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Third Edition James L. Nation

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Third Edition
Insect Physiology
and Biochemistry
James L. Nation, Sr.
Third Edition
Insect Physiology
and Biochemistry
Third Edition
Insect Physiology
and Biochemistry
UNIVERSITY OF FLORIDA, GAINESVILLE, USA
Boca Raton London New York
CRC Press is an imprint of the

Taylor & Francis Group, an informa business
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Contents
Preface��xix
Author��xxi
Chapter 1
Embryogenesis.................................................................................................................................... 1
Preview...............................................................................................................................................1
1.1 Introduction.............................................................................................................................. 2
1.2 Morphogenesis.........................................................................................................................3
1.2.1 Egg, Fertilization, and Zygote Formation...................................................................3
1.2.2 Variations in Zygotic Nucleus Cleavage, Formation of Energids,
and Blastoderm Formation.........................................................................................4
1.2.2.1 Apterygota................................................................................................... 5
1.2.2.2 Hemimetabola............................................................................................. 7
1.2.2.3 Holometabola.............................................................................................. 8
1.2.3 Formation of the Germ Band...................................................................................... 9
1.2.4 Gastrulation................................................................................................................9
1.2.5 Germ Band Elongation............................................................................................. 10
1.2.6 Blastokinesis and Extraembryonic Membranes........................................................ 11
1.3 Genetic Control of Embryogenesis........................................................................................ 15
1.3.1 Development of a Model for Patterning.................................................................... 16
1.3.1.1 Bicoid Gene and Anterior Determination in Drosophila......................... 16
1.3.1.2 Posterior Group Genes and Posterior Pattern Formation.......................... 18
1.3.1.3 Genes Required in the Acron and Telson.................................................. 18
1.3.1.4 Dorsal–Ventral Axis.................................................................................. 19
1.4 Segmentation Genes............................................................................................................... 19
1.5 Homeotic Genes.....................................................................................................................20
1.5.1 Homeobox................................................................................................................. 21
1.6 Organogenesis........................................................................................................................ 21
1.6.1 Neurogenesis............................................................................................................. 21
1.6.2 Development of the Gut............................................................................................ 22
1.6.3 Malpighian Tubules.................................................................................................. 23
1.6.4 Tracheal System........................................................................................................ 23
1.6.5 Oenocytes................................................................................................................. 23
1.6.6 Cuticle Secretion in the Embryo............................................................................... 23
1.6.7 Cell Movements during Embryogenesis................................................................... 23
1.6.8 Programmed Cell Death: Apoptosis.........................................................................24
1.7 Hatching.................................................................................................................................24
1.8 Imaginal Discs.......................................................................................................................24
1.9 Review and Self-Study Questions..........................................................................................28
References.........................................................................................................................................28
v
vi Contents
Chapter 2
Digestion........................................................................................................................................... 33
Preview............................................................................................................................................. 33
2.1 Introduction............................................................................................................................ 33
2.2 Relationships between Food Habits and Gut Structure and Function...................................34
2.2.1 Plant versus Animal Origin: Solid versus Liquid Diet.............................................34
2.3 Major Structural Regions of the Gut...................................................................................... 36
2.3.1 Foregut...................................................................................................................... 36
2.3.2 Midgut....................................................................................................................... 39
2.3.3 Hindgut..................................................................................................................... 39
2.4 Midgut Cell Types..................................................................................................................40
2.4.1 Columnar Cells.........................................................................................................40
2.4.2 Regenerative Cells....................................................................................................40
2.4.3 Goblet Cells.............................................................................................................. 41
2.5 Microvilli or Brush Border of Midgut Cells..........................................................................44
2.6 Glycocalyx............................................................................................................................. 45
2.7 Peritrophic Matrix.................................................................................................................. 45
2.7.1 Functions of the Peritrophic Matrix..........................................................................46
2.8 Digestive Enzymes................................................................................................................. 47
2.8.1 Carbohydrate-Digesting Enzymes............................................................................ 48
2.8.2 Lipid-Digesting Enzymes......................................................................................... 49
2.8.3 Protein-Digesting Enzymes...................................................................................... 49
2.8.4 Do Proteinase Inhibitors in the Food Influence the Evolution of Proteinase
Secreted?.............................................................................................................. 51
2.9 Hormonal Influence on Midgut.............................................................................................. 51
2.10 Countercurrent Circulation of Midgut Contents and Absorption of Digested Products....... 53
2.11 Transepithelial and Oxidation–Reduction Potential of the Gut............................................. 55
2.12 Gut pH.................................................................................................................................... 55
2.13 Hematophagy: Feeding on Vertebrate Blood......................................................................... 57
2.14 Digestive System Morphology and Physiology in Major Insect Orders................................ 58
2.14.1 Orthoptera................................................................................................................. 58
2.14.2 Dictyoptera................................................................................................................ 59
2.14.3 Isoptera...................................................................................................................... 59
2.14.4 Hemiptera..................................................................................................................60
2.14.5 Homoptera................................................................................................................. 61
2.14.6 Coleoptera................................................................................................................. 61
2.14.7 Hymenoptera............................................................................................................. 62
2.14.8 Diptera...................................................................................................................... 62
2.14.9 Lepidoptera............................................................................................................... 63
2.15 Insect Gut as a Potential Target for Population Management and Control of the
Spread of Plant and Animal Disease Organisms...................................................................64
2.16 Review and Self-Study Questions.......................................................................................... 65
References......................................................................................................................................... 65
Chapter 3
Nutrition............................................................................................................................................ 75
Preview............................................................................................................................................. 75
3.1 Introduction............................................................................................................................ 75
3.2 Importance of Balance in Nutritional Components............................................................... 76
Contents vii
3.3 Ability of Insects to Self-Select Nutritional Components..................................................... 77

3.4 Requirements for Specific Nutrients...................................................................................... 77
3.4.1 Nitrogen Source: Proteins and Amino Acids............................................................ 78
3.4.2 Essential Amino Acids............................................................................................. 79
3.4.3 Carbohydrates........................................................................................................... 81
3.4.4 Lipids........................................................................................................................ 82
3.4.5 Sterols....................................................................................................................... 82
3.4.6 Polyunsaturated Fatty Acids..................................................................................... 83
3.4.7 Vitamins....................................................................................................................84
3.4.8 Minerals.................................................................................................................... 85
3.5 Techniques and Dietary Terms Used in Insect Nutrition Studies.......................................... 87
3.6 Criteria for Evaluating Nutritional Quality of a Diet............................................................. 87
3.7 Measures of Food Intake and Utilization............................................................................... 88
3.8 Phagostimulants.....................................................................................................................90
3.9 Feeding Deterrents.................................................................................................................92
3.10 Review and Self-Study Questions..........................................................................................92
References......................................................................................................................................... 93
Chapter 4
Integument and Molting....................................................................................................................99
Preview.............................................................................................................................................99
4.1 Introduction.......................................................................................................................... 100
4.2 Structure of the Integument................................................................................................. 100
4.2.1 Cuticulin Envelope.................................................................................................. 101
4.2.2 Epicuticle................................................................................................................. 101
4.2.3 Procuticle................................................................................................................ 102
4.2.4 Pore Canals and Wax Channels.............................................................................. 103
4.2.5 Epidermal Cells...................................................................................................... 103
4.3 Molting and Formation of New Cuticle............................................................................... 106
4.3.1 Apolysial Space....................................................................................................... 108
4.3.2 Molting Fluid Secretion.......................................................................................... 108
4.3.3 New Cuticle Formation........................................................................................... 108
4.3.4 Reabsorption of Molting Fluid................................................................................ 109
4.4 Ecdysis................................................................................................................................. 109
4.4.1 Shedding the Old Cuticle: Ecdysis.......................................................................... 112
4.4.2 Post-Ecdysis Wing Expansion and Waterproofing the New Cuticle...................... 113
4.4.3 Sclerotization of Cuticle.......................................................................................... 114
4.5 Chemical Composition of Cuticle........................................................................................ 116
4.5.1 Chitin...................................................................................................................... 116
4.5.2 Biosynthesis of Chitin............................................................................................. 121
4.5.3 Cuticular Proteins................................................................................................... 122
4.5.4 Resilin..................................................................................................................... 125
4.5.5 Stage-Specific Differences in Cuticle Proteins....................................................... 126
4.5.6 Protective Functions of Cuticle Proteins................................................................ 126
4.5.7 Cuticular Lipids...................................................................................................... 126
4.6 Mineralization of Insect Cuticles......................................................................................... 129
4.7 Capture of Atmospheric Water on Cuticular Surfaces........................................................ 129
4.8 Review and Self-Study Questions........................................................................................ 129
References....................................................................................................................................... 130
viii Contents
Chapter 5
Hormones and Development........................................................................................................... 135
Preview........................................................................................................................................... 135
5.1 Introduction.......................................................................................................................... 136
5.2 Historical Beginnings for the Concept of Hormonal Control of Molting
and Metamorphosis.............................................................................................................. 136
5.3 Interplay of PTTH, Ecdysteroids, and Juvenile Hormone Control Development............... 137
5.4 Brain Neurosecretory Cells and Prothoracicotropic Hormone............................................ 141
5.4.1 Source and Chemistry............................................................................................. 141
5.4.2 Bioassay for PTTH Activity................................................................................... 141
5.4.3 Stimuli for the Secretion of PTTH......................................................................... 143
5.4.4 PTTH Secretion after Brain Activation by Stretch Receptors................................ 143
5.4.5 Gated PTTH Secretion in Tobacco Hornworm...................................................... 144
5.4.6 Secretion of PTTH after Brain Activation by Cold Exposure................................ 144
5.4.7 Regulation of Tissue and Hemolymph Levels of PTTH......................................... 144
5.4.8 Mode of Action of PTTH........................................................................................ 145
5.5 Prothoracic Glands and Ecdysteroids.................................................................................. 146
5.5.1 Biosynthesis of Ecdysone....................................................................................... 146
5.5.2 Conversion of Ecdysone into 20-Hydroxyecdysone............................................... 148
5.5.3 Molecular Diversity in the Structure of the Molting Hormone.............................. 148
5.5.4 Calliphora Assay for Ecdysteroids......................................................................... 151
5.5.5 Radioimmunoassay for Ecdysone and Related Ecdysteroids................................. 151
5.5.6 Assay by Physicochemical Techniques................................................................... 152
5.5.7 Tissues and Cell Cultures Used in Assays.............................................................. 153
5.5.8 Degradation of Ecdysone........................................................................................ 153
5.5.9 Virus Degradation of Host Ecdysteroids................................................................ 155
5.5.10 Dependence of Some Parasitoids on Host Ecdysteroids......................................... 155
5.6 Corpora Allata and Juvenile Hormones............................................................................... 155
5.6.1 Glandular Source and Chemistry........................................................................... 155
5.6.2 Assays for JH Activity............................................................................................ 157
5.6.3 Regulation of the Tissue and Hemolymph Levels of JH........................................ 157
5.6.4 Insect Growth Regulators and Compounds That Are Cytotoxic
to the Corpora Allata.............................................................................................. 161
5.6.5 Cellular Mode of Action and Receptors for JH...................................................... 162
5.6.6 Downstream Transcription Factors......................................................................... 163
5.7 Mode of Action of Ecdysteroids at the Gene Level............................................................. 164
5.7.1 Chromosomal Puffs................................................................................................ 164
5.7.2 Isolation of an Ecdysteroid Receptor...................................................................... 166
5.7.3 Differential Tissue and Cell Response to Ecdysteroids.......................................... 168
5.8 Possible Timer Gene in the Molting Process....................................................................... 170
5.9 Ecdysone–Gene Interaction Ideas Stimulated Vertebrate Work......................................... 170
References....................................................................................................................................... 172
Chapter 6
Biological Rhythms........................................................................................................................ 181
Preview........................................................................................................................................... 181
6.1 Introduction.......................................................................................................................... 182
6.2 Characteristics of Circadian and Photoperiodic Rhythms.................................................. 182
Contents ix
6.3 Molecular Basis for the Circadian Clock............................................................................. 183

