Anatomy and Histology of the Domestic Chicken, 1st Edition
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Anatomy and Histology
of the Domestic Chicken
Edited by
Wael Khamas
College of Veterinary Medicine
Western University of Health Sciences
Pomona, USA
Josep Rutllant
College of Veterinary Medicine
Western University of Health Sciences
Pomona, USA
Copyright © 2024 by John Wiley & Sons, Inc. All rights reserved.
Published by John Wiley & Sons, Inc., Hoboken, New Jersey.
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Library of Congress Cataloging-in-Publication Data
Names: Khamas, Wael, editor. | Rutllant, Josep, editor.
Title: Anatomy and histology of the domestic chicken / edited by Wael
Khamas, Josep Rutllant.
Description: Hoboken, New Jersey : Wiley-Blackwell, [2024] | Includes
bibliographical references and index.
Identifiers: LCCN 2024003433 (print) | LCCN 2024003434 (ebook) | ISBN
9781119841722 (cloth) | ISBN 9781119841753 (adobe pdf) | ISBN
9781119841746 (epub) | ISBN 9781119841739 (oBook)
Subjects: MESH: Chickens–anatomy & histology
Classification: LCC SF767.P6 (print) | LCC SF767.P6 (ebook) | NLM SF
767.P6 | DDC 636.50891–dc23/eng/20240229
LC record available at https://lccn.loc.gov/2024003433
LC ebook record available at https://lccn.loc.gov/2024003434
Cover Design: Wiley
Cover Images: © Wael Khamas and Josep Rutllant
Set in 9.5/12.5pt STIXTwoText by Straive, Pondicherry, India
Dedication
To My beloved family. I dedicate this book to you with profound gratitude and a heart full of love. Your support, patience, and
endless encouragement have been the guide on this journey. May this book be a token of my appreciation and a reminder of
the unbreakable bonds we share.
Wael Khamas
This book is dedicated to my parents, who have consistently instilled in me the treasured values of learning, diligent work,
integrity, civility, and the significance of friendship. “Mai us podré tornar tot el que heu fet per mi” – I can never fully repay
you for all that you’ve done for me. To Helena and Laura, for the boundless love and unwavering support you have graciously
bestowed upon me. Lastly, to Joaquín Camón, a cherished mentor who introduced me to the world of anatomy and firmly
believed in my potential. You are all notes of my music.
Josep Rutllant
vii
Contents
List of Contributors xix
Foreword xxi
Preface xxiii
Acknowledgments xxv
Lists of Abbreviations xxvii
1 External Features of Chicken 1
Wael Khamas and Josep Rutllant
1.1 Skin 1
1.1.1 Epidermis 1
1.1.2 Dermis 2
1.1.3 Hypodermis (Subcutis) 2
1.1.4 Blood Supply of the Skin and Nerve Ending 2
1.2 Structures Associated with the Skin 2
1.2.1 Patagium (pl. patagia) 2
1.2.2 Operculum 3
1.2.3 Ornaments 3
1.2.3.1 Comb 3
1.2.3.2 Wattle 4
1.2.3.3 Ear Lobes 4
1.2.3.4 External Ear Opening 5
1.3 Feathers 5
1.3.1 Development 5
1.3.2 Description of Feathers 5
