Insect Physiology and Biochemistry, Second Edition

540 Index regenerative cells in the midgut of, 37f Sensilla campaniformia, 299 spiracular opening of, 390f Sensilla chaetica, 299 Scarab beetle. See Cotinus mutabilis Sensilla coeloconica, 299, 306 Scarabaeid Sensilla placodea, 299 digestive system morphology and physiology, 57 Sensilla styloconica, 299, 306 proline as a metabolic fuel for, 193 Sensilla trichodea, 299, 306, 310 Scarabaeidae, serine proteinases of, 46 Sensory fatigue, 472 Scarabaeus zambesianus (dung beetles), use of plane- Sensory hairs, structure of, 299–300 Sensory neurons, 206, 218–219, 296 polarized light by, 333 Schiff base, 326 chemoreceptor, 309 Schistocerca americana Sensory organs active ventilation of, 400 complex chordotonal, 301–305 flight muscle power of, 290 hygroreceptors, 305–306 Schistocerca gregaria (desert locust) simple chordotonal, 305 acid-base balance in, 433 thermoreceptors, 305–306 ammonia secretion of, 434 Sensory receptors, 515 breathing motor pattern in, 224–225 Septate junctions, 96, 97f DGC patterns in, 402 SER, 98 digestive system of, 55 Serine, 45 FaRPs in, 227 Serine endoprotease, 500 feeding deterrents for, 85 Serosa, 9–10 glucose conversion in, 181 Serotonin, 346, 424, 429. See also glutamate receptors in, 263 hemolymph buffering of, 358 5-hydroxytryptamine hemolymph pH of, 357 Serpins, 370, 376 ileum of, 427 Serratia marcescens, 378 jumping muscles in, 270–271 Sesquiterpenoid compounds, 142 molting fluid secretion by, 100 Sex chromosomes, ratio of to autosomes, 502 proline metabolism by, 197 Sex combs reduced gene, 17 tracheae of, 396 Sex pheromones, 448–449, 452 tympanum of, 304 Sex-determining genes, 502 use of plane-polarized light by, 332 Shaker potassium channel, 239 vitamin requirements of, 78 Sheath cells, 299 water balance during flight of, 404 Shielding pigment distribution, 318 Schistocerca nitens, hemolymph pH of, 357 Short germ band, 9 Schistocerca sp., active ventilation of, 399 Sialis lutaria, ammonia excretion of, 434 Sciara coprophila Lintner, tracheole formation in, Sieve plates, 389 Signal processing, pheromone, 464–466 391–392 Signal transduction, 458 Sclerotization, 92, 94–95 Silkmoth, 126. See also Antheraea polyphemus Silkmoths. See also Hyalophora cecropia cuticle, 103–105 hormonal control of, 105 crystalline tracts in eyes of, 320 model of, 106f cuticle of, 114–115 Scolopale cells, 300–301 Silkworm. See Bombyx mori Scolopidium, 300–301 Silverfish. See Thysanura Scotophase, 132 Simple diffusion, 397 Scotopic eyes, 318 Siphonaptera Seasonal biological clocks, 168–169 developmental hormones in, 128 Second messengers, 133, 146 panoistic ovaries in, 486 Secondary plant substances, 85 stridulation of, 273 Secondary sensory neurons (type II), 296 Siricidae, carbohydrate digesting enzymes of, 45 Secondary tracheae, 396 SIT, 37 Secretory vesicles, 36 Sitona cylindricollis (sweet clover weevil), feeding Segment polarity genes, 16 Segmentation genes, 12, 15–16 deterrents for, 85 Semiochemicals, 448 Skeletal muscle, 260 chemical characteristics of, 453–456 Slo gene, 239–240 classes of, 448–449 Sloppy paired genes, 16 detection of, 457 Slow axons, 262 Semper’s cells, 319 Slow neuron, 260 Sensilla, 297, 456–457 Slow potentials, 235 classification scheme for, 298t Slowpoke gene. See slo gene pheromone signal processing and, 464–465 Smooth endoplasmic reticulum. See SER Sensilla basiconica, 299, 306, 309 snail gene, 15 snake gene, 15