6.4 Evidence for Clock Genes in Many Insects......................................................................... 186
6.5 Examples of Circadian Functions in Insects........................................................................ 186
6.5.1 Circadian Regulation of Hormone Secretion.......................................................... 186
6.5.2 Circadian Clock Influence in Peripheral Organs and Tissues................................ 188
6.5.3 Circadian Clock Influence in Social Behavior of Honeybees................................. 189
6.5.4 Circadian Clock Influence in Reproduction........................................................... 190
6.6 Photoperiodic Response: One Clock, Two Clocks, or Multiple Clocks?............................. 193
6.7 Clock Models Based on Experimental Responses of Insects to Varying
Light/Dark Regimes................................................................................................. 195
6.7.1 Hourglass Model..................................................................................................... 196
6.7.2 External Coincidence Model.................................................................................. 197
6.7.3 Internal Coincidence Model................................................................................... 197
6.7.4 Resonance Model.................................................................................................... 197
6.7.5 Summary Results from Model Experiments.......................................................... 198
6.8 Conclusions.......................................................................................................................... 198
References....................................................................................................................................... 199
Chapter 7
Diapause..........................................................................................................................................207
Preview...........................................................................................................................................207
7.1 Introduction..........................................................................................................................207
7.2 Diapause: A Survival Strategy.............................................................................................208
7.3 Phases of Diapause............................................................................................................... 210
7.3.1 Prediapause: Induction and Preparation................................................................. 210
7.3.2 Diapause: Initiation and Maintenance.................................................................... 211
7.3.3 Diapause Termination............................................................................................. 212
7.4 Hormonal Control of Diapause............................................................................................ 212
7.4.1 Embryonic Diapause............................................................................................... 212
7.4.2 Larval Diapause...................................................................................................... 213
7.4.3 Pupal Diapause....................................................................................................... 213
7.4.4 Adult Diapause........................................................................................................ 214
7.5 Role of Daily and Seasonal Biological Clocks in Diapause................................................ 215
7.6 Diapause and Gene Expression............................................................................................ 215
7.7 Nutrient Accumulation for Diapause and the Storage and Conservation of Nutrients
during Diapause................................................................................................................... 216
7.8 Molecular Studies of Diapause............................................................................................ 217
References....................................................................................................................................... 219
Chapter 8
Intermediary Metabolism............................................................................................................... 223
Preview........................................................................................................................................... 223
8.1 Introduction: Nutrient Stores—The Fat Body......................................................................224
8.2 Energy Demands for Insect Flight....................................................................................... 225
8.3 Metabolic Stores................................................................................................................... 226
8.3.1 Carbohydrate Resources......................................................................................... 226
8.3.1.1 Trehalose Resources................................................................................ 226
8.3.1.2 Glycogen: Storage and Synthesis............................................................ 229
x Contents
8.4 Hormones Controlling Carbohydrate Metabolism.............................................................. 230

8.5 Pathways of Metabolism Supporting Intense Muscular Activity, Such as Flight................ 230
8.5.1 Glycolysis................................................................................................................ 230
8.5.1.1 Glycerol-3-Phosphate Shuttle and Regeneration of NAD+...................... 232
8.5.1.2 Significance and Control of the Glycerol-3-Phosphate Shuttle............... 235
8.5.2 Krebs Cycle............................................................................................................. 235
8.5.2.1 Control of Krebs Cycle Metabolism and the Regulation
of Carbohydrate Metabolism in Flight Muscles...................................... 237
8.5.3 Electron Transport System...................................................................................... 237
8.5.4 Proline as a Fuel for Flight.....................................................................................240
8.5.5 Mobilization and the Use of Lipids for Flight Energy............................................ 243
8.5.5.1 Transport of Lipids by Lipophorin.......................................................... 247
8.5.5.2 Activation of Fatty Acids, Entry into Mitochondria, and β-Oxidation........247
References....................................................................................................................................... 249
Chapter 9
Neuroanatomy................................................................................................................................. 255
Preview........................................................................................................................................... 255
9.1 Introduction.......................................................................................................................... 256
9.2 Central Nervous System....................................................................................................... 256
9.3 Brain..................................................................................................................................... 258
9.3.1 Protocerebrum........................................................................................................ 259
9.3.2 Deutocerebrum.......................................................................................................260
9.3.2.1 Antennal Mechanosensory and Motor Center Neuropil......................... 261
9.3.2.2 Antennal Lobe......................................................................................... 261
9.3.3 Tritocerebrum......................................................................................................... 263
9.4 Ventral Ganglia....................................................................................................................264
9.4.1 Abdominal Ganglia................................................................................................. 265
9.4.2 Lateral Nerves.........................................................................................................266
9.5 Oxygen and Glucose Supply to the Brain and Ganglia........................................................266
9.6 Neuropil............................................................................................................................... 267
9.7 Hemolymph–Brain (CNS) Barrier....................................................................................... 268
9.8 Neurons: Building Blocks of a Nervous System.................................................................. 269
9.8.1 Afferent or Sensory Neurons.................................................................................. 270
9.8.2 Efferent or Motor Neurons...................................................................................... 270
9.8.3 Interneurons............................................................................................................ 271
9.8.4 Glial Cells............................................................................................................... 272
9.9 Giant Axons in the Insect Central Nervous System............................................................ 273
9.10 Nervous System Control of Behavior: Motor Programs...................................................... 274
9.10.1 Motor Program That Controls Walking.................................................................. 274
9.10.2 Motor Pattern for Rhythmic Breathing.................................................................. 274
9.11 Neurosecretory Cells and Neurosecretion Products from the CNS..................................... 276
9.11.1 Neurosecretory Cells.............................................................................................. 276
9.11.2 Adipokinetic Hormone........................................................................................... 278
9.11.3 Proctolin.................................................................................................................. 278
9.11.4 FMRFamide-Related Peptides................................................................................ 278
9.11.5 Tachykinins: Locustatachykinins and Leucokinins............................................... 279
9.11.6 Pigment-Dispersing Factors.................................................................................... 279
Contents xi
9.11.7 Vasopressin-Like Peptide (Locust F2 Peptide)....................................................... 279

9.11.8 Allatotropins and Allatostatins...............................................................................280
9.11.9 Crustacean Cardioactive Peptide............................................................................280
9.11.10 Pheromone Biosynthesis Activating Neuropeptide................................................280
Review and Self-Study Questions................................................................................................... 281
References....................................................................................................................................... 281
Chapter 10
Neurophysiology............................................................................................................................. 285
Preview........................................................................................................................................... 285
10.1 Introduction.......................................................................................................................... 286
10.2 Nerve Cell Responses to Stimuli......................................................................................... 286
10.2.1 Graded Responses................................................................................................... 287
10.2.2 Spike Potentials....................................................................................................... 288
10.3 Physiological Basis for Neuronal Responses to Stimuli...................................................... 289
10.3.1 Membrane Ion Channels: Bioelectric Potentials..................................................... 289
10.3.2 Resting Potential..................................................................................................... 292
10.3.3 The Action Potential: Sodium Activation............................................................... 294
10.3.4 Sodium Inactivation and Repolarization................................................................ 296
10.3.5 Measurement of Ion Fluxes: Voltage Clamp Technique......................................... 297
10.4 Conduction of the Action Potential: Local Circuit Theory.................................................. 298
10.5 Physiology and Biochemistry at the Synapse: Excitatory and Inhibitory Postsynaptic
Potentials.............................................................................................................................. 299
10.6 Acetylcholine-Mediated Synapses....................................................................................... 301
10.6.1 Action of Acetylcholine at the Synapse.................................................................. 301
10.6.2 Nicotinic and Muscarinic Cholinergic Receptors in Insects..................................302
10.6.3 Acetylcholine Receptor Structure........................................................................... 303
10.7 Electric Transmission across Synapses................................................................................304
10.8 Neuromuscular Junctions.....................................................................................................304
10.9 Review and Self-Study Questions........................................................................................304
References....................................................................................................................................... 305
Chapter 11
Muscles...........................................................................................................................................309
Preview...........................................................................................................................................309
11.1 Introduction.......................................................................................................................... 310
11.2 Basic Muscle Structure and Function.................................................................................. 310
11.2.1 Macro- and Microstructure of Muscle.................................................................... 310
11.2.2 Muscle Attachments to the Exoskeleton................................................................. 313
11.2.3 Skeletal Muscle....................................................................................................... 314
11.2.4 Polyneuronal Innervation and Multiterminal Nerve Contacts............................... 314
11.2.5 Transmitter Chemical at Nerve–Muscle Junctions................................................. 316
11.3 Synchronous and Asynchronous Muscles............................................................................ 317
11.4 Muscle Proteins and Physiology of Contraction.................................................................. 320
11.4.1 Active State: Binding of Myosin Heads to Actin and the Sliding of Filaments..... 322
11.4.2 Release of Myosin Heads from Actin..................................................................... 322
11.5 Muscles Involved in General Locomotion, Running, and Jumping..................................... 323
11.5.1 Adaptations for Running, Walking, and Survival.................................................. 323
11.5.2 Adaptations for Jumping......................................................................................... 325
xii Contents
11.6 Sound Production: Tymbal and Stridulatory Muscle........................................................... 327

11.6.1 Tymbal Morphology and Physiology...................................................................... 327
11.6.2 Stridulatory Muscle Physiology.............................................................................. 328
11.7 Morphology and Physiology of Nonskeletal Muscle........................................................... 329
11.7.1 Visceral Muscles..................................................................................................... 329
11.7.2 Heart Muscle........................................................................................................... 329
11.7.3 Alary Muscles......................................................................................................... 330
References....................................................................................................................................... 330
Chapter 12
Flight............................................................................................................................................... 335
Preview........................................................................................................................................... 335
12.1 Introduction.......................................................................................................................... 335
12.2 Thoracic Structure, Wing Hinges, and Muscle Groups Involved in Flight......................... 336
12.3 Wing Strokes........................................................................................................................ 338
12.4 Multiple Contractions from Each Volley of Nerve Impulses to Asynchronous Muscles.... 339
12.5 Flight in Dragonflies and Damselflies.................................................................................340
12.6 Aerodynamics of Lift and Drag Forces Produced by Wings............................................... 341
12.6.1 Lift Forces Generated by Clap and Fling Wing Movements.................................. 342
12.6.2 Lift Forces Derived from Drag and Delayed Stall................................................. 343
12.7 Hovering Flight....................................................................................................................344
12.8 Control of Pitch and Twisting of Wings...............................................................................346
12.9 Power Output of Flight Muscles..........................................................................................346
12.10 Metabolic Activity of Wing Muscles................................................................................... 347
12.11 Flight Behavior.....................................................................................................................348
References....................................................................................................................................... 349
Chapter 13
Sensory Systems............................................................................................................................. 353
Preview........................................................................................................................................... 353
13.1 Introduction.......................................................................................................................... 353
13.2 External and Internal Receptors Monitoring the Environment........................................... 355
13.3 General Functional Classification of Sensory Receptors..................................................... 355
13.3.1 Receptors with Multiple Pores................................................................................ 356
13.3.2 Receptors with a Single Pore.................................................................................. 357
13.3.3 Receptors without Pores......................................................................................... 357
13.4 Mechanoreceptors................................................................................................................ 357
13.4.1 Structure of a Simple Tactile Hair: A Mechanoreceptor Sensillum....................... 357
13.4.2 Hair Plates............................................................................................................... 358
13.4.3 Chordotonal Sensilla............................................................................................... 358
13.4.4 Subgenual Organs................................................................................................... 359
13.4.5 Tympanal Organs: Specialized Organs for Airborne Sounds................................360
13.4.6 Johnston’s Organ.....................................................................................................364
13.4.7 Simple Chordotonal Organs....................................................................................364
13.4.8 Thermoreceptors and Hygroreceptors.................................................................... 365
13.4.9 Infrared Reception.................................................................................................. 366
Contents xiii
13.5 Chemoreceptors................................................................................................................... 368