1.3.3 Ptilosis and Pterylosis 5
1.3.4 Feathers Color 6
1.3.5 Types of Feathers 7
1.3.6 Parts of Feather 7
1.3.7 Microscopic Description of Feather 8
1.3.8 Types of Flying Feathers 8
1.3.8.1 Primary Remiges and Their Coverts (also Known as Cover Feathers) 8
1.3.8.2 Secondary Remiges and Their Coverts 9
1.3.8.3 Rectrices and Their Coverts 9
1.3.8.4 Contour Feathers 9
1.4 Lamellar Corpuscles 9
1.5 Molting 9
1.5.1 Induction of Molting by Food Deprivation 10
1.5.2 Feather Picking 10
1.6 Debeaking 11
viii Contents
1.7 yelids Including Third Eyelid
E 11
1.8 Vent 11
1.9 Uropygial Gland 12
1.10 Brood Pad 12
1.11 Bursa Sterni 13
1.12 Legs 13
1.12.1 Podotheca 13
1.12.2 Scutes (Scales) 13
1.12.3 Digits 13
1.12.4 Claws 14
1.12.5 Spur 14
1.12.6 Pads 14
1.12.6.1 Metatarsal Pad 14
1.12.6.2 Digital Pads 14
1.13 Thermoregulation 15
References 15
2 Skeletal System 19
Wael Khamas, Josep Rutllant, and Ken Noriega
2.1 Introduction 19
2.2 Cartilage and Bone Embryonic Origin 19
2.3 Cartilage Cell Types 19
2.3.1 Chondrogenic (Chondroprogenitor) 19
2.3.2 Chondroblast 19
2.3.3 Chondrocyte 19
2.3.4 Chondroclast 19
2.4 Type of Cartilages 20
2.4.1 Hyaline Cartilage 20
2.4.2 Fibrocartilage 20
2.4.3 Elastic Cartilage 20
2.5 Cartilage Development 20
2.6 Calcium Homeostasis 20
2.7 Bone Cell Types 21
2.7.1 Mesenchymal 21
2.7.2 Pericyte 21
2.7.3 Osteogenic 21
2.7.4 Osteoblast 21
2.7.5 Osteocyte 21
2.7.6 Osteoclast 21
2.8 Ossification 22
2.8.1 Bone Formation (Ossification) 22
2.8.2 Intramembranous Ossification 23
2.8.2.1 Flat Bones 23
2.8.3 Endochondral Ossification 23
2.8.3.1 Long Bone 23
2.9 Axial Skeleton 25
2.9.1 Skull 25
2.9.2 Vertebrae 25
2.9.3 Notarium 25
2.9.4 Ribs (Costae) 26
2.9.5 Sternum 26
Contents ix
2.10 esamoid Bones 26
S
2.11 Pneumatic Bones 27
2.12 Pectoral Girdle 27
2.12.1 Scapula 27
2.12.2 Coracoid 27
2.12.3 Clavicle 27
2.12.4 Triosseal Foramen 28
2.13 Pectoral Limb (Wing) 28
2.13.1 Humerus 28
2.13.2 Radius and Ulna 29
2.13.3 Carpometacarpal 29
2.14 Pelvic Girdle 29
2.15 Pelvic Limb 29
2.15.1 Femur 29
2.15.2 Patella 29
2.15.3 Tibiotarsus 30
2.15.4 Fibula 31
2.15.5 Tarsometatarsus 31
2.15.6 Digits 31
2.16 Arthrosis (Joints) 31
2.16.1 Joint Movements 31
2.16.2 List of Joints 32
2.16.2.1 Pectoral Limb Joints 32
2.16.2.2 Pelvic Limb Joints 32
References 32
3 Muscular System 35
Ken Noriega, Thomas Marino, and Wael Khamas
3.1 Introduction 35
3.2 Muscle Histology 35
3.2.1 Smooth Muscle 35
3.2.2 Cardiac Muscle 35
3.2.3 Skeletal Muscle 35
3.3 Muscle Nomenclature 37
3.3.1 Structure and Arrangement of Fibers 37
3.3.2 Number of Heads 37
3.3.3 Action 37
3.3.4 Shape/Morphology 37
3.4 Types of Skeletal Muscle Fibers 37
3.5 Muscle Cells 38
3.5.1 Myocyte 38
3.5.2 Satellite 38
3.6 Skeletal Muscle Blood Supply 38
3.7 Skeletal Muscle Innervation 38
3.8 Muscle Spindle 38
3.9 Tendon 39
3.9.1 Histology 39