Index 541 Sodium, 79, 345, 359 Storage proteins, 170, 360 activation, 242–244 Stretch receptors, secretion of PTTH and, 131–132 channels, 239 Striated border, 41 inactivation, 244 Stride indicator, 269 ion secretion of hematophagous insects, 429 Stridulation, 273–274 Stridulatory muscles, physiology of, 273–274 Solid vs. liquid food, 31–32 Strychnine-HCL, 85 Soma, 217, 234 Subalar muscles, 281 Somata, 215 Subesophageal ganglion, 206, 212 Sorbitol, 171–172 Sound production, 411 PBAN production in, 467 Subgenual organs, 301–302 effect of air sacs on, 393 Substance P, 227 tymbal muscles, 272 Sucrose, 85 South African long-horned beetle. See Phrymeta spinator Sugars, 85 Sparing sterol, 76 Summation, 262 Speophyes lucidulus, thermoreceptors of, 306 Supella longipalpa (brown-banded cockroaches), self- Sperm apyrene and eupyrene of Lepidoptera, 500 selection of nutritional components by, 71 maturation divisions of, 4f Supernumerary larvae, 125 survival of in female genital tract, 500 Superposition, 318 transfer of, 501 swallow gene, 13 Spermatheca, 484 Sympetrum danguineum, flight of, 285 Spermatogonia, 499 Sympetrum sanguineum Spermatophore, 500 Spermatophorins, 501 clap and fling wing motion of, 286 Spermatozoa, 498 flight of, 285 Spherulocytes, 348–349 Synapses, 246–248 Sphingid moths acetylcholine-mediated, 248–250 dorsal aorta of, 342f electric transmission across, 250 hovering flight of, 289 Synchronous muscles, 260, 263–265 Spike potential, 236, 242–244. See also action potential stridulation and, 273 Spiracles, 386 Syncytial blastoderm, Drosphila melanogaster, 8 structure and function of, 389–391 Synomones, 449 Spodoptera exempta, photoreceptors of, 329 Spodoptera frugiperda (fall armyworm), 142 T female, vitellogenin synthesis in, 493 Mas-AT isolated from, 228 Tabanid flies. See also specific tabanid flies Spodoptera littoralis (cotton leafworm) male, visual acuity of, 335 inactivation of molting hormone in, 141f male, MGC of, 465 Tabanidae, corneal layering in eyes of, 321 PBAN and pheromone biosynthesis in, 467 Tachinaephagus zealandicus, muscle of, 258f Springtails. See Collembola Tactile hairs, 297 Spruce budworm. See Choristoneura fumiferana SR, 256–257f structure of, 299–300 synchronous muscles and, 263–264 Taenidia, 388 Stable fly. See Stomoxys calcitrans Taeniopoda eques (western lubber grasshopper) Standing gradient process, 423 staufen gene, 13 acid-base balance in, 433 influence of on posterior development, 14 DGC cycles in, 402 Steady-state aerodynamics, 286 hemolymph pH of, 358 Steinernema feltiae, 378 TAG, 214, 467 Stemmata, 316–317, 324 tailless gene, 14–15 Stereochemical theory, 311 takeout gene. See to gene Sterile insect technique. See SIT Tanning, 103–105. See also sclerotization Sternum, 514 Tannins, 40 Steroid hormones, 154–155 Tapetum, 320–321 Sterols, 76–77 tarazu gene, 16 Stimulus-receptor excitation coupling, 311 Techykinins, 227 Stomatograstric nervous system, 212 Tela polyphenus, chemoreceptors of, 310 Stomoxys calcitrans (stable fly) Teleogryllus commodus cuticle mineralization of, 118 female, role of JH III in ovarian development of, 489 digestive system of, 59 prothoracic leg tracheae of, 412 proteolytic enzymes of, 46 Telotrophic ovarioles, 486–487 Storage excretion, 418 Telson, genes required in, 15 Tenebrio molitor (mealworm). See also Coleoptera active ventilation of, 398 cardioactive peptides in, 229 cryptonephridial system of, 438–439

