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==Preface== | ==Preface== | ||
More than half a century ago Wilhelm Preyer published a monograph «specielle Physiologie des Embryo« embodying not only a review of the literature of that day, but a great many new observations of his own as well. This boolc has occupied a rather unique Position in that it was long the only source of summarized lcnowledge concerning the activities of embryos and fetuses of many species. That no other book has quite talcen its place is not surprising because, after initial exploration of the Held, physiologists turned their attention to more urgent problems concerning the adult. Within the last decade or two, interest has revived and a school of developmental physiology has come into being: Many of the subjects discussed by Preyer have been restudied prolitably by experimental methods not available in his time, and new observations have added facts of great signiiicance to our conception of life before birth. Few biologists, however, are aware of all that has been accomplished during the last few years. | |||
My main purpose in writing the present boolc was to assemble and summarize scattered physiologic observations on fetuses for my own information and for that of my students. I hope that the result will be useful to others who are working in fetal physiology and perhaps will help direct attention to problems which«need to be investigated Another purpose was to provide a supplement for courses in embryo1ogy to help stress functional aspects of de— velopment; this is in line with current trends of medical teaching. Finally, I had in mind those in allied lields, especially neurology, psychology and pediatrics, who are interested in problems of behavior and who have frequent occasion to desire lcnowledge of prenatal physiology. | |||
Originally I thought to produce a more comprehensive review somewhat similar to that of Preyer, but my first excursions into Helds with which I had been only slightly familiar before demonstrated the futility of doing so within a single small volume. some of the purposes of the taslc would have been defeated by undertaking lengthy detailed discussions Furthermore, a great deal has been written on subjects with· which I feel incompetent to deal critically. This is particularly true of nutrition and metabolism. To do justice to these subjects would have meant- duplication of much that is contained in Joseph Needham’s splendid «Chemical Embryology." Therefore I determined to place emphasis upon the more strictly physiologic aspects of prenatal life and to enter into the chemistry of the fetus only to the point of supplying the reader with brief up-to-date synopses for the salce of completeness. | |||
and | |||
It is clearly realized that the present boolc is incomplete in respect to many subjects which might have been included had arbitrary limits not been set down. It was in many instances diilicult | It is clearly realized that the present boolc is incomplete in respect to many subjects which might have been included had arbitrary limits not been set down. It was in many instances diilicult to draw the line between physiology and anatomy. Determinate growth in the early stages of development was of necessity ignored entirely, and many very interesting observations in experimental embryology of lower vertebrates (indeed, most of the studies in inframammalian species) ,have been purposely omitted or mentioned only brieHy. As much as possible, therefore, I have tried to limit consideration to mammals, and especially to the higher mammals, but I have brought in observations on other animals when these seem to add siguilicantly to our knowledge of human physiology. | ||
to draw the line between physiology and anatomy. Determinate | |||
growth in the early stages of development was of necessity ignored | |||
entirely, and many very interesting observations in experimental | |||
embryology of lower vertebrates (indeed, most of the studies in | |||
inframammalian species) ,have been purposely omitted or mentioned only brieHy. As much as possible, therefore, I have tried | |||
to limit consideration to mammals, and especially to the higher | |||
mammals, but I have brought in observations on other animals | |||
when these seem to add siguilicantly to our knowledge of human | |||
physiology. | |||
Aclcnowledgments are due a number of individuals and groups | Bibliographic references are placed at the end of each chaptetn Even though this involves some repetition, it was thought to be a more useful plan than to collect them at the end of the book. Citations have been used freely, but no attempt has been made to include all references on any subject: In most instances, pre— liminary articles and many of the older papers discussed by Preyer have been left out and only the more recent lcey references included. | ||