13.5.1 Olfactory Sensilla: Dendritic Fine Structure.......................................................... 368
13.5.2 Contact Chemoreceptors–Gustatory Receptors..................................................... 369
13.5.3 Specialists versus Generalists among Chemoreceptors.......................................... 371
13.5.4 Stimulus–Receptor Excitation Coupling................................................................ 371
References....................................................................................................................................... 372
Chapter 14
Vision.............................................................................................................................................. 377
Preview........................................................................................................................................... 377
14.1 Introduction.......................................................................................................................... 377
14.2 Compound Eye Structure..................................................................................................... 379
14.3 Dioptric Structures............................................................................................................... 381
14.4 Corneal Layering................................................................................................................. 383
14.5 Retinula Cells....................................................................................................................... 383
14.6 Rhabdomeres........................................................................................................................ 384
14.7 Electrical Activity of Retinula Cells.................................................................................... 384
14.8 Neural Connections in the Optic Lobe................................................................................ 384
14.9 Ocelli.................................................................................................................................... 385
14.10 Larval Eyes: Stemmata........................................................................................................ 386
14.11 Dermal Light Sense............................................................................................................. 387
14.12 Chemistry of Insect Vision.................................................................................................. 388
14.13 Visual Cascade..................................................................................................................... 389
14.14 Regulation of the Visual Cascade........................................................................................ 390
14.15 Color Vision......................................................................................................................... 391
14.16 Vision Is Important in Behavior........................................................................................... 394
14.17 Nutritional Need for Carotenoids in Insects........................................................................ 394
14.18 Detection of Plane-Polarized Light...................................................................................... 395
14.19 Visual Acuity....................................................................................................................... 397
References.......................................................................................................................................400
Chapter 15
Circulatory System.........................................................................................................................405
Preview...........................................................................................................................................405
15.1 Introduction: Embryonic Development of the Circulatory System and Hemocytes............405
15.2 Dorsal Vessel: Heart and Aorta...........................................................................................406
15.2.1 Alary Muscles.........................................................................................................409
15.2.2 Ostia........................................................................................................................ 410
15.2.3 Heartbeat................................................................................................................. 410
15.2.4 Ionic Influences on Heartbeat................................................................................. 411
15.2.5 Nerve Supply to the Heart...................................................................................... 411
15.2.6 Cardioactive Secretions.......................................................................................... 412
15.3 Accessory Pulsatile Hearts.................................................................................................. 413
15.4 Hemocytes............................................................................................................................ 414
15.4.1 Functions of Hemocytes......................................................................................... 416
15.4.2 Hemocytopoietic Tissues and Origin of Hemocytes.............................................. 417
15.4.3 Number of Circulating Hemocytes......................................................................... 418
xiv Contents
15.5 Hemolymph.......................................................................................................................... 420

15.5.1 Functions of Hemolymph and Circulation.............................................................. 420
15.5.2 Hemolymph Volume............................................................................................... 422
15.5.3 Coagulation of Hemolymph.................................................................................... 422
15.5.4 Hemolymph pH and Hemolymph Buffers.............................................................. 424
15.5.5 Chemical Composition of Hemolymph.................................................................. 425
15.5.5.1 Inorganic Ions.......................................................................................... 426
15.5.5.2 Free Amino Acids................................................................................... 426
15.5.5.3 Proteins.................................................................................................... 427
15.5.5.4 Other Organic Constituents..................................................................... 427
15.6 Rate of Circulation............................................................................................................... 428
15.7 Hemoglobin.......................................................................................................................... 428
References....................................................................................................................................... 429
Chapter 16
Immunity........................................................................................................................................ 433
Preview........................................................................................................................................... 433
16.1 Introduction.......................................................................................................................... 433
16.2 Physical Barriers to Invasion............................................................................................... 435
16.3 Cellular Immune Reactions................................................................................................. 436
16.4 Recognition of Nonself........................................................................................................ 437
16.5 Synthesis of Antifungal and Antibacterial Peptides............................................................ 438
16.6 Toll Pathway for the Synthesis of Antimicrobial Peptides.................................................. 439
16.7 IMD Pathway for the Synthesis of Antimicrobial Peptides................................................. 442
16.8 C-Type Lectins..................................................................................................................... 443
16.9 Serpins.................................................................................................................................. 443
16.10 Ecology, Behavior, and Immunity.......................................................................................444
16.11 Cost of Defense.................................................................................................................... 445
16.12 Coevolutionary Race between Parasitoid Escape Mechanisms and Host Defense
Mechanisms.........................................................................................................................446
16.13 Autoimmune Consequences of Some Defense Reactions.................................................... 447
16.14 Gender Differences in Immune Responses.......................................................................... 447
16.15 Conclusions.......................................................................................................................... 447
References.......................................................................................................................................448
Chapter 17
Respiration...................................................................................................................................... 455
Preview........................................................................................................................................... 455
17.1 Introduction.......................................................................................................................... 455
17.2 Structure of the Tracheal System......................................................................................... 457
17.2.1 Tracheae and Tracheole Structure.......................................................................... 457
17.2.2 Spiracle Structure and Function............................................................................. 459
17.2.3 Tracheal Epithelium................................................................................................ 462
17.2.4 Development of New Tracheoles............................................................................ 462
17.2.5 Air Sacs...................................................................................................................464
17.2.6 Molting of Tracheae................................................................................................464
17.3 Tracheal Supply to Tissues and Organs...............................................................................464
17.3.1 Adaptations of Tracheae to Supply Flight Muscles................................................465
Contents xv
17.4 Ventilation and Diffusion of Gases within the System........................................................466

17.4.1 Simple Diffusion Is Usually Not Adequate............................................................ 467
17.4.2 Active Ventilation of Tracheae............................................................................... 467
17.4.3 Diffusion from Tracheoles to Mitochondria........................................................... 471
17.5 Discontinuous Gas Exchange............................................................................................... 471
17.6 Water Balance during Flight................................................................................................ 475
17.7 Gas Exchange in Aquatic Insects......................................................................................... 475
17.7.1 Compressible Gas Gills........................................................................................... 476
17.7.2 Incompressible Gas Gills: A Plastron..................................................................... 477
17.7.3 Use of Aquatic Plants as Air Source....................................................................... 477
17.7.4 Cutaneous Respiration: Closed Tracheal System in Some Aquatic Insects........... 478
17.8 Respiration in Endoparasitic Insects.................................................................................... 481
17.9 Respiratory Pigments........................................................................................................... 481
17.10 Respiration in Eggs and Developing Embryos..................................................................... 481
17.11 Nonrespiratory Functions of the Tracheal System............................................................... 482
References.......................................................................................................................................484
Chapter 18
Excretion......................................................................................................................................... 489
Preview........................................................................................................................................... 489
18.1 Introduction.......................................................................................................................... 489
18.2 Malpighian Tubules.............................................................................................................. 490
18.3 Malpighian Tubule Cells...................................................................................................... 493
18.4 Formation of Primary Urine in Malpighian Tubules........................................................... 494
18.5 Proton Pump as a Driving Mechanism for Urine Formation and Homeostasis.................. 495
18.6 Selective Reabsorption in the Hindgut................................................................................ 499
18.6.1 Anatomical Specialization of Hindgut Epithelial Cells......................................... 499
18.6.2 Secretion and Reabsorption in the Ileum................................................................ 501
18.6.3 Reabsorption in the Rectum................................................................................... 501
18.7 Role of the Excretory System in Maintaining Homeostasis................................................ 502
18.7.1 Electrolyte Homeostasis......................................................................................... 502
18.7.2 Water Homeostasis.................................................................................................. 503
18.7.2.1 Diuretic Hormones..................................................................................504
18.7.2.2 Antidiuretic Hormones............................................................................506
18.7.3 Acid–Base Homeostasis..........................................................................................506
18.7.4 Nitrogen Homeostasis............................................................................................. 507
18.7.4.1 Ammonia Excretion................................................................................ 508
18.7.4.2 Uric Acid Synthesis and Excretion..........................................................509
18.8 Cryptonephridial Systems.................................................................................................... 511
18.9 Self-Study and Review Questions........................................................................................ 513
References....................................................................................................................................... 514
Chapter 19
Semiochemicals.............................................................................................................................. 523
Preview........................................................................................................................................... 523
19.1 Introduction.......................................................................................................................... 524
19.2 Classes of Semiochemicals.................................................................................................. 524
19.3 Importance of the Olfactory Sense in Insects...................................................................... 525
xvi Contents
19.4 Active Space Concept.......................................................................................................... 528

19.5 Pheromones Classified according to Behavior Elicited....................................................... 528
19.6 Pheromone Parsimony......................................................................................................... 529
19.7 Chemical Characteristics of Semiochemicals...................................................................... 529
19.8 Insect Receptors and the Detection Process........................................................................ 533
19.8.1 Pheromone-Binding Proteins.................................................................................. 533
19.8.2 Signal Transduction and Receptor Response.......................................................... 535
19.8.3 Pheromone Inactivation and Clearing of the Receptor........................................... 538
19.8.4 Do Insects Smell the Blend or Just the Major Components?.................................. 538
19.9 Information Coding and Processing.................................................................................... 539
19.9.1 Structure of Odor Plumes....................................................................................... 539
19.9.2 Pheromone Signal Processing................................................................................. 541
19.10 Hormonal Control of Pheromone Synthesis and Release.................................................... 543
19.10.1 Mode of Action of PBAN.......................................................................................544
19.11 Biosynthesis of Pheromones................................................................................................ 545
19.12 Geographical and Population Differences and Evolution of Pheromone Blends................. 547
19.13 Practical Applications of Pheromones................................................................................. 548
19.13.1 Mechanisms Operating in Mating Disruption........................................................ 549
19.13.1.1 Sensory Fatigue....................................................................................... 549
19.13.1.2 False Trail Following.............................................................................. 549
19.13.1.3 Camouflage of Natural Pheromone Plume.............................................. 550
19.13.1.4 Pheromone Antagonists and Imbalanced Blends.................................... 550
References....................................................................................................................................... 551
Chapter 20
Reproduction................................................................................................................................... 561
Preview........................................................................................................................................... 561
20.1 Introduction.......................................................................................................................... 562
20.2 Female Reproductive System............................................................................................... 562
20.2.1 Structure of Ovaries................................................................................................ 562
20.2.1.1 Panoistic Ovarioles..................................................................................564
20.2.1.2 Telotrophic Ovarioles..............................................................................564
20.2.1.3 Polytrophic Ovarioles.............................................................................. 565
20.2.1.4 Oviposition.............................................................................................. 565
20.2.2 Nutrients for Oogenesis.......................................................................................... 566
20.2.3 Hormonal Regulation of Ovary Development and Synthesis of Egg Proteins....... 567
20.3 Vitellogenins and Yolk Proteins.......................................................................................... 572
20.3.1 Biochemical Characteristics of Vitellogenins and Yolk Proteins........................... 572
20.3.2 Yolk Proteins of Higher Diptera............................................................................. 573
20.4 Sequestering of Vitellogenins and Yolk Proteins by Oocytes............................................. 574
20.4.1 Patency of Follicular Cells...................................................................................... 574
20.4.2 Egg Proteins Produced by Follicular Cells............................................................. 576
20.4.3 Proteins in Addition to Vitellogenin and Yolk Proteins in the Egg........................ 576
20.5 Formation of the Vitelline Membrane................................................................................. 576
20.6 Chorion................................................................................................................................. 576
20.7 Gas Exchange in Eggs.......................................................................................................... 577
Contents xvii
20.8 Male Reproductive System.................................................................................................. 577