542 Index cuticle proteins of, 115 Thoracic ganglia, 206 developmental hormones in, 128 Thoracic ganglion, 212–213f digestive system morphology and physiology, 57 Thoracic muscular pumping, 398–399 diuretic hormone of, 431–432 Thoracic structure, 280 female, vitellogenin synthesis in, 492 Thorax, 514 glutamate receptors in, 263 Threonine, 73 heart of, 344 Thysanura immunity in, 377 midgut chitinase of, 44 embryological development of, 2, 6–7 mineral requirements of, 80 excretory system of, 419 telotrophic ovary in, 487 ommatidia of, 317 vitamin requirements of, 78 panoistic ovaries in, 486 water absorption of cuticular layer of, 118 Tight junctions, 97 Tenebrio spp., carbohydrate needs of, 75 Timer gene concept, 154 Tenebrionid beetles. See also Cryptoflossus verrucosa; Timing genes, 169 Tineola bisselliella (clothes moth) Onymacris unguicularis digestive system of, 60 cuticle lipid layer of, 116 redox potential in the midgut of, 51 DGC patterns in, 402 Tipulidae, dorsal vessel of, 341 water absorption of cuticular layer of, 118 Tissue Tenethredinidae, blastokinesis, 11 hemocytopoietic, 351 Tenodora australasia (praying mantis), visual acuity of, 335 regulation of levels of juvenile hormone in, 144–147 Tenodora sinensis (praying mantis) regulation of levels of PTTH in, 132 crops of, 32 tracheal supply to, 394–396 digestive system of, 55 use of in assays of ecdysteroids, 140 TEP, 51 Tissue-specific factors, 153 Tephritid fruit flies. See also Anastrepha suspensa; TLC, 140 TMOF, 47–48, 490 Bactrocera tryoni TnC, 266 carbohydrate needs of, 75 TnI, 266 developmental hormones in, 128–129 TnT, 266 digestive system of, 59 to gene, 270 Tergosternal muscle, 396 Tobacco budworm. See Heliothis virescens (tobacco Tergum, 514 Terminal abdominal ganglia. See TAG budworm) Termites. See also Cubitermes fungifaber; Isoptera; Tobacco hornworm. See Manduca sexta Token stimuli, termination of diapause and, 165 Macrotermitinae; Rhinotermitidae; Termitidae Tokus, 394 antimicrobial peptides, 372 Toll pathway, synthesis of antimicrobial peptides and, carbohydrate digesting enzymes of, 45 cyclic release of CO2 in, 403 372–375 digestive system morphology and physiology, 55–56 Toll protein, 15 hindgut in, 36 Tomato hornworm. See Manduca quinquemaculata pH of hindgut of, 54 Tomato moth. See Lacanobia oleracea synthesis of polyunsaturated fats by, 78 Tomocerus minor, peritrophic matrix of, 43 TEP of, 51 Tonic receptors, 131, 296. See also stretch receptors trail hormones of, 452 Tonofibrillae, 259 Termitidae, digestive system of, 55 Tormogen, 299 Territory-marking pheromones, 452 Toroid, 465 Tetranchus urticae (two-spotted spider mites), essential torso gene, 15 torsolike gene, 15 amino acids for, 74t–75 Tracheae, 386 Tetrodontophora bielanensis, zygotic nucleus cleavage active ventilation of, 397–400 of, 4 adaptations of to supply flight muscles, 394 Tettigetta josei, auditory neurons of, 302 diffusion in, 397 Tettigoniidae. See also grasshoppers; katydids molting of, 394 structure of, 387–389 subgenual organs of, 301 Tracheal epithelium, 391 Thecogen, 299 Tracheal gills, 408 Thermobia domestica (thysanuran firebrat) Tracheal system closed, 408–410 female, ovarian development and oogenesis of, 492 connections of to Malpighian tubules, 419 panoistic ovary of, 486 development of, 19 water absorption of cuticular layer of, 118 nonrespiratory functions of, 411–412 Thermoreceptors, 305–306 primary tracheal supply to wing muscles, 396f Thermoregulation, role of circulation in, 355 structure of, 387–394 Thiamine, 78 Thin layer chromatography. See TLC Third axillary muscles, 281