who have contributed directly and indirectly to my program of | |||
investigation in fetal physiology and to the present book. In the | |||
Hrst place, I am grateful to my teachers and colleagues, especially | Aclcnowledgments are due a number of individuals and groups who have contributed directly and indirectly to my program of investigation in fetal physiology and to the present book. In the Hrst place, I am grateful to my teachers and colleagues, especially Professors L. B. Arey, s. W. Ranson and A. C. Ivy, for helpful criticisms and suggestions My graduate students and associates in research during the past ten years have contributed notably to the project and have provided the incentive to proceed with it. To sir Joseph Baker-oft, in whose laboratory I was a guest during the winter ok 1·935-36, belongs credit kor lcindling my interest in respiratory physiologzn Finally, to the National Research council, the Ella sachs Plotz Foundation, The American Academy ok Arts and sciences. The Chijd Neurology Research Council (Fried— sham Foundation) , and The J0hn and Mary R. Marlcle Foundation, all ok whom have generously supplied kunds to aid my investigations, I wish to extend thanks. | ||
Professors L. B. Arey, s. W. Ranson and A. C. Ivy, for helpful | |||
criticisms and suggestions My graduate students and associates | |||
in research during the past ten years have contributed notably to | |||
the project and have provided the incentive to proceed with it. | |||
To sir Joseph Baker-oft, in whose laboratory I was a guest during the winter ok 1·935-36, belongs credit kor lcindling my interest in | |||
respiratory physiologzn Finally, to the National Research council, the Ella sachs Plotz Foundation, The American Academy ok | |||
Arts and sciences. The Chijd Neurology Research Council (Fried— | |||
sham Foundation) , and The J0hn and Mary R. Marlcle Foundation, all ok whom have generously supplied kunds to aid my investigations, I wish to extend thanks. | |||
W. F. Wissens. | W. F. Wissens. | ||
==Contents== | ==Contents== |
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Windle WF. Physiology of the Fetus. (1940) Saunders, Philadelphia.
1940 Physiology of the Fetus: 1 Introduction | 2 Heart | 3 Circulation | 4 Blood | 5 Respiration | 6 Respiratory Movements | 7 Digestive | 8 Renal - Skin | 9 Muscles | 10 Neural Genesis | 11 Neural Activity | 12 Motor Reactions and Reflexes | 13 Senses | 14 Endocrine | 15 Nutrition and Metabolism | Figures
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This 1940 textbook by Prof William Frederick Windle (1898-1986) describes human fetal physiology as it was understood at that time. His research collection consists of paper documents, plus approximately 25,000 feet of motion pictures, over 400 slides, photographs, and a few artifacts. The materials span the years 1918-1986. (More? See Online Archives of California)
Also by this author: Windle WF. (1970). Development of neural elements in human embryos of four to seven weeks gestation. Exp. Neurol. , 28, Suppl:44-83. PMID: 4097652
|
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Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding. (More? Embryology History | Historic Embryology Papers) |
Physiology of the Fetus
Origin and Extent of Function in Prenatal Life
By
William Frederick Windle
Professor of Microscopic Anatomy, Northwestern University Medical School
W.B. Saunders Company Philadelphia and London, 1940
Preface
More than half a century ago Wilhelm Preyer published a monograph «specielle Physiologie des Embryo« embodying not only a review of the literature of that day, but a great many new observations of his own as well. This boolc has occupied a rather unique Position in that it was long the only source of summarized lcnowledge concerning the activities of embryos and fetuses of many species. That no other book has quite talcen its place is not surprising because, after initial exploration of the Held, physiologists turned their attention to more urgent problems concerning the adult. Within the last decade or two, interest has revived and a school of developmental physiology has come into being: Many of the subjects discussed by Preyer have been restudied prolitably by experimental methods not available in his time, and new observations have added facts of great signiiicance to our conception of life before birth. Few biologists, however, are aware of all that has been accomplished during the last few years.
