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TODAY: Structure & Function Part I • Muscles & Movement • Blood & Circulation • Tracheal System & Gas Exchange Muscles & Movement IMPORTANT CONCEPTS: • All striated (no smooth) • Major Types: 1) Synchronous 2) Asynchronous • Motion often driven by both muscles and cuticular flexure + energy storage Diagrammatic structure of a striated muscle fibril, the basic muscle unit. Z X-sec Z A muscle unit comprised of several fibers each made up of many fibrils from Chapman Typical ennervation of insect muscle; slow & fast axons in parallel (vs. graded response of vertebrates). from Chapman Muscle attachments to exoskeleton. from Gullen & Cranston the tentorium an internal framework for muscle attachment in some insectsfrom Snodgrass 1935 Muscles of honey bee abdomen. from Snodgrass Caterpillar body wall musculature; functions: undulatory movement & hydrostatic skeleton. Caterpillar gut musculature. Types of Movement Larvae Sinuous motion, lateral muscular waves, some primitive fly larvae e.g midges Undulatory movement, anterior + posterior waves, typical of moth & butterfly caterpillars Whip-like, posterior + anterior waves, used with turgor muscles some caterpillars such as inch worms Adults Walking, leg strokes Jumping, aided by cuticular flexion Swimming, aided by hairs, special appendages Flying, aided by cuticular flexion at wing base & whole thorax Typical “tripod gate” of an insect, maximum center-ofgravity stability with simplest mechanics and control. Action: Thoracic muscles pull on leg bases; fine control by extension & flexion of internal leg muscles Rhythm: slight offset between legs: 1-2-3 & 1-2-3 … it’s a waltz! Jumping The main power Resilin source comes from the release of energy in the cuticle, which has been “cocked” by the muscles. Super-flexible resilin allows extreme bending of the joint. distortion, cuticular energy storage Visible “chevrons” = muscle attachments to cuticle. Swimming Often assisted by paddlelike appendages &/or hairs that fold backward on protraction, reducing drag. In aquatic beetles, the different syncopation of swimming legs is characteristic of some families. Predaceous diving beetle swimming adult (top), walking larva (bottom). Thoracic musculature of honey bee. from Snodgrass 1935 Indirect flight muscle action within thorax Circulatory System Main Points: • Blood = “haemolymph” • Generally not pressurized • Does not distribute oxygen • Heart (“aorta”) is dorsal Generalized insect circulatory system. (Gullen & Cranston, 2000, Fig. 3.9) Haemolymph, insect blood Body composition Larvae 20-40% Adults <20% Constituents H2O (~90%) Plasma amino acids organic acids phosphates sugars, trehalose (energy rich disaccharide characteristic of insect blood) Haemocytes, diverse cells with numerous functions Haemolymph Functions • Chemical exchange (e.g. ion exchange in excretory system) • Nutrient distribution • Waste removal • Hormone transport • Pressure changes support: hydrostatic skeleton molting turgor ventilation • Thermoregulation (heat distribution, protection against freezing) • H2O reserve • Defense wound healing toxins haemocytic action Haemocytes Cell Type Major Function(s) Location Plasmatocytes, Granulocytes, Prohaemocytes Defense (e.g. phagothroughout hoemocoel cytosis, encapsulation, coagulation), storage & distribution of nutrients Cystocytes coagulation throughout hoemocoel Nephrocyes haemolymph filtering, metabolize wastes for excretion localized: near dorsal vessel Oenocytes lipid synthsis (haemoglobin synthesis, rare) localized: fat body, epidermis Origin: embryonic mesoderm, singular generation (no bloodmaking organs in adult insects) Defense Functions of Haemolymph Coagulation Phagocytosis Antibacterial protein reactions Immune response signaling Noxious/toxic compound reservoir & delivery: Encapsulation: from Chapman ca. 1970 from Gullen & Cranston 2000 from Gullen & Cranston 2000 Tracheal System Main Points: • Oxygenation of tissues is accomplished mostly by passive diffusion • Double diffusion gradients: O2 (in) & CO2 (out) • Basic structure: spiracles =>tracheal system=> ending in tracheoles • Insect size partially determined by physical limits to diffusion & tracheal system Tracheal System tracheole-tissue interface The end terminals of the tracheal system. Spiracles taenidia • Interface with environment • Beginning of O2 diffusion gradient • Generally one pair per segment (up to 10 seg.) but varies between species; position, shape, number may be characteristic of taxon Tracheoles • Interface with O -demanding tissue 2 • Beginning of CO2 diffusion gradient • Microscopic blind-end • Liquid-filled • May penetrate tissue • Most numerous at highly active tissue • Can proliferate in response to long-term O2 deprivation from Gullen & Cranston 2000 Physical Basis of Tracheal system: Diffusion Amongst tropical slow-moving • Limits: diffusion only works over thin layers of tissue; rhinocerous beetles are the increased requirement formost tracheation with increase massive modern insect in size. species. Surprizingly, mostMost can “large” • Implication: Insect size limited by air supply. fly. or Internally they metabolism, are filled insects are long and slender, have low or with a dense mass of tracheae. display short durations of activity&/or are highly tracheated. Tinyknown insects have insect reduced tracheae because they Longest modern (body): Megaphasma denticris (PHASMATODEA) can breathe through their outer cuticle. ~ 30 cm long, native of SE Asian tropics. Long, slender body => shorter diffusion distance Modification & Control of the Tracheal System Basic Division into sections: spiracle => (trunk) => trachea => tracheole Subdivision & specialization trunks & air sacs tracheole proliferation gills aeriferous tracheae Control of flow spiracular valves water-conserving matrices, filters atrial chambers Movement-assisted air flow (“breathing”) Thoracic/Abdominal pumping (trunks as “bellows”) Tracheal contracting Air Sacs • Adaptations for more effective air supply during flight, i.e. high oxygen expendature. • Expansion of lateral tracheal trunks. • Present in many flying insects. • May take up large proportion of body cavity. • “Bellows” or quasi-lung function. • Depends on adaptation of abdomen for “pumping” action. from Snodgrass ca. 1935 Air sacs in the honey bee. Open vs. Closed Tracheal Systems a) cockroach, lateral trunks b) honey bee, air sacs c) mosquito larva, siphon d) small fly larva, cutaneous gas exchange e) mayfly nymph, external gills f) dragon fly nymph, “internal” gills Gullen & Cranston, 2000, Fig. 3.11 Gills Closed system Thin membrane allowing diffusion of oxygen CADDISFLY abdominal (TRICHOPTERA) Configuration, extent, location may be diagnostic of taxon Some types: leaf-like abdominal lobes (ODONATA: ZYGOPTERA) from Gullen & Cranston 2000 internal gills internal chamber (ODONATA: ANISOPTERA) c.f. siphon (open, NOT gills) from Borror & White (HEMIPERA) GIANT WATER BUG The Physical or “Gas Bubble” Gill • Underwater respiration with an open tracheal system • A bubble of atmosphere is captured… • …and serves as a gas-transfer chamber and water-stopper Modifications of the Cuticle for Aquatic Respiration from Gullen & Cranston 2000 Subelytral space in a predaceous diving beetle; it physically traps a temporary sir supply. from Chapman ca. 1970 The “plastron”, physical gill integrated into the integument; channelized cuticle with hydrophobic hairs; it holds a bubble by physical entrapment and surface tension. ~ end ~