Index 543 Tracheoles, 386 Twisting, control of, 289–290 development of, 391–392 Two-spotted spider mites. See Tetranchus urticae diffusion from to mitochondria, 400–401 Two-striped grasshopper. See Melanoplus bivattatus structure of, 387–389 Tymbal muscles, 264–265, 272f Trail-following pheromones, 452 morphology and physiology of, 272–273 false, 473 Tympanal organs, 302–304 Transcellular resistance, 429 effect of air sacs on, 393 Transcription, 150 Tympanum, 297, 304 Type I PM, 41–42 factors, Bicoid protein, 14 products, 153 in Lepidoptera, 59 Transducin, 326 Type II PM, 41–42 Transepithelial potential. See TEP Transition temperature, 117–118 in Diptera, 58 Transmembrane proteins, 238 opsin, 316 U Transport, mean cost of, 269 Trehalase, 182 UDP-glucose-glycogen transglycosylase. See glycogen Trehalose, 171–172, 180 synthetase supply of to the brain and ganglia, 215 synthesis of from glucose, 182f UDP-glucosyl transferase, 141–142 Triacylglycerols, 45, 165, 360 Ultrabithorax gene, 17 Triad junctions, 257 Ultraspiracle. See USP Triatoma infestans Uniramia, 512 diuresis of, 430–431 Univoltine life cycle, obligatory diapause of insects with, thermoreceptors of, 306 Triatoma protracta, female, vitellogenin synthesis in, 492 162 Triatoma sp., antimicrobial peptides, 372 Unsteady forces, 286 Tribolium Unsteady lift, 286 carnitine requirements of, 78 Upstroke, 281, 288 homeotic genes in, 16 Uric acid, 361, 433 Tribolium confusum (confused flour beetles) food utilization of, 83–84f method, 82 self-selection of nutritional components by, 71 synthesis and excretion of, 435–437 Tribolium spp., carbohydrate needs of, 75 Uricase, 437 Trichogen, 299 Uricotelic insects, 437 Trichopria columbiana, use of aquatic plants as an air Uridine 5’-diphosphate-glucosyl transferase. See UDP- source, 407–408 glucosyl transferase Trichoptera Urine adult diapause of, 168 osmality of, 424 flight muscles of, 264 primary, formation of in Malpighian tubules, 421 midgut pH of, 54 proton pump and formation of, 422–424 Tricoplusia ni, male, MGC of, 465 Urotensin I, 431 Trilobita, 511 USP, 171 Triosephosphate isomerase, 185 Utetheisa ornatrix, male, sex pheromone of, 449 Tritocerebrum, 205–206, 208, 212 UV-sensitive photoreceptors, 328–329 Trityrosine, 114 Trophocytes, 487. See also nurse cells V Tropical cockroach. See Xestoblatta hamata Tropomyosin, 265 Valine, 73 Tropomyosin binding subunit. See TnT Valois gene, influence of on posterior development, 14 Troponin, 265 Vanessa atalanta (Red Admiral), photoreceptors of, 329 Trypanosoma rangeli, 377 Vanessa indica, vision and behavior of, 330 Trypsin, 45–46, 51 Varroa destructor, 378 early, 47 Vas deferens, 498 late, 48 vasa gene, influence of on posterior development, 14 Trypsin modulating oostatic factor. See TMOF Vasopressin, 228 Tryptophan, 73 Vasopressin-like peptide. See locust F2 peptide Tsetse flies. See also Glossina sp. Ventilation gas exchange in, 405 proline as a metabolic fuel for, 193 active, 397–400 Tubular muscle, 264 discontinuous cycle of, 401 Turnip moth. See Agrotis segetum Ventral diaphragm, 343 twist gene, 15 Ventral ganglia, 207, 212–215 Ventral nerve cords, 102, 207 tracheal supply to, 394 Ventral sensory neuropil, 214 Vertebrate steroid hormones, 154–155