3.9.2 Tendon Modifications 39
3.10 General Description of Skeletal Muscle 39
3.11 Cutaneous Muscles 40
3.11.1 Bird Muscles 40
x Contents
3.12 light Muscles 40
F
3.12.1 Pectoral 40
3.12.2 Wing Muscles 41
3.12.2.1 Ventral Wing Muscles 41
3.12.2.2 Dorsal Wing Muscles 41
3.13 Muscles of the Pelvic Region 44
3.13.1 Thigh Muscles 44
3.13.1.1 Iliotibialis Complex 44
3.13.1.2 Iliacus Muscle 45
3.13.1.3 Adductor (Pubischiofemoralis Muscle) 45
3.13.1.4 Semimembranosus Muscle 45
3.13.1.5 Semitendinosus Muscle 45
3.13.1.6 Quadratus Femoris Muscle 45
3.13.1.7 Ambiens (Pectineus) Muscle 45
3.13.2 Leg (Crural) Muscles 46
3.13.2.1 Anterior Tibial 46
3.13.2.2 Gastrocnemius Muscle 46
3.14 Abdominal Muscles 46
3.14.1 External Abdominal Oblique 46
3.14.2 Internal Abdominal Oblique 46
3.14.3 Rectus Abdominal 46
3.14.4 Transverse Abdominal 46
References 46
4 Digestive System 49
Josep Rutllant and Wael Khamas
4.1 Introduction 49
4.2 Oral Cavity and Pharynx 49
4.2.1 Beak (Rhamphotheca) 50
4.2.2 Tongue 52
4.2.3 Salivary Glands 52
4.2.4 Taste Buds 53
4.2.5 Pharyngoesophageal Junction 53
4.3 Hyoid Apparatus (Hyoglossal, Hyolingual, or Hyobranchial) 53
4.4 Esophagus (Pars cervicalis, Crop, and Pars thoracica) 54
4.5 Stomach (Proventriculus and Ventriculus) 56
4.5.1 Proventriculus (Glandular Stomach) 56
4.5.2 Intermediate Zone 57
4.5.3 Gizzard (Muscular Stomach, Ventriculus) 58
4.6 Small Intestine 59
4.6.1 Duodenum 60
4.6.2 Jejunum 61
4.6.3 Ileum 61
4.7 Large Intestine 62
4.7.1 Rectum (Colon, Colorectum) 62
4.7.2 Ceca 63
4.8 Cloaca 63
4.9 Vent 65
4.10 Liver (Hepar, Jecur) 65
4.10.1 Hepatic Portal System and Blood Supply 66
4.11 Pancreas 67
References 68
Contents xi
5 Respiratory System 71
Hrvoje Smodlaka and Wael Khamas
5.1 Introduction 71
5.2 Nostrils and Nasal Cavity 71
5.3 Larynx (Larynx Cranialis), Upper Larynx 73
5.4 Trachea 74
5.5 Syrinx (Larynx Caudalis), Lower Larynx 75
5.6 Lungs 76
5.7 Direction of Airflow 78
5.8 Blood-Gas Barrier 79
5.9 Air Sacs 79
References 80
6 Urinary System 83
Wael Khamas and Josep Rutllant
6.1 Introduction 83
6.1.1 Overall Composition and Function of the Urinary System 83
6.1.2 Development 83
6.2 Kidney 83
6.2.1 Macroscopic and Topographic Anatomy 83
6.2.1.1 Kidney Lobation 84
6.2.1.2 Kidney Lobulation 84
6.2.2 Histology 85
6.2.2.1 Kidney Capsule 85
6.2.2.2 Cortex 85
6.2.2.3 Medulla 86
6.2.2.4 The Nephron 86
6.2.2.5 Juxtaglomerular Apparatus 88
6.2.2.6 Collecting Ducts 89
6.3 Ureter 89
6.4 Urodeum 90
6.5 Blood Supply 90
6.6 Innervation 90
6.7 Urine 91
References 91
7 Reproductive System 93
Josep Rutllant and Wael Khamas
7.1 Male Reproductive System 93
7.1.1 Testes 93
7.1.2 Epididymis 96
7.1.3 Deferent Duct (ductus deferens) 96
7.1.4 Phallus 96
7.1.5 Spermatogenesis 97
7.1.6 Mature Spermatozoa 98
7.2 Female Reproductive System 99
7.2.1 The Ovary 100
7.2.2 The Oviduct 101
7.2.2.1 Infundibulum 102