20.8.1 Apyrene and Eupyrene Sperm of Lepidoptera....................................................... 579
20.8.2 Male Accessory Glands.......................................................................................... 580
20.8.3 Transfer of Sperm................................................................................................... 580
20.9 Gender Determination......................................................................................................... 581
References....................................................................................................................................... 583
Chapter 21
Insect Symbioses............................................................................................................................. 591
Preview........................................................................................................................................... 591
21.1 Introduction.......................................................................................................................... 592
21.2 Symbioses among Leaf-Cutting Ants, Fungi, and Bacteria................................................ 593
21.3 Biology of Termites.............................................................................................................. 595
21.3.1 Symbionts in Termites............................................................................................ 596
21.3.2 Lignocellulose Structure......................................................................................... 598
21.3.3 Nitrogen Metabolism.............................................................................................. 599
21.3.4 Fungal Culture........................................................................................................ 599
21.4 Bark and Ambrosia Beetles and Their Symbionts............................................................... 599
21.4.1 Ambrosia Beetles....................................................................................................600
21.4.2 Bark Beetles............................................................................................................603
21.4.3 Fungal Role in Supplementing Limited Nutrients in Wood and Phloem...............603
21.4.4 Evolution of Fungal Feeding in Bark Beetles.........................................................603
21.4.5 Bacteria as Part of the Bark Beetle Holobiont........................................................604
21.4.6 Anthropogenic Effects upon Bark Beetles and Their Symbionts...........................605
21.5 Buchnera in Aphids.............................................................................................................605
21.6 Tsetse Fly Symbionts...........................................................................................................606
21.7 Wolbachia............................................................................................................................607
21.7.1 Cytoplasmic Incompatibility Inducing the Effect of Wolbachia............................608
21.7.2 Parthenogenesis Inducing the Effect of Wolbachia................................................609
21.7.3 Feminizing Strains of Wolbachia........................................................................... 610
References....................................................................................................................................... 611
Index�� 623
Preface
Research work and literature in physiology and biochemistry of insects continues to expand in an
explosive and exponential fashion. New journals continue to appear; the literature is very wide-
spread. Experienced and established scientists are likely to find time constraints that make it dif-
ficult or impossible to read the general field of insect physiology, and the problem is even more acute
for graduate students or undergraduates who are just getting acquainted with the physiology of
insects. Even though the Internet and online journals and books offer wonderful advantages, consid-
erable time is still required. My aim in this third edition of Insect Physiology and Biochemistry is to
produce a book that will be useful to advanced undergraduate and graduate students who may have
to take a course in insect physiology, and I hope the book will aid working scientists in a variety
of disciplines who conduct research with insects but may have limited time to read general insect
physiology. All the topics included in the last edition have been retained, with the addition of recent
references to each chapter, more than 500 additional references to those previous chapters. Two new
chapters—Biological Rhythms and Insect Symbioses—have been added, with an additional 300+
references, so the book now includes about 2600 references to the literature. Another new feature
in this edition is the inclusion of Review and Self-Study Questions at the end of each chapter. I have
found these helpful, both in classroom teaching and in a distance education approach with students
when I could only communicate with them by e-mail.
I thank Dr. N. Krishnan for suggesting the topic of biological rhythms and for helpful comments
on an early version of the chapter and pointing me to some important resources. I am indebted to
Dr. Guy Bloch, who read a later version of the chapter and provided very helpful comments. The
chapter on insect symbioses was read by Dr. Jiri Hulcr and Dr. Kirsten S. Pelz-Stelinski, both of
whom provided very helpful suggestions. I thank again those who read and advised me on the
previous editions—Glenn Hall, Marie Nation Becker, Jon Harrison, Tom Miller, and anonymous
reviewers.
I am indebted to the many persons who sent photographs to use in the book: Ring Cardé,
Herb Oberlander, D.L. Musolin, Jiri Hulcr, M. T. Kasson, Lyle Buss, Andrei Sourakov, Gretert
Montano, Marie Nation Becker, Rochelle Nation, Hanife Genc, Maria and Tom Eisner, J. N.
Holland, W.B. Hunter, Jerry Butler, Ethel Villabos and Todd Shelly, Ritsuo Nishida, Clay Smith,
V. Leclerc and J.-M Reichhart, Alexandra Shapiro, Jimmy Becnel, K. Tomioka, N. Peschel
and C. Helfrich-Förster, Thomas Chouvenc, Guy Bloch, D. L. Musolin, C. Hermann, Brian
Forschler, Jarrod Scott, Y. Hongoh, M.-Y. Choi, Peter Teal, Paul Shirk, Al Handler, Coby Schal,
D.O. Deonier, Robin Giblin-Davis, C. Bordereau, and C.R. Currie.
I thank John Sulzycki, CRC Press, Taylor & Francis Group, for requesting a revision and for
offering 24 pages of color illustrations. David Fausel was initially helpful in handling questions
and the manuscript. I thank Todd Perry for managing the book project, Gopinath Chandrasekaran
for excellent copy editing, and the illustrations editor for arranging the color illustrations. I thank
Kathy Milne, who constructed many of the line drawings on a computer. I thank the Department of
Entomology and Nematology, University of Florida, Gainesville, for allowing me to hold the title
of emeritus professor after retirement and for giving me an office and computer facilities. Finally,
I thank my wife, Dorothy, who has been patient as I waded through a thousand research papers and
numerous books in order to update this edition.

Professor Emeritus
Entomology and Nematology
University of Florida
Gainesville, Florida
xix
Author
James L. Nation, Sr., PhD, is professor emeritus of entomol-
ogy at the University of Florida, Gainesville. He holds a BSc in
entomology from Mississippi State University, Starkville, and a
PhD in entomology from Cornell University, Ithaca, New York.
His special interest in entomology is the physiology and biochem-
istry of insects. Prior to retirement in 2003, he taught and con-
ducted research at the University of Florida for 43 years. Although
he sometimes taught other courses, he introduced a graduate-
level course and mainly taught graduate-level insect physiology.
Research activities included work in insect excretion, pheromones,
cuticular hydrocarbons, and insect nutrition. He served as an asso-
ciate editor of Florida Entomologist from 1967 to 1969, as an editor of the Journal of Chemical
Ecology from 1994 to 2000, and as editor of Florida Entomologist from 2004 to 2010. He has
continued to teach a course in the Undergraduate Honors Program called Global Environmental
Issues each fall term since retirement and introduced a graduate-level course in insect physiol-
ogy at Florida A&M University in Tallahassee, Florida, in 2006 (taught by interactive television).
He wrote both the first (2002) and the second (2008) edition of Insect Physiology and Biochemistry.
He is active in volunteer activities and gardening.
xxi
Chapter 1
Embryogenesis
PREVIEW
Insect eggs have a central yolk surrounded by a layer of cytoplasm. A proteinaceous chorion put
on the egg while it is in the ovary provides a protective covering for the egg. Sperm released from
the spermatheca of the female passes through the micropyle, a narrow channel through the chorion,
as the egg passes down the oviduct on its way to be deposited in the environment. Usually, the egg
nucleus is diploid until the entry of the sperm stimulates meiotic division leading to the haploid egg
nucleus. The union of a sperm nucleus with the egg nucleus produces the zygote and stimulates the
zygote to begin divisions. Complete cleavage of the zygotic yolk and cytoplasm occurs in eggs of
some species during the first few divisions, but yolk cleavage ceases after a few divisions. In most
species, cleavage of yolk and cytoplasm is incomplete from the beginning. Ultimately, zygotic divi-
sions in all insect eggs produce large numbers of nuclei lacking cell membranes but each surrounded
with a small field of cytoplasm. These nuclei and associated cytoplasm are called energids. Energids
gradually migrate into a single layer near the periphery of the egg, forming the blastoderm. Cell
membranes become complete after blastoderm formation. A few cells, the pole cells, aggregate at
the posterior end of the egg and are the first to become committed to a future developmental track;
they will become the gametes of the adult. Cells on the ventral side of the blastoderm enlarge and
become committed as the germ band, the cells that will become the embryo. Subsequent develop-
ment of the germ band is controlled by maternal and zygotic genes. Maternal genes are present and
active in the nurse cells of the mother during oogenesis. The mother’s nurse cells pass maternal
gene transcripts (mRNAs) into the developing oocyte in the ovary, and these begin to function in
the zygote. The maternal gene transcripts are translated into proteins in the zygote, and one of the
earliest actions of these proteins is control of anterior–posterior and dorsal–ventral axes orienta-
tion of the embryo. Later, acting zygotic genes include gap genes that divide the embryo into large
domains, pair-rule genes that divide the domains into parasegments, and finally segment polarity
genes that control formation of true segments. Homeotic genes begin to function during paraseg-
ment formation to give each segment its characteristic identity. Organogenesis leads to formation of
the organ systems of the embryo. Insects with complete development retain within the larval body
small embryonic clusters of cells called imaginal discs that divide, differentiate, and grow into adult
structures during pupation.
1
2 INSECT PHYSIOLOGY AND BIOCHEMISTRY
1.1 INTRODUCTION
The three major divisions in the Insecta, the Apterygota, Hemimetabola, and Holometabola, are
not directly ancestral to each other, and consequently embryological developments in the groups,
although similar in some respects, often are divergent. The Apterygota (Protura, Collembola,
Diplura, and Thysanura) never evolved wings and lack metamorphosis. The immatures look just
like small versions of the adults. The Hemimetabola (Orthoptera, Hemiptera, Heteroptera, and oth-
ers) evolved wings and have gradual metamorphosis. Immatures have some adult features but lack
wings. The Holometabola (Coleoptera, Hymenoptera, Lepidoptera, Diptera, and others) evolved
wings and have complete metamorphosis. Immature forms are typically wormlike and thus look
very different from the adults. Wingless adults occur in the Hemimetabola and Holometabola, but
the wingless condition evolved secondarily from winged forms.
The goals for this chapter are to describe the morphogenetic events and the action of some genes
in formation of the embryo. The work by Johannsen and Butt (1941) is still a very valuable source
for understanding variations in morphogenesis, as are more recent reviews by Anderson (1972a,b),
Jura (1972), Sander et al. (1985), Campos-Ortega and Hartenstein (1985), Panfilio et al. (2006), and
Panfilio (2008). A review of the morphology of embryogenesis in the silkworm, Bombyx mori, has
been provided by Miya (1985), and the early stages of embryogenesis are described for several spe-
cies of fireflies by Kobayashi and Ando (1985).
More details of genetic control of insect embryogenesis are available for Drosophila melano-
gaster than for any other insect, and a timeline for some of the major morphogenetic events may
be helpful (Table 1.1), but one should keep in mind that Drosophila is a fast-developing insect,
and many other insects do not develop so rapidly. Good reviews of the development and genet-
ics of Drosophila are provided by Gehring and Hiromi (1986), Gehring (1987), French (1988),
Table 1.1 Developmental Stages of Drosophila Embryogenesis

