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PHYLUM ARTHROPODA “joint” “footed” animals Reticulitermes Termite Fortress 0.2 mm Coconut Crab hellgrammite Dust Mite Spider Crab holding man Zoologists Are Interested in Embryology WHY? Arthropods are “EUCOELOMATE PROTOSTOMES” Deuterostomes Protostomes e.g. Echinoderms Vertebrates e.g. Molluscs Annelids Arthropods Deuterostomes Protostomes e.g. Echinoderms Vertebrates e.g. Molluscs Annelids Arthropods Arthropods exhibit metamerism 3 3P3 Phyla insects Origin (Chapter 9) “if my theory be true, it is indisputable that before the lowest Silurian stratum … the world swarmed with living creatures.” “Cambrian Explosion” ARTHROPOD SUCCESS …. WHY?? * * * Insects 750,000 named species 30 million total ?? 1 Reason for Success: the Exoskeleton = Cuticle - protective - allows flexibility & mobility - versatile “Quads” Mobility Grasshopper Hind leg “Hamstrings” human elbow joint Joint-Muscle Similarity? Arthropod Cuticle Layered Protein Endocuticle Layered Glycoprotein The “glyco” in glycoprotein: CHITIN Chitin is a polysaccharide similar to cellulose: Light weight Mechanically tough Chemically inert Unmodified Chitin: clear, flexible, resilient Contrasting Cuticles Contrasting Cuticles Contrasting Cuticles Cuticle Modifications for Toughening e.g., Grasshopper thorax Exocuticle becomes “tanned” or sclerotized (increased cross-linking within & between glycoprotein layers) e.g., Crab “shell”, lobster claw Procuticle becomes impregnated with Ca++ salts Cuticle Modifications for Waterproofing Insect Problem: Dessication - small size - air is dessicating - cuticles not water-tight CUTICLE WAX GLAND Solution??? Structure of a WAX O C Fatty Acid O CH2 CH2 CH2 CH2 (CH2)n (CH2)n CH2 CH2 CH3 CH3 Alcohol Pillbugs are land crustaceans!! NO WAX Arthropod cuticles have 1 BIG drawback … They don’t expand to allow growth!!! Vertebrate Bone Growth RADICAL arthropod solution: the MOLT http://www.youtube.com/watch?v=rDK3IT29uoQ http://www.youtube.com/watch?v=2A1i10ZIB-w Ecdysone Secretes Molting Fluid (chitinase, proteases) Molting Fluid attacks Endocuticle Old endocuticle dissolved away Phylum Arthropoda Subphylum Myriapoda “many footed” Class Chilopoda centipedes -2 legs per segment (mostly) -Predators (venom claws) -Fast Subphylum Myriapoda “many footed” Class Diplopoda millipedes -”1000 feet” -Secretive & slow -4 legs per segment (mostly) -herbivorous Subphylum Chelicerata Class Arachnida Order Araneae http://video.nationalgeographic.com/video/kids/animals-pets-kids/bugskids/leafcutter-ant-kids/ Wolf Spider with egg sac Phylum Arthropoda Subphylum Hexapoda Class Entognatha (springtails, snow fleas, etc.) Class Insecta “BUGS” 7,000,000,000 humans on earth … for every person … 200,000,000 insects !! We live in “The Age of Insects” Surprise: not found in world’s largest habitat Breath-taking diversity … Master Niche Fillers Friend and Foe The Honeybee BROWN MARMORATED STINK BUG Halyomorpha halys Basic Body Plan 3 “Tagmata” (body regions) Head Thorax Abdomen Body Variety Body Plan Variety H T A Earwig Cerci Defense Prey Capture Mouthpart Variety (sponges) Sensory Variety Darwin saw usefulness Vision Leg Variety INSECT MOBILITY All insect muscles are striated! Muscles & Endoskeletons (flexion) Antagonistic Pairs (extension) Muscles & Exoskeletons Antagonistic Pairs Q: Which muscle produces the leap? 1st in Flight??? Not Birds … Not Bats Flight has huge implications (foraging, reproduction, migration, escape, etc.) 24 - 26 Orders of insects … 22 have winged forms Wingless orders in this group: silverfish, bristletails, proturans, and springtails 2 Pairs of Wings BEETLES Forewings are protective Hindwings for flight “flies” – Order Diptera 2 Wings Mud Dauber collecting and… transporting mud Spiders