7.2.2.2 Magnum 102
7.2.2.3 Isthmus 104
7.2.2.4 Uterus (Shell Gland, Eggshell Gland) 104
xii Contents
7.2.2.5 Vagina 104
7.2.3 Cloaca 105
7.2.4 The Formation of a Chicken’s Egg 105
7.2.4.1 Yolk 106
7.2.4.2 Synthesis of Albumen 106
7.2.4.3 Production of Shell Membranes 106
7.2.4.4 Formation of the Testa or Hard Shell 106
References 106
8 Endocrine System 109
Manel Lopez-Bejar
8.1 Introduction 109
8.1.1 Chicken Endocrine System: A Key Regulator of Physiological Processes 109
8.2 Physiological Balance 110
8.2.1 Homeostasis 110
8.3 Major Endocrine Glands 110
8.3.1 Hypothalamus 111
8.3.1.1 Anatomy 111
8.3.1.2 Histology 111
8.3.1.3 Function 111
8.3.2 Pituitary Gland (Hypophysis Cerebri) 112
8.3.2.1 Anatomy 112
8.3.2.2 Histology 112
8.3.2.3 Adenohypophysis Cells 113
8.3.2.4 Neurohypophysis 113
8.3.2.5 Function 113
8.3.3 Adrenal Gland 114
8.3.3.1 Anatomy 114
8.3.3.2 Histology 114
8.3.3.3 Function 115
8.3.4 Thyroid Gland 115
8.3.4.1 Anatomy and Histology 115
8.3.4.2 Function 116
8.3.5 Parathyroid Glands 116
8.3.5.1 Anatomy and Histology 116
8.3.5.2 Function 116
8.3.6 Ultimobranchial Bodies 117
8.3.6.1 Anatomy and Histology 117
8.3.6.2 Function 117
8.3.7 Carotid Body 118
8.3.8 Pineal Gland (Epiphysis Cerebri) 118
8.3.8.1 Anatomy and Histology 118
8.3.8.2 Function 118
8.3.9 Islets of Langerhans 119
8.3.9.1 Anatomy and Histology 119
8.3.9.2 Function 119
8.3.10 Gonads 120
8.3.10.1 Anatomy and Histology 120
8.3.10.2 Function 120
8.3.11 Digestive Enterochromaffin Cells 120
References 121
Contents xiii
9 Sense Organs 125
Wael Khamas and Josep Rutllant
9.1 The Eye 125
9.1.1 Eyeball 125
9.1.1.1 Fibrous Tunic 125
9.1.1.2 Vascular Tunic (Uvea) 126
9.1.1.3 Nervous Tunic (Retina) 128
9.1.2 Lens 128
9.1.3 Eye Chambers 129
9.1.4 Barriers 129
9.1.5 Vitreous Body 129
9.1.6 Adnexa 129
9.1.6.1 Eyelids (Palpebra; Plural Palpebrae) 129
9.1.6.2 Third Eyelid 130
9.1.6.3 Extraocular Muscles 131
9.1.6.4 Lacrimal Apparatus 131
9.1.7 Vision 132
9.2 The Ear 132
9.2.1 Mature Chicken Ear Anatomy 132
9.2.1.1 External Ear 133
9.2.1.2 Middle Ear 133
9.2.1.3 Inner Ear 134
9.2.2 Hearing 135
References 135
10 The Cardiovascular System 139
Thomas Marino and Wael Khamas
10.1 Introduction 139
10.2 Heart 139
10.2.1 Heart Topography 139
10.2.2 Pericardial Sac 139
10.2.3 Heart Surface Anatomy 140
10.2.4 Chambers of the Heart 141
10.2.5 Sinus Venosus 141
10.2.6 Right Atrium 141
10.2.7 Right Ventricle 141
10.2.8 Left Atrium 141
10.2.9 Left Ventricle 142
10.2.10 Atrioventricular Valves 142
10.2.11 Cardiac Skeleton 142
10.2.12 Conducting System 142
10.2.13 Heart Arterial Supply 142
10.2.14 Venous Drainage of the Heart 143
10.3 General Arterial System 143
10.3.1 Pulmonary Trunk 143
10.3.2 Systemic Arteries 143
10.3.2.1 Cervical Region 143
10.3.2.2 Head 143
10.3.2.3 Breast and Thoracic Limb 145
10.3.2.4 Spinal Cord 145
10.3.2.5 Abdominal Viscera 145