Morphological Events (25°C) Hoursa
Stage 1 25 min Cleavage divisions 1 and 2 0:25
Stage 2 40 min Divisions 3–8 occur 1:05
Stage 3 15 min Pole bud formation, division 9 occurs 1:20
Stage 4 50 min Final four divisions, syncytial blastoderm formed, stage 4 ends at beginning 2:10
of cellularization
Stage 5 40 min Cellularization occurs 2:50
Stage 6 10 min Early stages of gastrulation 3:00
Stage 7 10 min Gastrulation complete 3:10
Stage 8 30 min Formation of amnioproctodeal invagination and rapid germ band elongation 3:40
Stage 9 40 min Transient segmentation of mesodermal layer, stomodeal invagination 4:20
Stage 10 60 min Stomodeum invaginates, germ band growth continues 5:20
Stage 11 120 min Growth stage, no major morphogenetic changes, parasegmental furrows 7:20
develop
Stage 12 60 min Germ band shortens 9:20
Stage 13 60 min Germ band shortening complete, head involution begins 10:20
Stage 14 60 min Closure of midgut, dorsal closure 11:20
Stage 15 30 min Gut forms complete tube and encloses yolk sac 13:00
Stage 16 3h Intersegmental grooves evident, shortening of ventral nerve cord 16:00
Stage 17 Stage 17 extends to hatching
Source: Data from Campos-Ortega, J.A. and Hartenstein, V., The Embryonic Development of Drosophila melano-
gaster, Springer-Verlag, Berlin, Germany, 1985, 227pp.
Note: Times and stages probably will be different in other species of insects.
a The time is elapsed time after the egg has been laid in hours.
Embryogenesis 3
Nüsslein-Volhard (1991), Lawrence (1992), and Bate and Martinez-Arias (1993). Melton (1991)
provides a good comparative review of certain aspects of animal development.
1.2 MORPHOGENESIS
1.2.1 Egg, Fertilization, and Zygote Formation
Insect eggs are centrolecithal, which means that the eggs have a central yolk surrounded by a
layer of cytoplasm. The yolk is a nutrient source to be used by the developing embryo. A vitelline
membrane surrounds the peripheral cytoplasm (sometimes called the periplasm), and a protein-
aceous chorion provides a protective cover for the egg contents (Figure 1.1). The cytoplasm is a
layer of variable thickness in eggs of different groups. In some, there is so little cytoplasm that it
is not visually obvious, as for example, in eggs of the Apterygota. The egg nucleus may lie at the
periphery of the egg, on top of the yolk and surrounding cytoplasm, or it may lie in the cytoplasm.
When an egg is laid, the nucleus usually is still in the diploid state. The entry of sperm as the egg
passes down the oviduct of the female often initiates maturation divisions. The first maturation
division divides the chromosomes equally, but the nuclear plasma is divided unequally, resulting
in a large egg nucleus and a small polar body (Figure 1.2). The egg nucleus divides once more to
become the haploid female gamete, with production of another small polar body. The first polar
body may or may not divide again. If it does divide, two more polar bodies are produced; in any
case, polar bodies eventually are reabsorbed into the yolk. The haploid female nucleus usually
migrates toward the center of the egg and unites on the way with the sperm nucleus; the developing
organism is then called the zygote.
Anterior end
Micropyle region
Chorion
Dorsal side
Vitelline membrane
Periplasm
Subcortical layer
Yolk
Figure 1.1 Diagram of egg structure.

4 INSECT PHYSIOLOGY AND BIOCHEMISTRY
Spermatocyte
Oogonium
Polar
body
Figure 1.2 Maturation divisions of oocyte and sperm. Oogonia in the germarial region of an ovary divide by
meiosis to produce an oocyte and a polar body. A second meiotic division, which may not occur
until the oocyte is united with the sperm, produces the final oocyte. The polar bodies are reab-
sorbed as food for the developing oocyte. Spermatocytes in the germarium of the testes give rise
to mature spermatozoa by meiotic divisions. The union of a sperm and egg produces the zygote.
1.2.2 Variations in Zygotic Nucleus Cleavage, Formation

of Energids, and Blastoderm Formation
Zygotic nucleus divisions are influenced by the quantity of yolk and cytoplasm. The division in
eggs with little yolk, such as in the collembolan Tetrodontophora bielanensis (Apterygota), parti-
tions the yolk in a few early divisions (Figure 1.3), but not after the eight-cell stage. In the great
majority of insect groups, the zygotic nuclei divide from the beginning without cleavage of the
yolk and without formation of cell membranes between nuclei. Repeated nuclear divisions produce
(a) (b)
(c) (d)
Figure 1.3 The first few cleavages of the yolk may be complete, as in some Collembola, but complete cleav-
age ceases after a few divisions. (a) The first division is depicted as beginning after the nucleus
has divided by mitosis; (b) division into two cells is illustrated; (c) cleavage into four cells is under-
way, and the four may divide into eight cells, after which, the yolk usually is not cleaved equally
with subsequent nuclear divisions; (d) a ball of cells has formed with yolk that has not been parti-
tioned accumulated in the center (not shown) of the mass of cells. (Stages modified from Jura, C.,
Development of apterygote insects, in J. Counce and C.H. Waddington (eds.), Developmental
Systems: Insects, vol. 1, Academic Press, New York, 1972, pp. 49–94.)
Embryogenesis 5
Energids
Figure 1.4 An example of an egg in which the yolk is not partitioned and cleavage nuclei (energids) are
produced and surrounded by a small amount of cytoplasm. Yolk remains in the interior of the
egg. (Redrawn and modified from Johannsen, O.A. and Butt, F.H., Embryology of Insects and
Myriapods, McGraw-Hill, New York, 1941.)
thousands of nuclei, each surrounded by a small island of cytoplasm. Each nucleus with its island of
cytoplasm is called an energid (Figure 1.4). Energids migrate toward the periphery, and when a few
thousand nuclei have been formed, they distribute themselves in a single layer around the perimeter.
Some energids remain in the yolk and become vitellophages that digest (liquify) the yolk and make
the nutrients available to the developing embryo. Cytoplasmic strands extend from the blastomeres,
as the energids are now usually called, into the yolk as a route for nutrient uptake. Eventually, cell
membranes become complete, the cytoplasmic strands disappear, and the layer of cells is called the
blastoderm (Figure 1.5). There are numerous differences in the way the blastoderm forms and in
subsequent morphogenetic movements among the different groups of insects. A brief summary of
major differences is given in the following text; the reviews and reference works cited in the intro-
duction should be consulted if more details about specific groups are desired.
1.2.2.1 Apterygota
Apterygotes are small, wingless insects with ametabolous development (no metamorphosis and
no major changes in morphology between immatures and adults) and include the orders Protura
(small insects in soil and leaf litter), Collembola (commonly called springtails), Diplura (called
bristletails), Archaeognatha (also called bristletails), and Thysanura (some bristletails, silverfish, and
firebrats) (Romoser and Stoffolano, 1998). Details and variations in development of the Apterygota
have been reviewed by Jura (1972). Even in the Apterygota, the processes of division and cleavage
are not the same in all members of the group. In some, the yolk is cleaved at each division, but in
others, nuclear division occurs without yolk cleavage. Division continues to make many small blas-
tomeres that move toward the periphery of the egg and gradually align themselves in a single layer
around the perimeter of the egg to form the blastoderm. At one pole of the blastoderm, blastomeres
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Now, it may be said that there is no reason for this distinction; but
I claim that there is. And there is no man that can look upon this
crime, horrid as it is, diabolical as it is when committed by the white
man, and not say that such a crime committed by a negro upon a
white woman deserves, in the sense and judgment of the American
people, a different punishment from that inflicted upon the white
man. And yet the very purpose of this section, as I contend, is to
abolish or prevent the execution of laws making a distinction in
regard to the punishment.
But, further, it is said the negro race is weak and feeble; that they
are mere children—“wards of the Government.” In many instances it
might be just and proper to inflict a less punishment upon them for
certain crimes than upon men of intelligence and education, whose
motives may have been worse. It might be better for the community
to control them by milder and gentler means. If the judge sitting
upon the bench of the State court shall, in carrying out the law of the
State, inflict a higher penalty upon the white man than that which
attaches to the freedman, not that I suppose it is ever contemplated
to enforce that, yet it would be equally applicable, and the penalty
would be incurred by the judge in the same manner precisely.
But I proceed to the section I was about to remark upon when the
gentleman interrupted me. The marshals who may be employed to
execute warrants and precepts under this bill, as I have already
remarked, are offered a bribe for the execution of them. It creates
marshals in great numbers, and authorizes commissioners to appoint
almost anybody for that purpose, and it stimulates them by the offer
of a reward not given in the case of the arrest of persons guilty of any
other crime.
It goes further. It authorizes the President, when he is
apprehensive that some crime of that sort may be committed, on
mere suspicion, mere information or statement that it is likely to be
committed, to take any judge from the bench or any marshal from
his office to the place where the crime is apprehended, for the
purpose of more efficiently and speedily carrying out the provisions
of the bill.
The gentleman from Pennsylvania (Mr. Thayer) tells us that it is
very remarkable that it should be claimed that this bill is intended to
create and continue a sort of military despotism over the people
where this law is to be executed. It seems to me nothing is plainer.
Where do we find any laws heretofore passed having no relation to
the negro in which such a provision as this tenth section is to be
found? Generally the marshal seeks by himself to execute this
warrant, and failing, he calls out his POSSE COMITATUS. But this bill
authorizes the use in the first instance of the Army and Navy by the
President for the purpose of executing such writs.
The gentlemen who advocate this bill are great sticklers for
equality, and insist that there shall be no distinction made on
account of race or color.
Why, sir, every provision of this bill carries upon its face the
distinction, and is calculated to perpetuate it forever as long as the
act shall be in force. Where did this measure originate but in the
recognition of the difference between races and colors? Does any one
pretend that this bill is intended to protect white men—to save them
from any wrongs which may be inflicted upon them by the negroes?
Not at all. It is introduced and pressed in the pretended interest of
the black man, and recognizes and virtually declares distinction
between race and color.
I deprecate all these measures because of the implication they

carry upon their face, that the people who have heretofore owned
slaves intend to do them harm. I do not believe it. So far as my
knowledge goes, and so far as my information extends, I believe that
the people who have held the freedmen as slaves will treat them with
more kindness, with more leniency, than those of the North who
make such loud professions of love and affection for them, and are so
anxious to pass these bills. They know their nature; they know their
wants; they know their habits; they have been brought up together;
none of the prejudices and unkind feelings which many in the north
would have toward them.
I do not credit all these stories about the general feeling of hostility
in the South toward the negro. So far as I have heard opinions
expressed upon the subject, and I have conversed with many persons
from that section of the country, they do not blame the negro for
anything that has happened. As a general thing, he was faithful to
them and their interests, until the army reached the place and took
him from them. He has supported their wives and children in the
absence of the husbands and fathers in the armies of the South. He
has done for them what no one else could have done. They recognize
his general good feeling toward them, and are inclined to reciprocate
that feeling toward him.
I believe that is the general feeling of the southern people to-day.
The cases of ill-treatment are exceptional cases. They are like the
cases which have occurred in the northern States where the
unfortunate have been thrown upon our charity.
Take, for instance, the stories of the cruel treatment of the insane
in the State of Massachusetts. They may have been barbarously
confined in the loathsome dens as stated in particular instances; but
is that any evidence of the general ill-will of the people of the State of
Massachusetts toward the insane? Is that any reason why the Federal
arm should be extended to Massachusetts to control and protect the
insane there?
It has also been said that certain paupers in certain States have
been badly used, paupers, too, who were whites. Is that any reason
why we should extend the arm of the Federal Government to those
States to protect the poor who are thrown upon the charity of the
people there?
Sir, we must yield to the altered state of things in this country. We
must trust the people; it is our duty to do so; we cannot do otherwise.
And the sooner we place ourselves in a position where we can win the
confidence of our late enemies, where our counsels will be heeded,
where our advice may be regarded, the sooner will the people of the
whole country be fully reconciled to each other and their changed
relationship; the sooner will all the inhabitants of our country be in
the possession of all the rights and immunities essential to their
prosperity and happiness.
Hon. A. K. McClure on What of the Republic?
Annual Address delivered before the Literary Societies of Dickinson

