apoptosis
		5.5.1 Deformation of Frontal Bones Induced by the Mechanical Force from the Contact of the Up-Migrating Eye on the Blind Side
		5.5.2 Classical Apoptosis Pathway Involved in the Distortion of Frontal Bones
	5.6 The Role of Thyroid Hormone for Front Bone Deformation
	5.7 Summary
	References
Chapter 6: Molecular Basis of Dorsal Fin Elongation and Regression During Metamorphosis
	6.1 Dorsal Fin Development in Fish
	6.2 The Process of Dorsal Fin Ray Elongation and Regression in some Flatfishes
	6.3 The Molecular Basis of Dorsal Fin Elongation in Japanese Flounder
		6.3.1 Morphological Changes During Dorsal Fin bud Formation and Skeletogenesis Process in Japanese Flounder Larvae
		6.3.2 Cellular Origins of the Dorsal Fin bud
		6.3.3 Position and Formation of Dorsal Fin bud
		6.3.4 Dorsal Fin bud Formation Regulated by Shh
		6.3.5 Some Genes Expressed in both Dorsal fin bud and fin Fold
	6.4 The Role of Thyroid Hormone in Regulating Dorsal Fin Development in Japanese Flounder
		6.4.1 The Distribution of Thyroid Hormone and Gene Expression of THs Signal Pathway in the bud During Dorsal Fin Development
		6.4.2 Role of the Thyroid Hormones in Dorsal Fin bud Formation and Ray Development
		6.4.3 The Role of the Thyroid Hormones in Dorsal Fin Elongation and Regression During Metamorphosis of Japanese Flounder
	6.5 Summary
	References
Chapter 7: Molecular Basis of Left/Right Asymmetrical Pigmentation during Metamorphosis
	7.1 The Establishment of Left/Right Asymmetrical Pigmentation in Flatfish
	7.2 Hypotheses on Left/Right Asymmetrical Pigmentation in Flatfish
		7.2.1 Previous Hypotheses
		7.2.2 Our Hypothesis on Left/Right Asymmetrical Pigmentation in Flatfish
	7.3 The Roles of Phototransduction Pathways and Retinoic Acid Signaling in Establishing Asymmetric Pigmentation
		7.3.1 The Roles of Phototransduction Pathways in Establishing Asymmetric Pigmentation in Japanese Flounder
		7.3.2 The Roles of Retinoic Acid Signaling in Establishing Asymmetric Pigmentation in Japanese Flounder
	7.4 Malpigmentation in Flatfish Aquaculture Industry
		7.4.1 Malpigmentation Pattern in Flatfish
			7.4.1.1 Types of Pigmentation Defects
		7.4.2 The Progression of Hypermelanosis in Blind Side
		7.4.3 The Progression of Pseudoalbino in Ocular Side
	7.5 Nutrition and Environmental Factors Causing Malpigmentation in Ocular Side in Flatfish Aquaculture
		7.5.1 Nutrition
			7.5.1.1 Vitamin A
			7.5.1.2 Vitamin D3
			7.5.1.3 Polyunsaturated Fatty Acid
		7.5.2 Light
		7.5.3 Temperature
	7.6 Environmental Factors Causing Staining-Type Hypermelanosis in Flatfish Aquaculture
		7.6.1 Environmental Factors
			7.6.1.1 Rearing Density
			7.6.1.2 Background Color
			7.6.1.3 Burrowing Substratum
		7.6.2 Hypothesis on Occurrence of Staining-Type Hypermelanosis
	7.7 Genetic Screening for Malpigmentation in Artificial Breeding Flatfish
		7.7.1 Genetic Screening for Malpigmentation in Artificial Breeding Population of Japanese Flounder
		7.7.2 Genetic Screening for Malpigmentation in Artificial Breeding Population of Chinese Tongue Sole
	7.8 Summary
	References
Chapter 8: Molecular Basis of Body Depth Change during Metamorphosis
	8.1 Brief Introduction on Body Shape Change during Early Life History in Various Fishes
	8.2 The Body Height Increased Gradually during Flatfish Metamorphosis
	8.3 The Role of Cell Proliferation for the Change of Body Depth During Flatfish Metamorphosis
	8.4 The Role of Thyroid Hormone in Regulating Body Depth Change in Flatfish
		8.4.1 The Body Height Change Regulated by Thyroid Hormone Via Cell Proliferation During Metamorphosis
		8.4.2 Strong Expression of Thyroid Hormone Receptors Along Dorsal-Ventral Margins During Metamorphosis
		8.4.3 Does the Change of Body Shape During Early Development of Non-Flatfish Is Regulated by Thyroid Hormone?
	8.5 Summary
	References
Chapter 9: Genetic Basis for Eye Migration in Flatfish
	9.1 Different Molecular Mechanisms to Determine Which Eye Migrates and the Migration Distance
	9.2 A New Hypothesis to Explain Eye Asymmetry During Early Life History of Flatfishes
	9.3 Is Eye Asymmetry Regulated by the Nodal-Lefty-pitx2 (NLP) Pathway?
	9.4 Screening of Reverse Eye Genes by Using Comparative Transcriptomic Analysis
	9.5 Duplication of Thyroid Hormone Receptor TRβ Gene Found in Flatfishes
	9.6 Summary
	References
Chapter 10: Evolutionary Origin of Left-Right Eye Asymmetry
	10.1 Evolutionary Debate on the Origin of Asymmetry in Flatfish
	10.2 Phylogenetic Pattern of Asymmetry in Flatfish
	10.3 Our Hypothesis for the Evolutionary Origin of Left-Right Eye Asymmetry
	10.4 Is the Evolution of Eye Migration in Flatfishes Convergent?
	10.5 Is it Possible That One Gene Determines Eye Asymmetry?
	10.6 Inheritance of Asymmetry in Flatfish
	10.7 Genome-Wide Screening for the Locus of Eye Migration in Flatfish
	10.8 Summary
	References