THE INSECTS Structure and Function R. F. CHAPMAN Assistant Director and Chief of Research, Centre for Overseas Pest Research, London and Visiting Professor, Department of Zoology, University of Hull FACHBEREICH 8IOI.QGIE (10) der Techniscbe" Modischiii D - 6100 D a r m s t a d t / 8. R. D. o HODDER AND STOUGHTON LONDON SYDNEY AUCKLAND TORONTO CONTENTS Preface v SECTION A THE HEAD, INGESTION AND UTILISATION OF THE FOOD I. The Head and its Appendages . . . . . . 1.1 Head 1.2 Neck 1.3 Antennae 1.4 Mouthparts II. Feeding . . . . . . . . 2.1 Feeding habits 2.2 Sense organs associated with feeding 2.3 Finding and recognising the food 2.4 Significance of food preferences 2.5 Conditioning to food 2.6 Feeding and ingestion 2.7 Control of feeding 2.8 The timing of feeding activity and amount eaten 2.9 Fungus-growing insects 2.10 Food storage 2.11 Social feeding III. The Alimentary Canal . . 3.1 General structure 3.2 Foregut 3.3 Midgut 3.4 Hindgut 3.5 Innervation of the gut 3.6 Passage of food through the gut 3.7 Head glands IV. Digestion and Absorption 4.1 Digestion 4.2 Absorption 4.3 Efficiency of food utilisation -.- . . ix . . . . . . . . . . 3 . 22 . . 4 6 . 66 X CONTENTS V. Nutrition 5.1 Dietary requirements 5.2 Storage 5.3 Micro-organisms 84 VI. The Fat Body and General Metabolism 6.1 Fat body 6.2 Intermediate metabolism 6.3 Respiratory metabolism 6.4 End products of catabolism 6.5 Metabolic rate 6.6 Control of metabolism 6.7 Luminescence . . . . . VII. Colour 7.1 The nature of colour 7.2 Physical colours 7.3 Pigmentary colours 7.4 The colours of insects 7.5 Colour change 7.6 Significance of colour VIII. 8.1 8.2 8.3 127 SECTION B THE THORAX AND MOVEMENT The Thorax and Legs Segmentation Thorax Legs IX. Locomotion . . . . 9.1 Walking 9.2 Jumping 9.3 Crawling 9.4 Movements on the surface of water 9.5 Movement under water X. The 10.1 10.2 10.3 10.4 10.5 10.6 100 . . . . 153 . . Wings Occurrence and structure of wings Modifications of the wings Wing coupling Articulation of the wings with the thorax Sensilla on the wings and the haltere Muscles associated with the wings XI. Movement and Control of the Wings 11.1 Mechanisms of wing movement 11.2 Movements of the wings 11.3 Aerodynamics 169 197 . . . . . 216 CONTENTS 11.4 11.5 11.6 Xi Control of wingbeat Stability in flight Landing XII. The Muscles . . r 12.1 Structure 12.2 Control of muscular contraction 12.3 Patterns of muscular contraction 12.4 Energetics of muscle contraction XIII. Flight Activity . . . 13.1 Factors limiting flight activity 13.2 Factors promoting take-off 13.3 Stimuli leading to landing 13.4 Speed of flight *3-5 Types of flight 13.6 Migration . . . . . . 244 .271 SECTION C THE ABDOMEN, REPRODUCTION AND DEVELOPMENT XIV. The Abdomen . . . . . . . . . 14.1 Segmentation of the abdomen r14.2 Abdominal appendages XV. The Reproductive System . . . . . 305 . .318 MALE 15.1 Anatomy of male internal reproductive organs 15.2 Spermatogenesis 15.3 Transfer of sperm to the seminal vesicle FEMALE 15.4 Anatomy of female internal reproductive organs 15.5 Oogenesis 15.6 Resorption of oocytes 15.7 Ovulation XVI. Mating Behaviour and the Transferof Sperm to the Female MATING 16.1 Aggregation 16.2 Recognition and courtship 16.3 Pairing 16.4 Male aggression SPERM TRANSFER 16.5 External reproductive organs of the male 16.6 Copulation 16.7 Insemination 16.8 Post-copulatory behaviour 16.9 Effects of male accessory glands on oogenesis and oviposition . 350 xii CONTENTS XVII. Oviposition and the Egg . 17. i Female genitalia: the ovipositor 17.2 Oviposition r 17.3 The egg XVIII. 