Development of the Avian Embryo: A Behavioural and Physiological Study

378 Pages·1974·13.679 MB·English

by  B. M. Freeman, Margaret A. Vince (auth.)

Development of the Avian Embryo
Development of the
Avian Embryo
A Behavioural and Physiological Study
B. M. FREEMAN
Houghton Poultry Research Station,
Houghton, Huntingdon
and
Margaret A. VINCE
Psychological Laboratory,
Downing Street, Cambridge
LONDON
CHAPMAN AND HALL
First published 1974
by Chapman and Hall Ltd
11 New Fetter Lane, London EC4P 4EE
© 1974 B. M. Freeman and Margaret A. Vince
Soj'tcover reprint of the hardcover 1st edition 1974
All rights reserved. No part of this book
may be reprinted, or reproduced or utilized in
any form or by any electronic, mechanical or
other means, now known or hereafter invented,
including photocopying and recording, or in
any information storage and retrieval system,
without permission in writing from the
Publisher.
Distributed in the U.S.A.
by Halsted Press, a Division
of John Wiley & Son, Inc., New York
Library of Congress Catalog Card Number 74-4340
ISBN -13: 978-94-009-5712-1 e-ISBN-13:978-94-009-571 0-7
DOT: 10.1007/978-94-009-5710-7
Contents
Preface page xiii
Acknowledgements xiv
List of Generic and Specific Names xv
PART ONE: BEHAVIOUR by Margaret A. Vince
1. Incubation Requirements 3
1.1 Introduction 3
1.2 Requirements of the embryo of the domestic fowl 4
Heat - Humidity - The gaseous environment - Egg position
and change of position
1.3 Incubation requirements of other domestic species 5
1.4 Incubation in wild birds 6
Introduction - Incubation sites - Warming of the eggs -
The problem of cooling - The continuity of attentiveness -
Humidity - The gaseous environment - Egg shifting or
turning - Hatching success
2. General Development, Postural Changes, Activity and
Relationship between the Embryo and Other Structures
within the Shell 20
2.1 Changes in size, posture and position in the domestic fowl
embryo during incubation 20
2.2 Embryonic activity during incubation in the domestic fowl 25
embryo
Passive movements - Active, spontaneous movements -
Significance of spontaneous activity
2.3 Hatching and emergence in the fowl 30
2.4 Structural and behavioural development in species other
than the fowl 32
Structural and behavioural development during the first two
thirds of incubation - Pre-hatching and hatching behaviour in
species other than the fowl
v
CONTENTS
2.5 The characterization and scoring of embryonic movements 36
Terms used to describe embryonic activity - The quantifica-
tion of embryonic motility
3. Vocalization and Communication in the Natural
Situation 38
3.1 Species which can vocalize before hatching 38
3.2 Prerequisites for the onset of vocalization 38
3.3 Types of stimulation followed by embryonic vocalization 39
3.4 Effects of embryonic vocalization on the parent 40
3.5 Long-term effects of embryo/parent interaction 40
3.6 The earliest stage at which embryos become responsive to
parental calls 42
4. Effects of External Stimulation on Embryonic Activity,
Rate of Development and Time of Hatching 43
4.1 Introduction 43
4.2 The synchronization of hatching in quail 43
Effects of stimulation on the time of hatching - Problems
arising from changes in hatching time - Signals which have
accelerating or retarding effects on neighbouring eggs - Effects
ofa ccelerating and retarding signals on the rate of development
4.3 Effects of accelerating and retarding stimulation in
domesticated species 55
4.4 Effects of light in the fowl 56
4.5 Embryonic interactions in wild species 57
4.6 The embryonic response to stimulation 59
5. The Nervous System 62
5.1 Introduction 62
5.2 Development of activity in different parts of the nervous
system 62
Spontaneous activity - Responses of the brain to stimulation
5.3 Neural mechanisms mediating early motility in the embryo
of the domestic fowl 66
The development of motor and sensory areas in the spinal cord
-The part played by the brain in controlling periodic, random
motility - Independence of periodic, random motility from
sensory input - Areas of the brain which influence periodic
random motility
vi
CONTENTS
5.4 Neural mechanisms mediating pre-hatching and hatching
behaviour 71
Introduction - The spinal cord and pre-hatching behaviour
- The midbrain and pre-hatching behaviour - The fore-
brain and hatching - The cerebral EEG during hatching
5.5 Neural correlates of embryonic motility 73
Nervous activity in the spinal cord between 15 and 19 days-
The polyneuronal burst discharge between days 5 and 20 -
Neural correlates of motility - Communication between
different parts of the spinal cord
5.6 Inhibition 78
5.7 Sleep and arousal 80
6. The Development of Sensory Systems 84
6.1 Introduction 84
6.2 The non-visual response to light 85
Introduction - Evidence for a non-visual response to light in
newly hatched pigeons - Evidence for a non-visual response
to light in pigeon embryos
6.3 Tactile sensitivity 86
The onset of sensitivity - The type of movement evoked by
tactile stimulation - Response decrements with repeated
stimulation
6.4 Vestibular sensitivity 89
6.5 Proprioception 90
6.6 Hearing 91
Development of the auditory system - The age of onset of
auditory sensitivity - The development of auditory sensitivity
- The ontogeny of auditory discrimination
6.7 Taste 96
The sense of taste in birds - Development of a taste sense in
the embryo
6.8 Vision 98