xiv Contents
10.3.2.6 Aorta Termination 147
10.4 Venous System 148
10.4.1 Pulmonary Veins 148
10.4.2 Systemic Veins 148
10.4.2.1 Cranial Vena Cavae 148
10.4.2.2 Caudal Vena Cava 148
10.5 Sites for Blood Collection 148
10.6 Hepatic Portal System 149
10.7 Renal Portal System 149
10.8 Carotid Body/Carotid Sinus 149
References 150
11 The Lymphatic System 153
Wael Khamas and Josep Rutllant
11.1 Introduction 153
11.2 Cells of the Lymphatic System 153
11.2.1 Lymphocytes 153
11.2.1.1 Small-Size Lymphocyte 153
11.2.1.2 Medium-Size Lymphocyte 154
11.2.1.3 Large-Size Lymphocyte 154
11.2.1.4 Naive Lymphocyte 154
11.2.1.5 B Lymphocyte 154
11.2.1.6 Plasma Cell 155
11.2.1.7 T Lymphocyte 155
11.2.2 Monocyte 155
11.2.3 Macrophage, Resident, and Wandering 155
11.2.3.1 Dendritic 155
11.2.3.2 Microfold (M) 155
11.2.4 Reticular Cell 155
11.2.5 Epithelial-Reticular 156
11.3 Common Terms Used with the Lymphatic System 157
11.3.1 Capsule 157
11.3.2 Trabecula(e) 157
11.3.3 Cortex 157
11.3.4 Medulla 157
11.3.5 Lymphatic Nodule (Follicle) 157
11.3.6 Primary Lymphatic Follicle 157
11.3.7 Secondary Lymphatic Follicle 157
11.3.7.1 Germinal Center 157
11.3.7.2 Mantle 157
11.3.8 High Endothelial Cell Venules 157
11.4 Primary Lymphatic Organs 157
11.4.1 Bursa of Fabricius (Cloacal Bursa) 157
11.4.2 Thymus 159
11.4.2.1 Blood Thymic Barrier in Chicken 161
11.5 Secondary Lymphatic Organs 161
11.5.1 Spleen 161
11.5.1.1 Spleen Blood Supply 162
11.5.2 Cecal Tonsils 163
11.5.3 Gut-Associated Lymphoid Tissue 164
11.5.3.1 Peyer’s Patches 165
11.5.3.2 Meckel’s Diverticulum 166
Contents xv
11.5.4 Harderian Gland 166
11.5.5 Mucosal Associated Lymphoid Tissue 166
11.5.5.1 Conjunctival Associated Lymphoid Tissue, Conjunctival Lymph Nodules 166
11.5.5.2 Nasal Associated Lymphoid Tissue 166
11.5.5.3 Trachea and Lung Lymphoid Tissue, Mucosal Associated Lymphoid Tissue 166
11.5.5.4 Bronchiole-Associated Lymphoid Tissue 166
11.5.6 Lymph Nodes 168
11.6 Maternal Transfer of Immunity 168
11.7 Lymphatic Vessels 168
References 168
12 Nervous System 171
Wael Khamas, Josep Rutllant, and Hrvoje Smodlaka
12.1 Introduction 171
12.2 Neuron 171
12.2.1 Type of Neurons 171
12.2.1.1 Unipolar Neuron 171
12.2.1.2 Bipolar Neuron 171
12.2.1.3 Multipolar Neuron 172
12.2.2 Synapse 172
12.2.2.1 Types of Synapses 172
12.3 Ganglion 172
12.3.1 Sensory 172
12.3.2 Autonomic 172
12.4 Glial Cells (Neuroglia) 172
12.4.1 Microglial Cell (Microgliocyte) 172
12.4.2 Macroglial Cells 173
12.4.2.1 Astrocyte 173
12.4.2.2 Oligodendrocyte 174
12.4.3 Ependymal Cell 174
12.5 Peripheral Nervous System Cells 174
12.5.1 Schwann Cell (Lemmocyte, Neurolemmocyte) 174
12.5.2 Satellite Cell 174
12.6 Meninges 174
12.7 Brain Ventricles 175
12.8 Cerebrospinal Fluid (CSF) 175
12.8.1 Choroid Plexus 175
12.8.2 CSF Flow Pattern 175
12.9 Blood Brain Barrier 175
12.10 Cerebrum 176
12.10.1 Telencephalon 176