544 Index Vespa crabro Weigert, Carl, 206 activate ventilation of, 397 Western lubber grasshopper. See Taeniopoda eques active ventilation of, 399 Wheast, 72 Wholfahrtia vigil, ammonia secretion of, 434 Visceral muscles, 274 Willow beetle. See Chrysomela aeneicollis Visceral organs, tracheal supply to, 394 Wing hearts, 346 Vision, insect, chemistry of, 325–326 Wing muscles. See also flight Visual acuity, 334–335 Visual cascade, 326–327 tracheal supply to, 396f wingless gene, 16 regulation of, 328 Wings, 515. See also flight Visual pigment, 316 Vitamin A, 78, 331 aerodynamics of lift and drag forces, 285–289 Vitamin B12, 78 beat frequency, 265, 284t Vitamin BT, 78. See also carnitine control of pitch and twisting of, 289–290 Vitamin E, 79 hemolymph circulation in, 354–355 Vitamin K, 79 hinges, 280 Vitamins, 78–79 loading, 265 Vitelline membrane, 3 metabolic activity of muscles of, 291 movement of in dragonflies and damselflies, 284–285 formation of, 497 strokes of, 281 Vitellins, 493, 496 Woodwasps Vitellogenic stage, 490 carbohydrate digesting enzymes of, 45 Vitellogenins, 488 digestive system of, 58 Worker jelly, 75 biochemical characteristics of, 493–495 genetic controls of, 494 X sequestration of by oocytes, 495–497 Vitellophages, 4, 8 Xanthine dehydrogenase, 436 Voltage clamp, 245 Xestoblatta hamata (tropical cockroach), paternal Voltage gated channels, 238 investment behavior of, 437 W Xyleborus ferrugineus, cholesterol utilization by, 76 Wake capture, 288 Y Walking Yellow fever mosquito. See Anopheles gambiae; adaptations for, 269–270 mosquitoes motor program to control, 223–224 Wasps. See also Vespa crabro Yolk activate ventilation of, 397 body, 496 Water cleavages of, 5f absorption of through cuticular layer, 118 proteins balance of during flight, 404–405 biochemical characteristics of, 493–495 conservation, 36 genetic controls of, 494–495 excretion of, 31 sequestration of by oocytes, 495–497 Water homeostasis, 429–430 Water soluble vitamins, 78 Z Wax blooms, 94, 116 Wax channels, 95 Z lines, 258 Wax esters, 116 Zigzag flight behavior, 462–464 Wax layer, 116. See also chemical waxes Zinc, 80 Wax moths. See Galleria mellonella Zinc fingers, 151 Websites Zones of maturation, spermatid formation in, 499 clap and fling wing motion, 286 Zootermopsis angusticollis, synthesis of polyunsaturated cuticular proteins, 113 dragonfly flight, 285 fats by, 78 Drosophila courtship, 498 Zygotes, 3 flea jumping, 271 genome projects, 379 nucleus cleavage variations, 4–9 hemolymph flow, 355 Zygotic genes, 12 hovering flight, 289 Zymogen, 43, 100, 370 tympanum deflection, 304

Coxa orax Trochanter A P Femur Tibia Tarsus Figure 1.12  An imaginal leg disc of Drosophila melanogaster showing regions of the disk that will give rise to parts of the leg and some other parts of the body during pupation. (From Bryant, 1993. With permission.) Figure 1.13  A basic view of the growth-pro- moting glycoprotein (IDGF-DRW protein 3-D structure) from Diaprepes root weevil. The struc- ture was predicted with ROBETTA, a full-chain protein prediction server. The beta barrel motif is shown in yellow in the center. The conserved resi- dues in green (in space fill) are apparently essen- tial to maintain the barrel folding as predicted by interface alanine scanning. Residues are Gly109, Gly110, Asp152, Gly153, Leu218, Asp242, Lys295, Gly412, and Asp421. The residue at position 159 is Glu (in brown space fill), which is replaced by Gln in other known IDGY proteins. Structure key: alpha helix (pink), beta strand (yellow), turn (blue), and other (gray). (From Huang et al., 2006, by W.B. Hunter. With permission.)

Figure 4.17  Color phases of the beetle Cryptoglossa verrucosa (Le Conte) (Coleoptera: Tenebrionidae) in response to humidity. The beetle is dark, nearly black (right) at higher humidity and blue (left) at low humidity. Wax filaments secreted during low humidity by tubercles on the elytra disperse the light and give the beetles a beautiful blue color. (The photos were taken in Bahia de Kino, Sonora, Mexico, and are courtesy of J. Nathan- iel Holland, Department of Ecology and Evolutionary Biology, University of Arizona, Tucson.) A B Figure 6.1  A: An adult monarch butterfly taking nectar from a flower on a sunny day in mid-January at the El Rosario colony of diapausing and overwintering monarchs in the mountains near Mexico City. B: Several diapausing monarchs feeding from a flower in the Sierra Chincua colony in the mountains near Mexico City on a warm day in mid-January. (Photos courtesy of Rochelle Carlson Nation.)