Development of the visual system - Response to light in the
domestic fowl - Colour vision - The behavioural response
to light - Effects of previous experience on the responsiveness
of the visual system - A comparative view of the onset of
visual function - Effects of light on the embryo in ovo
6.9 Olfaction 102
The olfactory sense in birds - Development of olfaction in
the embryo
vii
CONTENTS
7. Conditioning of the Chick Embryo and Conclusions to 104
Chapters 1-7
8. The Newly Hatched Bird 108
8.1 Introduction 108
8.2 Requirements of the neonate 109
Warming and cooling - Righting and standing - Social
attachments in young birds - Feeding behaviour - Oiling and
preening
8.3 What is new in the life of the neonate? 113
8.4 Effects of new environmental stimuli on neural
mechanisms 114
8.5 Conclusions 116
PART TWO: PHYSIOLOGY by B. M. Freeman
9. Gaseous Exchange and Oxygenation of the Embryo 119
9.1 Gross oxygen consumption and carbon dioxide
production 119
9.2 The shell and the chorio-allantois 123
Structure of the shell - Development and structure of the
chorio-allantois
9.3 Physical aspects of gaseous exchange 130
Oxygen - Carbon dioxide - The air space - Maximum
gaseous flux during incubation
9.4 The blood and vascular system 138
Red blood cells - Blood volume and pressure - The heart
Haemoglobins - Oxygen affinity of haemoglobins - Blood
gases
9.5 Efficiency of the chorio-allantois as a surface for gaseous
exchange 154
9.6 Effects of altitude 156
10. Nutrition and Utilization of Albumen and Yolk 160
10.1 Nutrition of the developing bird 160
Essential amino acids - Carbohydrate - Essential fatty acids
- Minerals, trace elements and vitamins - Energy sources
10.2 Gross composition of the egg and the changes during
incubation 163
10.3 The albumen and its utilization 164
Composition - Absorption - Functions
viii
CONTENTS
10.4 The yolk and its utilization 168
Composition - The yolk sac membrane - Utilization of yolk
10.5 Energy metabolism 183
10.6 The sub-embryonic fluid 184
11. Acid-base Balance 186
11.1 Blood pH and P 186
C02
11.2 Plasma bicarbonate 186
11.3 The compensation of respiratory acidosis 187
The sources of bicarbonate - The possible role of the
kidney in excreting bicarbonate
12. Excretion and Water Balance 191
12.1 Introduction 191
12.2 The development of the excretory system 191
The pronephros - The mesonephros - The metanephros
12.3 Synthesis of nitrogenous excretory products 194
Metabolism of the amino group - Urea - Uric acid
12.4 The pattern of excretion during development 198
Excretory products - The organs of excretion
12.5 The allantoic sac 200
Storage of excretory products - Active transport by the
allantoic endoderm
12.6 Water balance 204
Sources of water - Factors affecting water loss - Gross
movements of water within the egg - Conservation of water
13. Hormones in Development 208
13.1 General introduction 208
13.2 Thyroid hormones 208
Development of the thyroid glands - Synthesis and secretion
of hormones - Metabolic effects of the thyroid hormones-
The thyroid and hatching
13.3 Parathyroid hormone and calcitonin 213
Introduction - Development oft he parathyroids and ultimo
branchial bodies - Secretion of parathormone and calcitonin
- Significance in development
13.4 Adrenal hormones 216
Development of the adrenal glands - Synthesis and secretion
of adrenal hormones - Significance of adrenal cortical hor
mones in development - Significance of adrenal medullary
hormones in development
*
~
CONTENTS
13.5 Pancreatic hormones 223
Introduction - Development of the endocrine pancreas -
Structure and secretion of insulin and glucagon - Role of
insulin in development - Role of glucagon in development
13.6 Gonadal hormones 226
Development of the genital system - Synthesis and secretion
of gonadal hormones - Significance in development
13.7 Hypophyseal hormones 232
Introduction - Development of the hypophysis - Secretion
of hormones by the pars distalis - Significance of hypo-
physeal hormones in development
13.8 Glands of uncertain endocrine status 235
Introduction - Bursa of Fabricius - The pineal gland
14. Mobilization and Utilization of Calcium Stores 237
14.1 Physiological aspects of calcium transport 237
14.2 The sources of calcium and gross changes during
incubation 237
14.3 The role of the chorio-allantois in calcium transport 239
Solubilization of calcium - Active transport by the chorionic
ectoderm
14.4 Changes following calcium resorption 246
The shell - Plasma calcium
14.5 Mineralization of the bones 247
15. Physiology of Hatching 249
15.1 General introduction 249
15.2 Pulmonary respiration 249
Introduction - The respiratory system - The onset of
breathing: the pulmonary stimulus - The development of
the breathing pattern
15.3 Circulatory and associated changes 254
15.4 Pipping 256
15.5 Active hatching 257
15.6 Withdrawal and fate of the yolk sac 258
15.7 Oxygenation and energy metabolism during hatching 258
16. The Neonate 261
16.1 Introduction 261
16.2 Thermoregulation 261
Ontogeny of thermoregulation - Mechanisms of thermo
regulation
x
CONTENTS
16.3 The alimentary tract 263
16.4 Immunological competence 264
16.5 Concluding remarks 265
Appendix 1: Chronology of development in the domestic fowl 269
Appendix 2: Development of the chick embryo in relation to the
shell, yolk, albumen and extra-embryonic membranes
by Beryl Tolhurst 277
References 295
Index 351
xi