College, June 26th, 1873.
Gentlemen of the Literary Societies:—What of the Republic?
The trials and triumphs of our free institutions are hackneyed
themes. They are the star attractions of every political conflict. They
furnish a perpetual well-spring of every grade of rhetoric for the
hustings, and partisan organs proclaim with the regularity of the
seasons, the annual perils of free government.
But a different occasion, with widely different opportunities and
duties, has brought us together. The dissembling of the partisan
would be unwelcome, but here truth may be manfully spoken of that
which so profoundly concerns us all. I am called to address young
men who are to rank among the scholars, the teachers, the
statesmen, the scientists of their age. They will be of the class that
must furnish a large proportion of the executives, legislators,
ministers, and instructors of the generation now rapidly crowding us
to the long halt that soon must come. Doubtless, here and there,
some who have been less favored with opportunities, will surpass
them in the race for distinction; but in our free government where
education is proffered to all, and the largest freedom of conviction
and action invites the humblest to honorable preferment, the learned
must bear a conspicuous part in directing the destiny of the nation.
Every one who moulds a thought or inspires a fresh resolve even in
the remotest regions of the Continent, shapes, in some measure, the
sovereign power of the Republic.
The time and the occasion are alike propitious for a dispassionate
review of our political system, and of the political duties which none
can reject and be blameless. Second only to the claims of religion are
the claims of country. Especially should the Christian, whether
teacher or hearer, discharge political duties with fidelity. I do not
mean that the harangue of the partisan should desecrate holy places,
or that men should join in the brawls of pot-house politicians; but I
do mean that a faithful discharge of our duty to free government is
not only consistent with the most exemplary and religious life, but is
a Christian as well as a civil obligation. The government that
maintains liberty of conscience as one of its fundamental principles,
and under which Christianity is recognized as the common law, has
just claims upon the Christian citizen for the vigilant exercise of all
political rights.
If it be true, as is so often confessed around us, that we have
suffered a marked decline in political morality and in our political
administration, let it not be assumed that the defect is in our system
of government, or that the blame lies wholly with those who are
faithless or incompetent. Here no citizen is voiceless, and none can
claim exemption from just responsibility for evils in the body politic.
Ours is, in fact as well as in theory, a government of the people; and
its administration is neither better nor worse than the people
themselves. It was devised by wise and patriotic men, who gave to it
the highest measure of fidelity; and so perfectly and harmoniously is
its framework fashioned, that the sovereign power can always
exercise a salutary control over its own servants. An accidental
mistake of popular judgment, or the perfidy of an executive, or the
enactment of profligate or violent laws, are all held in such
wholesome check by co-ordinate powers, as to enable the supreme
authority of the nation to restrain or correct almost every conceivable
evil.
Until the people as a whole are given over to debauchery the safety
of our free institutions cannot be seriously endangered. True, such a
result might be possible without the demoralization of a majority of
the people, if good citizens surrender their rights, and their duties,
and their government to those who desire to rule in profligacy and
oppression.
If reputable citizens refrain from active participation in our
political conflicts, they voluntarily surrender the safety of their
persons and property, and the good order and well-being of society,
to those who are least fitted for the exercise of authority. When such
results are visible in any of the various branches of our political
system, turn to the true source and place the responsibility where it
justly belongs. Do not blame the thief and the adventurer, for they
are but plying their vocations, and they rob public rather than private
treasure, because men guard the one and do not guard the other.
Good men employ every proper precaution to protect their property
from the lawless. When an injury is done to them individually they
are swift to invoke the avenging arm of justice. They are faithful
guardians of their own homes and treasures against the untitled
spoiler, while they are criminally indifferent to the public wrongs
done by those who, in the enactment and execution of the laws,
directly affect their happiness and prosperity. Do not answer that
politics have become disreputable. Such a declaration is a confession
of guilt. He who utters it becomes his own accuser. If it be true that
our politics, either generally or in any particular municipality or
State, have become disreputable, who must answer for it? Who have
made our politics disreputable? Surely not the disreputable citizens,
for they are a small minority in every community and in every party.
If they have obtained control of political organizations, and thereby
secured their election to responsible trusts, it must have been with
the active or passive approval of the good citizens who hold the
actual power in their own hands. There is not a disgraceful official
shaming the people of this country to-day, who does not owe his
place to the silent assent or positive support of those who justly claim
to be respectable citizens, and who habitually plead their own wrongs
to escape plain and imperative duties. If dishonest or incompetent
appointments have been made, in obedience to the demands of mere
partisans, a just expression of the honest sentiments of better
citizens, made with the manliness that would point to retribution for
such wrongs, would promptly give us a sound practical civil service,
and profligacy and dishonesty would end.
Our Presidents and Governors are not wholly or even mainly
responsible for the low standard of our officials. If good men concede
primary political control to those who wield it for selfish ends, by
refraining from an active discharge of their political duties, and make
the appointing powers dependent for both counsel and support upon
the worst political elements, who is to blame when public sentiment
is outraged by the selection of unworthy men to important public
trusts? The fruits are but the natural, logical results of good citizens
refusing to accept their political duties. There is not a blot on our
body politic to-day that the better elements of the people could not
remove whenever they resolved to do so,—and they will so resolve in
good time, as they have always done in the past. There is not a defect
or deformity in our political administration that they cannot, and will
not correct, by the peaceful expression of their sober convictions, in
the legitimate way pointed out by our free institutions.
You who are destined to be more or less conspicuous among the
teachers of men, should study well this reserved power so
immediately connected with the preservation of our government.
The virtue and intelligence of the people is the sure bulwark of safety
for the Republic. It has been the source of safety in all times past, in
peace and in war, and it is to-day, and will ever continue to be, the
omnipotent power that forbids us to doubt the complete success of
free government. It may, at times, be long suffering and slow to
resent wrongs which grow gradually in strength and diffuse their
poison throughout the land. It may invoke just censure for its
forbearance in seasons of partisan strife. It may long seem lost as a
ruling element of our political system, and may appear to be faithless
to its high and sacred duties. It may be unfelt in its gentler
influences, which should ever be active in maintaining the purity and
dignity of society and government. But if for a season the better
efforts of a free people are not evident to quicken and support public
virtue, it must not be assumed that the source of good influences has
been destroyed, or that public virtue cannot be restored to its just
supremacy. When healthful influences do not come like the dew
drops which glitter in the morning as they revive the harvest of the
earth, they will most surely come in their terrible majesty, as the
tempest comes to purify the atmosphere about us. The miasmas
which arise from material corruption, poison the air we breathe and
disease all physical life within their reach. The poison of political
corruption is no less subtle and destructive in its influences upon
communities and nations. But when either becomes general or
apparently beyond the power of ordinary means of correction, the
angry sweep of the hurricane must perform the work of regeneration.
In our government the mild, but effectual restraints of good men
should be ceaseless in their beneficent offices, but when they fail to
be felt in our public affairs, and evil control has widened and
strengthened itself in departments of power, the storm and the
thunderbolt have to be invoked for the public safety, and our
convulsive but lawful revolutions attest the omnipotence of the
reserved virtue of a faithful and intelligent people.
I am not before you to garner the scars and disjointed columns of
free government. The Republic that has been reared by a century of
patriotic labor and sacrifice, more than covers its wounds with the
noblest achievements ever recorded in man’s struggle for the rights
of man. It is not perfect in its administration or in the exercise of its
vast and responsible powers; but when was it so? when shall it be so?
No human work is perfect. No government in all the past has been
without its misshaped ends; and few, indeed, have survived three
generations without revolution. We must have been more than
mortals, if our history does not present much that we would be glad
to efface. We should be unlike all great peoples of the earth, if we did
not mark the ebb and flow of public virtue, and the consequent
struggles between the good and evil elements of a society in which
freedom is at times debased to license. We have had seasons of war
and of peace. We have had tidal waves of passion, with their
sweeping demoralization. We have enlisted the national pride in the
perilous line of conquest, and vindicated it by the beneficent fruits of
our civilization. We have had the tempest of aggression, and the
profound calm that was the conservator of peace throughout the
world. We have revolutionized the policy of the government through
the bitter conflicts of opposing opinions, and it has been
strengthened by its trials. We have had the fruits of national
struggles transferred to the vanquished, without a shade of violence;
and the extreme power of impeachment has been invoked in the
midst of intensest political strife, and its judgment patriotically
obeyed. We have had fraternal war with its terrible bereavements
and destruction. We have completed the circle of national perils, and
the virtue and intelligence of the people have ever been the safety of
the Republic.
At no previous period of our history have opportunity and duty so
happily united to direct the people of this country to the triumphs
and to the imperfections of our government. We have reached a
healthy calm in our political struggles. The nation has a trusted ruler,
just chosen by an overwhelming vote. The disappointments of
conviction or of ambition have passed away, and all yield cordial
obedience and respect to the lawful authority of the country. The
long-lingering passions of civil war have, for the last time, embittered
our political strife, and must now be consigned to forgetfulness. The
nation is assured of peace. The embers of discord may convulse a
State until justice shall be enthroned over mad partisanship, but
peace and justice are the inexorable purposes of the people, and they
will be obeyed. Sectional hatred, long fanned by political necessities,
is henceforth effaced from our politics, and the unity of a sincere
brotherhood will be the cherished faith of every citizen. We first
conquered rebellion, and now have conquered the bitterness and
estrangement of its discomfiture.
The Vice-President of the insurgent Confederacy is a
Representative in our Congress. One who was first in the field and
last in the Senate in support of rebellion has just died while
representing the government in a diplomatic position of the highest
honor. Another who served the Confederacy in the field and in the
forum, has been one of the constitutional advisers of the national
administration. One of the most brilliant of Confederate warriors
now serves in the United States Senate, and has presided over that
body. The first Lieutenant of Lee was long since honored with
responsible and lucrative official trust, and many of lesser note,
lately our enemies, are discharging important public duties. The war
and its issues are settled forever. Those who were arrayed against
each other in deadly conflict are now friends. The appeal from the
ballot to the sword has been made, and its arbitrament has been
irrevocably ratified by the supreme power of the nation. Each has
won from the other the respect that is ever awarded to brave men,
and the affection that was clouded by the passion that made both
rush to achieve an easy triumph, has returned chastened and
strengthened by our common sacrifices. Our battle-fields will be
memorable as the theatres of the conflicts of the noblest people the
world had to offer to the god of carnage, and the monuments to our
dead, North and South, will be pointed to by succeeding generations
as the proud records of the heroism of the American people.
The overshadowing issues touching the war and its logical results
are now no longer in controversy, and in vain will the unworthy
invoke patriotism to give them unmerited distinction. No supreme
danger can now confront the citizen who desires to correct errors or
abuses of our political system. He who despairs of free institutions
because evils have been tolerated, would have despaired of every
administration the country has ever had, and of every government
the world has ever known. If corruption pervades our institutions to
an alarming extent, let it not be forgotten that it is the natural order
of history repeating itself. It is but the experience of every nation,
and our own experience returning to us, to call into vigorous action
the regenerating power of a patriotic people. We have a supreme
tribunal that is most jealous of its high prerogatives, and that will
wield its authority mercilessly when the opportune season arrives.
We have just emerged from the most impassioned and convulsive
strife of modern history. It called out the highest type of patriotism,
and life and treasure were freely given with the holiest devotion to
the cause of self-government. With it came those of mean ambition,
and of venal purposes, and they could gain power while the unselfish
were devoted to the country’s cause. They could not be dethroned
because there were grave issues which dare not be sacrificed. Such
evils must be borne at times in all governments, rather than destroy
the temple to punish the enemies of public virtue. To whatever extent
these evils exist, they are not the legitimate creation of our free
institutions. They are not the creation of maladministration, nor of
any party. They are the monstrous barnacles spawned by unnatural
war, which clogged the gallant ship of State in her extremity, and had
to be borne into port with her. And now that the battle is ended, and
the issues settled, do not distrust the reserved power of our free
institutions. It will heal the scars of war and efface the stains of
corruption, and present the great Republic to the world surpassing in
grandeur, might and excellence, the sublimest conceptions ever
cherished of human government.
As you come to assume the responsibilities which must be
accepted by the educated citizen, you will be profoundly impressed
with the multiplied dangers which threaten the government. They
will appear not only to be innumerable and likely to defy correction,
but they will seem to be of modern creation. It is common to hear
intelligent political leaders declaim against the moral and intellectual
degeneracy of the times, and especially against the decline in public
morality and statesmanship. They would make it appear that the
people and the government in past times were models of purity and
excellence, while we are unworthy sons of noble sires. Our rulers are
pronounced imbecile, or wholly devoted to selfish ends.
Our law-makers are declared to be reeking with corruption or
blinded by ambition, and greed and faithlessness are held up to the
world as the chief characteristics of our officials. From this painful
picture we turn to the history of those who ruled in the earlier and
what we call the better days of the Republic, and the contrast sinks
us deep in the slough of despair. I am not prepared to say that much
of the complaint against the political degeneracy of the times, and
the standard of our officials, is not just; but in the face of all that can
be charged against the present, I regard it as the very best age this
nation has ever known. The despairing accusations made against our
public servants are not the peculiar creation of the times in which we
live, and the allegation of wide spread demoralization in the body
politic, was no more novel in any of the generations of the past than
it is now. We say nothing of our rulers that was not said of those
whose memory we so sacredly worship. License is one of the chief
penalties, indeed the sole defect of liberty, and it has ever asserted its
prerogatives with tireless industry. It was as irreverent with
Washington as it is with Grant. It racked Jefferson and Jackson, and
it pained and scarred Lincoln and Chase, and their compatriots. It
criticised the campaigns and the heroes of the revolutionary times, as
we criticise the living heroes of our day. It belittled the statesmen of
every epoch in our national progress, just as we belittle those who are
now the guardians of our free institutions. Perhaps we have more
provocation than they had; but if so, they were less charitable, for the
tide of ungenerous criticism and distrust has known no cessation. I
believe we have had seasons when our political system was more free
from blemish than it is now, and that we have had periods when both
government and people maintained a higher standard of excellence
than we can boast of; but it is equally true that we have, in the past,
sounded a depth in the decline of our political administration that
the present age can never reach.
You must soon appear in the active struggles for the perpetuity of
free government, and some of the sealed chapters of the past are
most worthy of your careful study. I would not efface one good
inspiration that you have gathered from the lives and deeds of our
fathers, whose courage and patriotism have survived their
infirmities. Whatever we have from them that is purifying or
elevating, is but the truth of history; and when unborn generations
shall have succeeded us, no age in all the long century of freedom in
the New World, will furnish to them higher standards of heroism and
statesmanship than the defamed and unappreciated times in which
we live. And when the future statesmen shall turn to history for the
most unselfish and enlightened devotion to the Republic, they will
pause over the records we have written, and esteem them the
brightest in all the annals of man’s best efforts for his race. We can
judge of the true standard of our government and people only by a
faithful comparison with the true standard of the men and events
which have passed away. You find widespread distrust of the success
of our political system. It is the favorite theme of every disappointed
ambition, and the vanquished of every important struggle are
tempted, in the bitterness of defeat, to despair of the government.
Would you know whence comes this chronic or spasmodic political
despair? If so, you must turn back over the graves of ages, for it is as
old as free government. Glance at the better days of which we all
have read, and to which modern campaign eloquence is so much
indebted. Do not stop with the approved histories of the fathers of
the Republic. They tell only of the transcendent wisdom and
matchless perfections of those who gave us liberty and ordained
government of the people. Go to the inner temple of truth. Seek that
which was then hidden from the nation, but which in these days of
newspapers and free schools, and steam and lightning, is an open
record so that he who runs may read. Gather up the few public
journals of a century ago, and the rare personal letters and sacred
diaries of the good and wise men whose examples are so earnestly
longed for in the degenerate present, and your despair will be
softened and your indignation at current events will be tempered, as