My main purpose in writing the present boolc was to assemble and summarize scattered physiologic observations on fetuses for my own information and for that of my students. I hope that the result will be useful to others who are working in fetal physiology and perhaps will help direct attention to problems which«need to be investigated Another purpose was to provide a supplement for courses in embryo1ogy to help stress functional aspects of de— velopment; this is in line with current trends of medical teaching. Finally, I had in mind those in allied lields, especially neurology, psychology and pediatrics, who are interested in problems of behavior and who have frequent occasion to desire lcnowledge of prenatal physiology.
Originally I thought to produce a more comprehensive review somewhat similar to that of Preyer, but my first excursions into Helds with which I had been only slightly familiar before demonstrated the futility of doing so within a single small volume. some of the purposes of the taslc would have been defeated by undertaking lengthy detailed discussions Furthermore, a great deal has been written on subjects with· which I feel incompetent to deal critically. This is particularly true of nutrition and metabolism. To do justice to these subjects would have meant- duplication of much that is contained in Joseph Needham’s splendid «Chemical Embryology." Therefore I determined to place emphasis upon the more strictly physiologic aspects of prenatal life and to enter into the chemistry of the fetus only to the point of supplying the reader with brief up-to-date synopses for the salce of completeness.
It is clearly realized that the present boolc is incomplete in respect to many subjects which might have been included had arbitrary limits not been set down. It was in many instances diilicult to draw the line between physiology and anatomy. Determinate growth in the early stages of development was of necessity ignored entirely, and many very interesting observations in experimental embryology of lower vertebrates (indeed, most of the studies in inframammalian species) ,have been purposely omitted or mentioned only brieHy. As much as possible, therefore, I have tried to limit consideration to mammals, and especially to the higher mammals, but I have brought in observations on other animals when these seem to add siguilicantly to our knowledge of human physiology.
Bibliographic references are placed at the end of each chaptetn Even though this involves some repetition, it was thought to be a more useful plan than to collect them at the end of the book. Citations have been used freely, but no attempt has been made to include all references on any subject: In most instances, pre— liminary articles and many of the older papers discussed by Preyer have been left out and only the more recent lcey references included.
Aclcnowledgments are due a number of individuals and groups who have contributed directly and indirectly to my program of investigation in fetal physiology and to the present book. In the Hrst place, I am grateful to my teachers and colleagues, especially Professors L. B. Arey, s. W. Ranson and A. C. Ivy, for helpful criticisms and suggestions My graduate students and associates in research during the past ten years have contributed notably to the project and have provided the incentive to proceed with it. To sir Joseph Baker-oft, in whose laboratory I was a guest during the winter ok 1·935-36, belongs credit kor lcindling my interest in respiratory physiologzn Finally, to the National Research council, the Ella sachs Plotz Foundation, The American Academy ok Arts and sciences. The Chijd Neurology Research Council (Fried— sham Foundation) , and The J0hn and Mary R. Marlcle Foundation, all ok whom have generously supplied kunds to aid my investigations, I wish to extend thanks.