Figure 11.6  A male tabanid fly hovering in flight waiting for a female to fly by so that it can give chase. (Photo courtesy of Jerry Butler.) Figure 13.4  A tabanid fly with a dark band across the lower part of the compound eyes. (Photo courtesy of Jerry Butler, professor emeritus, Dept. of Entomology and Neu , University of Florida, Gainesville, FL.)

G+ Lys-PGN? Osiris Fungus PGRP-SA ? nec Hades ? psh clip spz G– spz spz PGRP-LC DAP-PGN Toll ? PGRP-LE Toll pathway Hemolymph IMD pathway Fat body cell Figure 15.3  The recognition of microbial infection in Drosophila leading to activation of the Toll and IMD pathways. Gram (+) bacteria are recognized by PGRP-SA, a peptidoglycan recognition protein that binds the lysine-type peptidoglycan of Gram (+) bacteria. Osiris is another peptidoglycan recognition protein that participates in the initial recognition reaction. A proteolytic kinase cascade is initiated that finally removes a clip domain from the dimer Späetzle, the ligand for Toll, a transmembrane protein in the fat body cell mem- brane. Fungal invasion also elicits recognition proteins, one of which may be Hades. A proteolytic cascade is activated and one of the kinases, Persephone, proteolytically removes the clip domain from Späetzle, which then binds Toll. Toll dimerizes, perhaps as a result of binding spz, and then binds two molecules of spz. Gram (–) bacteria, which have diaminopimelic acid (DAP)-type peptidoglycan in the outer coat, is recognized and bound by PGRP-LDC and possibly PGRP-LE. The immune deficiency (IMD) pathway is activated. (From Leclerc and Reichhart, 2004. With permission.)

TIRspz spzTIR TIR TIR Toll Myd88 Tube Pelle DD DD DD DD DD DD DIF Kinase Cactus ? β-TrCP DIF aPKC P Cactus Ref(2)P Proteasome DIF Nucleus DIF Figure 15.4  An illustration of selected details of intracellular Toll pathway that culminates in synthesis of the antimicrobial peptide Drosomysin. Each of the Toll receptors recruits several additional proteins, includ- ing MYD88, Tube, and Pelle, and an intracellular proteolytic kinase cascade is activated. At least one of the kinases in the cascade hydrolyzes the Cactus domain from the Cactus-DIF complex in the cell cytoplasm. Cactus is then further degraded by enzymes, and DIF is translocated to the nucleus where it acts as a tran- scription factor for drosomysin. Expression of Drosomysin results in synthesis of the antimicrobial peptide Drosomysin and its secretion into the circulating hemolymph where it attacks the invading microorganisms. (From Leclerc and Reichhart, 2004. With permission.)

PGRP-LC DD PGRP-LEDD dTak! ? DID ImdDID dFADD DREDD kenny Caspase ? IKK Ird5 P Rel -49 Relish Relish Rel-68 Figure 15.5  The IMD pathway results in synthesis of an antimicrobial peptide, Diptericin, effective against Gram (–) bacteria. Several factors are recruited when the IMD receptor is activated and two intracellular pro- teolytic cascades are initiated. One cascade results in phosphorylation of Relish. The second cascade cleaves Relish into an inhibitory molecule, Relk-49, which remains in the cytoplasm, while Rel-68 is translocated to the nucleus where it serves as a transcription factor for diptericin. The antimicrobial peptide Diptericin is then synthesized and secreted into the hemolymph. (From Leclerc and Reichhart, 2004. With permission.)

Figure 18.13B  B. dorsalis photographed on the Hawaiian rainbow shower tree, a Cassia sp., that releases on its flower methyl eugenol, a pheromone precursor for the Oriental fruit fly male. (Photograph courtesy of Ethel Villalobos and Todd Shelly.)