packed into a cell larva Mud Dauber Nests Adult Escape Holes Optimal Foraging? Creationist WING Argument: Only fully-formed wings could be beneficial How did Darwin respond?? Weak Flying? Gliding? Thermoregulation? Gas Exchange? Behavioral Signal? Communication Function?? Sustainable Flight Speeds Small grasshoppers Malaria mosquitos Housefly Blowfly Honeybee Large dragonfly Deer bot fly 1.8 km/hr 3.2 6.4 11.0 22.4 30.0 40.0 !! Roughly Bolt’s Top Speed Forward Thrust Downstroke AND Upstroke FLIGHT MUSCLES Direct & Indirect Tergum Indirect Flight Muscle Direct Flight Muscle Sternum Bird-like Bat-like Note locations of fulcrum and muscle insertion Downstroke Both Muscle Groups are Indirect Fulcrum … yes Muscle insertion … no http://www.youtube.com/watch?v=iUNRkvwIUQ0 Slow wing beats … slow neuromuscular activity EXAMPLE: butterflies @ 4 beats/sec Synchronous Neural Stimulation 1 action potential 1 set of muscle cell contractions NMJ 1 wing beat Extreme wing speed … Can nervous systems keep up?? Asynchronous Stimulation e.g., some midges exceed 1000 beats/sec midge 1000 beats/sec = 1 beat per ms !! This should be IMPOSSIBLE … why? Neuron refractory periods: 5 – 10 ms In other words, muscles are contracting faster than nervous systems operate !! MECHANISM When the upstroke muscles contract, they stretch the downstroke muscles. Downstroke muscles respond by contracting which stretches the upstroke muscles. This antagonism may continue several cycles before a neural input is needed to keep it going. In extreme cases, 40 wingbeats have been recorded for each neural input. FLIGHT TEMPERATURES Thoracic temperature of a flying sphinx moth (Manduca sexta) mammals Flying and running are energetically demanding … super lungs and heart?? NO (!) (at first look) Insect Respiration Spiracle Body wall branching resembles that inside mammalian lungs Flying insects have large aerobic requirements while having a crude OPEN circulatory system. PARADOX? How do insects “get away with” this system? Danger of Dessication High SA:VOL exacerbated by … flying ?? high metabolic rate? producing urine? H2O Spiracles are Valved Water Loss In Urine Hemolymph (blood) in the Hemocoel (body cavity) [NO CAPILLARIES] Marcello Malpighi (1628-1694) NITROGENOUS Waste Products of Animals Ammonia Urea Uric Acid Cost of Synthesis none low high Toxicity high moderate low Solubility high high Diffusion Coefficient high intermediate low 1 2 4 N per osmotic particle very low Active in re-claiming water Hyperosmotic Urine (U/P> 1) Significance?? What animals can produce it?? - Mammals - Birds Potent Kidneys - Arthropods (Malpighian Tubules) Animal urine can be dilute … or concentrated relative to body fluids U/P osmotic pressure of urine osmotic pressure of plasma INSECT DEVELOPMENT Holometabolous Development (88% of all insects) Maggot Grubs Radical reconstruction HEMIMETABOLOUS DEVELOPMENT A gradual metamorphosis e.g., grasshoppers, cicadas, preying mantis, mayflies, dragonflies Class Insecta Order Coleoptera Lepidoptera Hymenoptera Know examples Diptera from each Order Hemiptera Homoptera Orthoptera Friday’s Lab Optimal Foraging by Butterflies Natural selection pushes species TO BE ADAPTED to their environment. Useful traits often appear optimized. Why not maximized?? Swimming in cold water Blubber: What is optimal for a seal? optimal Food to stomach? Air to lungs? Blood to brain? Wouldn’t an even longer neck be better?? Trade-offs? How long is long enough? OPTIMALITY MODEL Benefit Amount of Cost or Benefit MAX Opt “Extent” of Phenotype Cost “proboscis” Flow = ΔPπr4 8lη P = pressure inside proboscis r = proboscis radius l = proboscis length η = fluid viscosity Lab Variable Amount/mL Amount/mL ? Costs? Concentration SUGAR CONCENTRATIONS (%): 8.75, 17.5, 35, 50, & 80 (η)