18.1 18.2 18.3 18.4 18.5 18.6 18.7 18.8 . . / . . 378 Embryology . Fertilisation Maturation of the oocytes Cleavage and formation of the blastoderm Early development of the embryo Movements of the embryo Development of organ systems Metabolic changes and control of organ development Duration of embryonic development XIX. U n u s u a l T y p e s o f D e v e l o p m e n t 19.1 Viviparity 19.2 P o l y e m b r y o n y 19.3 Parthenogenesis 19.4 Paedogenesis . . . XX. Hatching and Postembryonic Development 406 . . . . . . 433 . 449 HATCHING 20.1 Escape from the egg 20.2 Intermediate moult POSTEMBRYONIC DEVELOPMENT 20.3 20.4 20.5 20.6 20.7 Numbers of instars Growth Types of development Types of larvae Heteromorphosis XXI. Metamorphosis . . . 21.1 The pupa 21.2 Development of adult features 21.3 Adult emergence . . . . . . 473 SECTION D THE CUTICLE, RESPIRATION AND EXCRETION XXII. The Integument . . . . . . . . .501 22.1 Epidermis 22.2 Basic structure of cuticle 22.3 Different types of cuticle 22.4 Moulting 22.5 Cuticle formation 22.6 Expansion of the new cuticle 22.7 Control of moulting and cuticle production 22.8 Functions of the cuticle CONTENTS Xlll XXIII. 23.1 23.2 23.3 23.4 529 23.5 XXIV. 24.1 24.2 24.3 24.4 24.5 The Tracheal System and Respiration in Terrestrial Insects . The tracheal system Spiracles Moulting the tracheal system Gaseous exchange ^ Other functions of the tracheal system Respiration in Aquatic and Endoparasitic Insects . Aquatic insects obtaining oxygen from the air Insects obtaining oxygen from the water Insects subject to occasional submersion Respiration in endoparasitic insects Haemoglobin XXV. Excretion and Salt and Water Regulation 25.1 Excretory organs 25.2 Nitrogenous excretion 25.3 Excretion of ingested organic molecules 25.4 Maintenance of haemolymph ionic levels 25.5 Water regulation 25.6 Other functions of the Malpighian tubules . . . . . . 554 574 o SECTION E THE NERVOUS AND SENSORY SYSTEMS XXVI. 26.1 26.2 26.3 26.4 The Nervous System Structure of the nervous system Physiology of the nervous system Integration in the nervous system Learning 607 XXVII. 27.1 27.2 27.3 27.4 27.5 27.6 The Eyes and Vision Occurrence and structure of compound eyes Reception of light Functioning of the eye Dorsal ocelli Stemmata, Dermal light sense 642 XXVIII. Sound Production 28.1 Mechanisms and the sounds produced 28.2 Significance of the sounds produced 28.3 Control of sound production XXIX. 29.1 29.2 29.3 29.4 Mechanoreception . Mechanoreceptor neurones Trichoid sensilla Campaniform sensilla Chordotonal organs . . 675 . . . . .705 XIV CONTENTS 29.5 Stretch receptors 29.6 Statocysts 29.7 Pressure receptors XXX. 30.1 30.2 30.3 Chemorecption . . Olfaction Contact chemoreception Common chemical sense XXXI. 31.1 31.2 31.3 . . Temperature and Humidity Temperature Humidity Cryptobiosis . . . . . . . . . . 736 . 756 SECTION F THE BLOOD, HORMONES AND PHEROMONES XXXII. The Circulatory System 781 32.1 Structure 32.2 Circulation . XXXIII. The Haemolymph 33.1 Haemocytes 33.2 Connective tissue 33.3 Plasma . . XXXIV. The Endocrine Organs and Hormones 34.1 Endocrine organs 34.2 Dispersal of hormones 34.3 Mode of action of hormones 34.4 Chemical nature of hormones 34.5 Hormones and their functions 34.6 The rabbit flea and hormones . XXXV. 35.1 35.2 35.3 35.4 . . . . 796 . . Exocrine glands, pheromones and defensive secretions . The nature of exocrine secretions^ Structure of exocrine glands Pheromones Defensive secretions Taxonomic Index. Subject Index . . . . . . . . .851 . . 818 . 887 . 895
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