W. F. Wissens.
Contents
CHAPTTER I
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I
Relation ok Fetal to Maternal Organism . . . . . . . . . . . . . 2 Experimental Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Rekerences Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. to
CHAPTER II
THE FETAL HEART . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1 Initiation of the Heart Beat . . . . . . . . . . . . . . . . . . . . . . . . 1 1 The Fetal Electrocardiogram . . . . . . . . . . . . . . . . . . . . . . . 13 The Fetal Pulse Rate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . « 15 Nervous control of the Fetal Heart and circulation . . . 18 Arteria1 B1ood Pressure . . . . . . . . . . . . . . . . . . . . . . . . . .. 22 Venous Blood Pressure . . . . . . . . . . . . . . . . . . . . . . . . . . . · 25 References Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 26
CHAPTER III
THE FETAL cmCULATIoN . . . . . . . . . . . . . . . . . . . . . . .. 29
Volume of B1ood and Rate of Circulation . . . . . . . . . . . . 29
The course ok the Fetal B1ood .- . . . . . . . . . . . . . . . . . . . 32
changes in the circulation at Birth . . . . . . . . . . . . . . . . . 44
Rekerences Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 47
oHAPTER 1v THE BLOOD OF THE FETUs . . . . . . . . . . . . . . . . . . . . . . . . . . 49
The Red B1ood Corkzuscles . . . . . . . . . . . . . . . . . . . . . . . . 49
Oxygen Carrying Power ok Fetal B1ood . . . . . . . . . . . . . . « 53
The Leulcocytes and Platelets . . . . . . . . . . . . . . . . . . . . . . 55
The Plasma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .- . . . . . . 56
The cerebrospinal Fluid . . . . . . . . . . . . . . . . . . . . . . . . .. 56
Rekerences Cited . . . . . . . . . . . . .- . . . . . . . . . . . . . . . . . . .. 57
CHAPTER V Page
FETAL RESPIRATION . . . . . : . . . . . . . . . . . . . . . . . . . . . . . . . . 59 The Mechanism of Gaseous Exchange . . . . . . . . . . . . . . . 59 The Oxygen capacity ok the Blood . . . . . . . . . . . . . . . . . 61 Dissociation Curves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 Oxygen and Carbon Dioxide contents ok Fetal Blood . . 69 Asphyxia at Birth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 75 Rekerences Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 76
» CHAPTER vI
FETAL REsPIRAToRY MovEMENTs . . . . . . . . . . . . . . . . .. 78 Respiratory Movements in the Ititact Animal . . . . . . . . . 78
Respiratory Movernents under Experimental Conditions 8o Relation ok Oxygen and Carbon Dioxide to Respiratory
Movements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Other Factors in the Development ok Respiration .. .. 88 Aspiration ok Amniotic contents . . . . . . . . . . . . . . · . . .. 91 Fetal Hiccup . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 94 summary . . . . . . . . . . . . . . . . . . . . . . .« . . . . . . . . . . . . . . . . . 95 References cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 96
CHAPTER VII
THE FETAL D1GEsT1VE sYsTEM . . . . . . . . . . . . . . . . . . . . .. 99
Fetal swallowing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 99
Fetal Gastric Motility . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Fetal Intestinal Movements . . . . . . . . . . . . . . . . . . . . . . , . 103
Absorption in the Fetal Digestive Tract . . . . . . . . . . . ». 106
Defecation and Meconiophagy in Amnio . . . . » . . . . . . . . . 1o7
The Fetal Digestive Glands Enzymes . . . . . : . . . . . . . . . 108
References Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 11o
CHAPTER VIII
THE FETAL KIDNEY AND FLUIDS THE FETAL SKIN . . 1 12
Development ok Kidney Function . . . . . . . . . . . . . . . . .. 112 Physiologic Development of the Nephric Tubules . . . . . 1 14 Conditions Regulating Renal F unction . . . . . . . . . . . . . . 1 17 Changes in Elimination at. Birth . . . . . . . . . . . . . . . . . . . 119
The Fetal Urine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 120 The Allantoic F1uid . . . . . . .- . . . . . . . . . . . . . . . . . . . . . .. 121 The Amniotic Fluid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 The Fetal slcin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124 lseferences Cited . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . 126
CHAPTER 1X
THE FETAL MUsCLEs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
Tissue Cultures and In-vitro Experiments . . . . . . . . . .. 128
spontaneous Activity of Intact slceletal Musc1e . . . . . .. 130
Faradic and Mechanical stimu1ation . . . . . . . . . . . . . . .. 131
The Fetal Tetanic Reaction . . . . . . . . . . . . . . . . . . . . . .. 133 Excitation of Fetal Musc1e by Nerve stimülation . . . . . 135 Efkects of curare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 135 Feta1 Rigor Mortis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136 References Cited , . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
CHAPTER X
THE. cENEsls OF FUNCTION IN THE NERVOUS sYsTEM 138
Myogenic Responses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
N euromotor Responses . . . . . . . . . . . . . . . . . . . . . . . . . . .. 14o
Genesis of ReAex Behavior . . . . . . . . . . . . . . . . . . . . . . . . . 141
The concept of a Total Pattern . . . . . . . . . . . . . . . . . . . . 142
EarIy Reflexes in Mammalian Embryos . . . . . . . . . . . . . . 147
0ther sirnple Reilexes and Their Integration . . . . . . . . 153
References Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 156
CHAPTER xl
CONDIT10Ns REGULATING FETAL NERVOUS ACT1VITY 158
The Plan of structural Development of the Fetal Brain 158 (a) Order of Development in Functional Systems 16o (l)) Growth of 0ther circuits for Reliexes and.