12.10.1.1 Olfactory Bulbs 176
12.10.2 Diencephalon 177
12.10.2.1 Thalamus 177
12.10.2.2 Epithalamus 177
12.10.2.3 Hypothalamus 177
12.10.3 Mesencephalon 177
12.10.3.1 Optic Tectum 177
12.10.4 Metencephalon (Cerebellum) 177
12.10.5 Myelencephalon (Medulla Oblongata) 178
12.11 Reticular Formation 179
12.12 Spinal Cord 179
xvi Contents
12.13 lycogen Bodies 180
G
12.14 Autonomic Nervous System 180
12.14.1 Sympathetic 181
12.14.2 Parasympathetic 181
References 181
13 Applied Chicken Anatomy 183
Teresa Y. Morishita and Robert E. Porter Jr
13.1 Examining from a Distance 183
13.1.1 Daily Behavioral Deviations 183
13.1.2 Abnormal Mentation 183
13.1.3 Abnormal Posture 183
13.2 Physical Examination 185
13.2.1 Feathers 185
13.2.2 Head Region 186
13.2.3 Wattles 187
13.2.4 Eyes 187
13.2.5 External Auditory Canal (Ears) 188
13.2.6 Thorax 188
13.2.7 Cloaca 188
13.2.8 Legs 189
13.3 Specialized Clinical Procedures 190
13.3.1 Identification Placement 190
13.3.2 Culture Techniques: Choana, Oropharynx, and Trachea 191
13.3.3 Blood Collection 191
References 192
14 Chicken Necropsy 193
Miguel D. Saggese
14.1 Introduction 193
14.2 Euthanasia and Carcass Disposal 193
14.3 Chicken Disposal 194
14.4 Facilities and Materials Needed 194
14.4.1 Personal Protective Equipment 194
14.4.2 Histopathological Examination 194
14.4.3 Collection of Microbiological Samples for Culture and Molecular Diagnostics 195
14.4.4 Molecular Diagnosis of Microorganisms 196
14.4.5 Toxicological Investigation 196
14.5 Whole Carcass Conservation 196
14.6 Necropsy Protocol 196
14.6.1 First Thing First 196
14.6.2 External Examination 197
14.6.3 Internal Examination 199
14.6.3.1 Subcutaneous and Muscles 199
14.6.3.2 Celomic Cavities 200
14.6.3.3 Heart 201
14.6.3.4 Liver and Gall Bladder 202
14.6.3.5 Air Sacs 203
14.6.3.6 Thyroid and Parathyroid Glands 204
14.6.3.7 Proventriculus 204
14.6.3.8 Lungs 204
14.6.3.9 Spleen 204
Contents xvii
14.6.3.10 Gastrointestinal Tract 205
14.6.3.11 Pancreas 206
14.6.3.12 Gonads and Associated Organs 206
14.6.3.13 Kidneys and Adrenal Glands 207
14.6.3.14 Bursa of Fabricius 207
14.6.4 Bones and Bone Marrow 207
14.6.5 Uropygial Gland 207
14.6.6 Skull 207
14.6.7 Vertebral Column and Spinal Cord 207
Further Reading 208
15 The Egg Anatomy 209
Josep Rutllant and Wael Khamas
15.1 Introduction 209
15.2 The Germinal Disc 209
15.3 The Yolk 209
15.4 The Albumen 211
15.5 The Eggshell 212
15.5.1 Shell Membranes 212
15.5.2 Testa 212
15.5.2.1 Inorganic Portion 212
15.5.2.2 Organic Portion 213
15.5.3 Cuticle 214
References 214
16 Fertilization and Chick Embryo Development 217
Josep Rutllant and Wael Khamas
16.1 Introduction 217
16.2 Avian Fertilization 217
16.2.1 Sperm Deposition, Storage, and Transport 217
16.2.2 Ovulation (Oviposition) 219
16.2.3 Fertilization 219
16.3 Chick Embryo Development 220
References 224
Index 227