you learn that our history is steadily repeating itself, and that with all
our many faults, we grow better as we progress.
Do you point to the unfaltering courage and countless sacrifices of
those who gave us freedom, so deeply crimsoned with their blood? I
join you in naming them with reverence, but I must point to their
sons, for whom we have not yet ceased to mourn, who equalled them
in every manly and patriotic attribute. When wealth and luxury were
about us to tempt our people to indifference and ease, the world has
no records of heroism which dim the lustre of the achievements we
have witnessed in the preservation of the liberty our fathers
bequeathed to us. Have corruption and perfidy stained the triumphs
of which we boast? So did corruption and perfidy stain the
revolutionary “times that tried men’s souls.” Do we question the
laurels with which our successful captains have been crowned by a
grateful country? So did our forefathers question the just distinction
of him who was first in war and first in peace, and he had not a
lieutenant who escaped distrust, nor a council of war that was free
from unworthy jealousies and strife. Do politicians and even
statesmen teach the early destruction of our free institutions? It is
the old, old story; “the babbling echo mocks itself.” It distracted the
cabinets of Washington and the elder Adams. It was the tireless
assailant of Jefferson and Madison. It made the Jackson
administration tempestuous. It gave us foreign war under Polk. It
was a teeming fountain of discord under Taylor, Pierce and
Buchanan. It gave us deadly fraternal conflict under Lincoln. Its
dying throes convulsed the nation under Johnson. The promise of
peace, soberly accepted from Grant, was the crown of an unbroken
column of triumphs over the distrust of every age, that was attacking
free government. Do we complain of violent and profligate
legislation? Hamilton, the favorite statesman of Washington, was the
author of laws, enacted in time of peace, which could not have been
enforced in our day even under the necessities and passions of war.
And when the judgment of the nation repealed them, he sought to
overthrow the popular verdict, because he believed that the
government was overthrown. Almost before order began after the
political chaos of the revolution, the intensest struggles were made,
and the most violent enactments urged, for mere partisan control.
Jefferson, the chief apostle of government of the people, did not
always cherish supreme faith in his own work. He trembled at the
tendencies to monarchy, and feared because of “the dupery of which
our countrymen have shown themselves susceptible.” He rescued the
infant Republic from the centralization that was the lingering dregs
of despotism, and unconsciously sowed the seeds which ripened into
States’ rights and nullification under Jackson, and into rebellion
under Lincoln. But for the desperate conflict of opposing convictions
as to the corner-stone of the new structure, Jefferson would have
been more wise and conservative. He was faithful to popular
government in the broadest acceptation of the theory. He summed it
up in his memorable utterance to his neighbors when he returned
from France. He said:—“The will of the majority, the natural law of
every society, is the only sure guardian of the rights of man. Perhaps
even this may sometimes err, but its errors are honest solitary and
short-lived.” Politically speaking, with the patriots and statesmen of
the “better days” of the Republic, their confidence in, or distrust of,
the government, depended much upon whether Hamilton or
Jefferson ruled. Dream of them as we may, they were but men, with
the same ambition, the same love of power, the same infirmities,
which we regard as the peculiar besetting sins of our times. If you
would refresh your store of distrust of all political greatness, study
Jefferson through Burr and Hamilton, or Washington and Hamilton
through Jefferson, or Jackson through Clay and the second Adams,
or Clay and Adams through Jackson and Randolph, and you will
think better of the enlightened and liberal age in which you live.
No error is so common among free people as the tendency to
depreciate the present and all its agencies and achievements.
We all turn with boundless pride to the Senate of Clay, Webster
and Calhoun. In the period of their great conflicts, it was the ablest
legislative tribunal the world has ever furnished. Rome and Greece in
the zenith of their greatness, never gathered such a galaxy of
statesmen. But not until they had passed away did the nation learn to
judge them justly. Like the towering oaks when the tempest sweeps
over the forest, the storm of faction was fiercest among their crowns,
and their struggles of mere ambition, and their infirmities, which
have been kindly forgotten, often made the thoughtless or the
unfaithful despair of our free institutions. Not one of them escaped
detraction or popular reprobation. Not one was exempt from the
grave accusation of shaping the destruction of our nationality, and
yet not one meditated deliberate wrong to the country on which all
reflected so much honor. Calhoun despaired of the Union, because of
the irrepressible antagonism of sectional interests, but he cherished
the sincerest faith in free institutions. But when the dispassionate
historian of the future is brought to the task of recording the most
memorable triumphs of our political system, he will pass over the
great Senate of the last generation, and picture in their just
proportions the grander achievements of the heroes and statesmen
who have been created in our own time. If we could draw aside the
veil that conceals the future from us, and see how our children will
judge the trials and triumphs of the last decade, we would be shamed
at our distrust of ourselves and of the instruments we have employed
to discharge the noblest duties. Our agents came up from among us.
We knew them before they were great, and remembered well their
common inheritance of human defects.—They are not greater than
were men who had lived before them, but the nation has had none in
all the past who could have written their names higher on the scroll
of fame. We knew Lincoln as the uncouth Western campaigner and
advocate; as a man of jest, untutored in the graces, and unschooled
in statesmanship. We know him in the heat and strife of the political
contests which made him our President, and our passions and
prejudices survived his achievements. If his friends, we were brought
face to face with his imperfections, and perhaps complained that he
was unequal to impossibilities. If his enemies, we antagonized his
policy and magnified his errors. We saw him wrestle with the greed
of the place-man, with the ambitious warrior and with the
disappointed statesman. We received his great act of Emancipation
as a part of the mere political policy of his rule, and judged it by the
light of prejudiced partisan convictions.
But how will those of the future judge him? When the hatreds
which attached to his public acts have passed into forgetfulness;
when his infirmities shall have been buried in oblivion, and when all
his master monuments shall stand out in bold relief, made stainless
by the generous offices of time, his name will be linked with devotion
wherever liberty has a worshipper. And it will be measurably so of
those who were his faithful co-laborers. It will be forgotten that they
were at times weak, discordant, irresolute men when they had to
confront problems the solution of which had no precedents in the
world’s history. It will not be conspicuous in the future records of
those great events, that the most learned and experienced member of
his cabinet would have accepted peace by any supportable
compromise, and that one of the most trusted of his constitutional
advisers would have assented to peaceable dismemberment to escape
internecine war. Few will ever know that our eminent Minister of
War was one of those who was least hopeful of the preservation of
the unity of the States, when armed secession made its first trial of
strength with the administration. It will not be recorded how the
surrender of Sumter was gravely discussed to postpone the presence
of actual hostilities, and how the midsummer madness of rebellion
made weakness and discord give way to might and harmony, by the
first gun that sent its unprovoked messenger of death against the flag
and defenders of the Union. It will not be remembered that faction
ran riot in the highest places, and that the struggle for the throne
embittered cabinet councils and estranged eminent statesmen, even
when the artillery of the enemy thundered within sound of the
Capital.
It will not be declared how great captains toyed with armies and
decimated them upon the deadly altar of ambition, and how blighted
hopes of preferment made jangled strife and fruitless campaigns.
Nor will the insidious treason that wounded the cause of free
government in the home of its friends, blot the future pages of our
history in the just proportions in which the living felt and knew it. It
will be told that in the hour of greatest peril, the administration was
criticised, and the constitution and laws expounded, with supreme
ability and boldness, while the meaner struggles of the cowardly and
faithless will be effaced with the passions of the times that created
them. And it is best that these defects of greatness should slumber
with mortality. Not only the heroes and rulers, but the
philanthropists as well, of all nations and ages, have had no
exemption from the frailties which are colossal when in actual view.
That we have been no better than we have seen ourselves, does not
prove that we are a degenerate people. On the contrary, it teaches
how much of good and great achievement may be hoped for with all
the imperfections we see about us. In our unexampled struggle, when
faction, and corruption, and faithlessness had done their worst, a
regenerated nationality, saved to perfected justice, liberty and law,
was the rich fruits of the patriotic efforts of the people and their
trusted but fallible leaders. There is the ineffaceable record we have
written for history, and it will be pointed to as the sublimest tribute
the world has given to the theory of self-government. The many
grievous errors and bitter jealousies of the conflict which weakened
and endangered the cause; the venality that grew in hideous
strength, while higher and holier cares gave it safety; the
incompetency that grasped place on the tidal waves of devotion to
country, and the widespread political evils which still linger as
sorrowful legacies among us, will in the fulness of time be healed and
forgotten, and only the grand consummation will be memorable.
This generous judgment of the virtue and intelligence of the people,
that corrects the varying efforts and successes of political
prostitution; that pardons the defects of those who are faithful in
purpose, and without which the greatest deeds would go down to
posterity scarred and deformed, is the glass through which all must
read of the noblest triumphs of men.
Our Republic stands alone in the whole records of civil
government. In its theory, in its complete organization, and in its
administration, it is wholly exceptional. We talk thoughtlessly of the
overthrow of the old Republics, and the weak or disappointed turn to
history for the evidence of our destruction. It is true that Republics
which have been mighty among the powers of the earth have
crumbled into hopeless decay, and that the shifting sands of time
have left desolate places where once were omnipotence and
grandeur. Rome made her almost boundless conquests under the
banner of the Republic, and a sister Republic was her rival in
greatness and splendor. They are traced obscurely on the pages of
history as governments of the people. Rome became mistress of the
world. Her triumphal arches of costliest art recorded her many
victories. Her temples of surpassing elegance, her colossal and
exquisite statues of her chieftains, her imposing columns dedicated
to her invincible soldiery, and her apparently rapid progress toward
a beneficent civilization, give the story of the devotion and heroism
of her citizens. But Rome never was a free representative
government. What is called her Republic was but a series of surging
plebeian and patrician revolutions, of Tribunes, Consuls and
Dictators, with seasons of marvelous prowess under the desperate
lead of as marvelous ambition. The tranquillity, the safety, and the
inspiration of a government of liberty and law, are not to be found in
all the thousand years of Roman greatness. The lust of empire was
the ruling passion in the ancient Republics. Hannibal reflected the
supreme sentiment of Carthage when he bowed at the altar and
swore eternal hostility to Rome; and Cato, the Censor, as faithfully
spoke for Rome when he declared to an approving Senate
—“Carthago delenda!” Such was the mission of what history hands
down to us as the great free governments of the ancients. Despotism
was the forerunner of corruption, and the proudest eras they knew
were but hastening them to inevitable destruction.
The imperial purple soon followed in Rome, as a debauched people
were prepared to accept in form what they had long accepted with
the mockery of freedom. Rulers and subjects, noble and ignoble,
church and state, made common cause to precipitate her decay. At
last the columns of the barbarian clouded her valleys. The rude hosts
of Attila, the “Scourge of God,” swarmed upon her, and their battle-
axes smote the demoralized warriors of the tottering empire. The
Goth and the Vandal jostled each other from the degraded sceptre
they had conquered, and Rome was left widowed in her ruins. And
Carthage!—she too had reared a great government by spoliation, and
called it a Republic. It was the creation of ambition and conquest.
Her great chieftain swept over the Pyrenees and the Alps with his
victorious legions, and even made the gates of the Eternal City
tremble before the impetuous advance of the Carthaginians. But
Carthage never was free until the cormorant and the bittern
possessed it, and the God of nations had “stretched out upon it the
line of confusion and the stones of emptiness.” Conqueror and
conquered are blotted from the list of the nations of the earth. We
read of the Grecian Republic; but it was a libel upon free
government. Her so-called free institutions consisted of a loose,
discordant confederation of independent States, where despotism
ruled in the name of liberty. Sparta has made romance pale before
the achievements of her sons, but her triumphs were not of peace,
nor were they for free government. Athens abolished royalty more
than a thousand years before the Christian era, and made Athenian
history most thrilling and instructive, but her citizens were strangers
to freedom. The most sanguinary wars with sister States, domestic
convulsions almost without cessation, and the grinding oppression of
caste, were the chief offerings of the government to its subjects.
Solon restored her laws to some measure of justice, only to be cast
aside for the usurper. Greece yet has a name among the nations of
the world, but her sceptre for which the mightiest once warred to
enslave her people under the banner of the Republic, has long since
been unfelt in shaping the destiny of mankind. Thus did Rome and
Carthage and Greece fade from the zenith of distinction and power,
before constitutional government of the people had been born
among men. To-day there is not an established sister Republic that
equals our single Commonwealth in population. Spain, France and
Mexico have in turn worshiped Emperors, Kings, Dictators and
popular Presidents. Yesterday they were reckoned Republics. What
they have been made to-day, or what they will be made to-morrow, is
uncertain and unimportant. They are not now, and never have been,
Republics save in name, and never can be free governments until
their people are transformed into law-creating and law-abiding
communities. With them monarchy is a refuge from the license they
miscall liberty, and despotism is peace. Switzerland is called a
Republic. She points to her acknowledged independence four
hundred years ago, but not until the middle of the present century
did the Republic of the Alps find tranquillity in a constitutional
government that inaugurated the liberty of law. Away on a rugged
mountain-top in Italy, is the only Republic that has maintained
popular government among the States of Europe. For more than
fourteen hundred years a handful of isolated people, the followers of
a Dalmatian hermit priest, have given the world an example of
unsullied freedom. Through all the mutations, and revolutions, and
relinings of the maps of Europe, the little territory of San Marino has
been sacredly respected. Her less than ten thousand people have
prospered without interruption; and civil commotions and foreign
disputes or conflicts have been unknown among them. She has had
no wealth to tempt the spoiler; no commerce or teeming valleys to
invite conquest; no wars to breed dictators; no surplus revenues to
corrupt her officials; and in patient and frugal industry her citizens
have enjoyed the national felicity of having no history. They have had
no trials and no triumphs, and have made civilization better only by
the banner of peace they have worshipped through all the
convulsions and bloody strife of many centuries.
The world has but one Republic that has illustrated constitutional
freedom in all its beneficence, power and grandeur, and that is our
own priceless inheritance. As a government, our Republic has alone
been capable of, and faithful to, representative free institutions, with
equal rights, equal justice, and equal laws for every condition of our
fellows. All the nations of the past furnish no history that can
logically repeat itself in our advancement or decline. Created through
the severest trials and sacrifices; maintained through foreign and
civil war with unexampled devotion; faithful to law as the offspring
and safety of liberty; progressive in all that ennobles our peaceful
industry, and cherishing enlightened and liberal Christian
civilization as the trust and pride of our citizens, for our government
of the people, none but itself can be its parallel.
In what are called free governments of antiquity, we search in vain
for constitutional freedom, or that liberty that subordinates passion
and license to law. The refuge from the constant perils of an
unrestrained Democracy was always found in despotism, and when
absolutism became intolerable, the tide of passion would surge back
to Democracy. The people, in mass councils, would rule Consuls,
Presidents and Generals, but it was fruitful only of chaos and
revolution. The victorious chieftain and the illustrious philosopher
would be honored with thanksgivings to the gods for their
achievements, and their banishment or death would next be
demanded by the same supreme tribunal. Grand temples and
columns and triumphal arches would be erected to commemorate
the victories of the dominant power, and the returning waves of
revolution would decree the actors and their monuments to
destruction. Ambitious demagogues prostituted such mockeries of
government to the basest purposes. The Olympic games of Greece
became the mere instruments of unscrupulous leaders to lure the
people, in the name of freedom, to oppression and degradation, and
the wealth of Rome was lavishly employed to corrupt the source of
popular power, and spread demoralization throughout the Republic.
The debauched citizens and soldiers were inflamed by cunning and
corrupt devices, against the purest and most eminent of the sincere
defenders of liberty; and the vengeance of the infuriated mob,
usurping the supreme power of the State, would doom to exile or to
death, honest Romans who struggled for Roman freedom. Cato, the
younger, Tribune of the people, and faithful to his country, took his
own life to escape the reprobation of a polluted sovereignty. Cicero
was Consul of the people, made so by his triumph over Cæsar. But
the same people who worshipped him and to whose honor and
prosperity he was devoted, banished him in disgrace, confiscated his
wealth and devastated his home. Again he was recalled through a
triumphal ovation, and again proscribed by the triumvirs and
murdered by the soldiers of Antony. The Grecian Republic banished
“Aristides the just,” and Demosthenes, the first orator of the world,
who withstood the temptations of Macedonian wealth, was fined,
exiled and his death decreed. He saved his country the shame of his
murder by suicide. Miltiades won the plaudits of Greece for his
victories, only to die in prison of wounds received in fighting her
battles. Themistocles, orator, statesman and chieftain, was banished
and died in exile. Pericles, once master of Athens, and who gave the
world the highest attainments in Grecian arts, was deposed from
military and civil authority by the people he had honored. Socrates,
immortal teacher of Grecian philosophy, soldier and senator, and
one of the most shining examples of public virtue, was ostracised and
condemned and drank the fatal hemlock. The Republic of Carthage
gave the ancients their greatest general, and as chief magistrate, he