Higher Integration . . . . . . . . . . . . . . . . . . . ·. 162
(c·) Myelogeny as Related to Function in the Ner— « vous System . . . . . . . . . . . . . . . . . . . . . . . . . . .. 163 Factors Other than structural Growth . . . . . . . . . . . . . . 164 (a) The Quiescence of Intra-uterine Life . . . . . . . 164
(b) Afkerent stirnulation in«Utero . . . . . . . . . . . .. 165 xä coNTENTs
. Fast? (c) Neura1 Thresholds to stimulation . . . . . . . . .. 166 (d) Musc1e Tonus and Mass Movements . . . . . . . . 168 (e) susceptibility Gradients to Asphyxia . . . . . . . . 169 (j) Inhibition ok Motor Mechanisms by Higher centers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . «« 170 References cited . . . . . . . . . . . . . . « . . . . . . . . . . . . . . . . . . 171
CHAPTER Xll
FETAL MOTOR REACTIONS AND REFLEXES . . . . . . . . . . 173
Development of Feeding Reactions . . . . . . . . . . . . . . . . . 173
Development ok Posture and Progression . . . . . . . . . . . . 175
Development of Eye ReHexes . . . . . . . . . . . . . . . . . . . . . . 1.79
Development of Palmar and Plantar Reflexes . . . . . . . . 180
Other Reiiexes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Rekerences Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 181
CHAPTER X1II
THE FETAL sENsEs . . . . . . . . . . . . . . . ., . . . . . . . . . . . . . . .. 183
The Fetal slcin as a Receptor Organ . . . .« . . . . . . . - . . . . . 184
(a) Pressure, Touch and Pain . . . . . . . . . . . . . . . . . 184
(b) Temperature sensitivity . . . . . . . . . . . . . . . . .. 186
Proprioceptive Function in the Fetus . . . . . . . . . . . . . . . . 186
0lkactory, Gustatory and Visceral senses . . . . . . . . . . .. 188
Hearing and Vision . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
References Cited » . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 189
CHAPTER XIV THE FETAL ENDOCRINE GLANDS . . . . . . . . . . . . . . . . . . . . 191
The suprarenal Cortex . . . . . . . . . . . . . . . . . . . . . . . .. 191
The suprarenal Medulla . . . . . . . . . . . . . . . . . . . . . . . . .. 194
The sex Horrnones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 195
The Thyroid Gland . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 197
The Parathyroid Glands . . . . . . . . . . . . . . . . . . . . . . . . .. 198
The Thymus . . - . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
The Hypophysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 199
secretin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 200
The, Endocrine Pancreas , ». . . . . . . . . . . . . . . . . . . . . . . .. 200
References Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202
FETAL NUTRITION AND METABcLlsM . . . . . . . . . . . . . . 206
Paraplacental Nutrition . . . . . . . . . . . . . . . . . . . . . . . . . .. 206
Placental Permeability . . . . . . . . . . . . . . . . . . . . . . . . . . .. 207
Metabolism ok Carbohyclrates . . . . . . . . . . . . . . . . . . . . . . 2 Io
Metabolism ok Lipids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212
Metabolism of Protein . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
Inorganic Metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . .. 219
Energy Metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 222
Rekerences Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 226 PHYSIOLOGY OF THE FETUS
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