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Insect physiology and biochemistry

Third Edition James L. Nation

Glutamine : biochemistry, physiology, and clinical

Reviews of Physiology Biochemistry and Pharmacology

Reviews of Physiology Biochemistry and Pharmacology

Handbook of Biochemistry and Molecular Biology Roger L.

Chemistry for today : general, organic, and

Tickets to Paradise D L James James D L Daniel Lewis

The American Psychiatric Association Publishing

Seeley's essentials of anatomy & physiology Tenth

Boca Raton London New York

CRC Press is an imprint of the

No claim to original U.S. Government works

International Standard Book Number-13: 978-1-4822-4760-2 (eBook - PDF)

3.3 Ability of Insects to Self-Select Nutritional Components..................................................... 77

6.3 Molecular Basis for the Circadian Clock............................................................................. 183

8.4 Hormones Controlling Carbohydrate Metabolism.............................................................. 230

9.11.7 Vasopressin-Like Peptide (Locust F2 Peptide)....................................................... 279

11.6 Sound Production: Tymbal and Stridulatory Muscle........................................................... 327

13.5 Chemoreceptors................................................................................................................... 368

15.5 Hemolymph.......................................................................................................................... 420

17.4 Ventilation and Diffusion of Gases within the System........................................................466

19.4 Active Space Concept.......................................................................................................... 528

20.8 Male Reproductive System.................................................................................................. 577

James L. Nation, Sr.

Table 1.1 Developmental Stages of Drosophila Embryogenesis

1.2.1 Egg, Fertilization, and Zygote Formation

Figure 1.1 Diagram of egg structure.

1.2.2 Variations in Zygotic Nucleus Cleavage, Formation

I deprecate all these measures because of the implication they

Annual Address delivered before the Literary Societies of Dickinson

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Figure 1.1 Diagram of egg structure.

1.2.2 Variations in Zygotic Nucleus Cleavage, Formation