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11/12/2015 ocelli frontal nerve (to cibarium) compound eye PROTOCEREBRUM red & black = CNS blue = VNS lateral connectives neurosecretory cells DEUTOCEREBRUM FRONTAL GANGLION to antenna BRAIN TRITOCEREBRUM frontal connective with tritocerebrum MOUTHPARTS: to labrum SUBESOPHAGEAL GANGLION recurrent nerve postesophageal (paraesophageal) connectives 1 2 3 4 2 3 4 5 6 CORPUS CARDIACUM circumesophageal connective to mandible to maxilla to labium & hypopharynx HYPOCEREBRAL GANGLION 1 CORPUS ALLATUM SUBESOPHAGEAL GANGLION VENTRICULAR (INGLUVIAL) GANGLION ventral nerve cord TRITOCEREBRUM DEUTOCEREBRUM BRAIN: PROTHORACIC GANGLION PROTOCEREBRUM to anterior body wall to leg to posterior body wall to spiracle & alary muscles to anterior body wall to wing to leg to posterior body wall MESOTHORACIC GANGLION to spiracle & alary muscles CAUDAL GANGLION (METATHORACIC & ABDOMINAL ganglia not shown) caudal nerve Centralization of CNS (1) (2) (0) Apterygota (unspecialized) e.g. 8 abdominal ganglia (7 in Odonata) (5/6 in Orthoptera) 1 11/12/2015 Types of neurons dendrites UNIPOLAR (motor & associative) axon cell body BIPOLAR (sensory) axon dendrites MULTIPOLAR (sensory) neurofibrillae axon dendrites Nerve connections axons of unipolar motor neurons muscle GANGLION multipolar stretch receptor (sensory) associative neuron (unipolar) seta epidermis cuticle axons of bipolar sensory neurons INTEGUMENT 2 11/12/2015 Brain or ganglion, cross-section neural lamella Typical nerve perineurium (secretes the neural lamella) giant fibers nerve cell bodies fiber tracts Glial cells: 1. Protection (cushioning) 2. Nutrition (passing nutrients & salts to neurons) 3. Food storage (glycogen) 4. Insulation (for delicate, leaky neurons/axons) neuropile (principal site of synapses) The insect brain ocellar nerve optic lobe antennal nerve (frontal ganglion) Protocerebrum Deutocerebrum Tritocerebrum (2nd ant.) circumesophageal connective postesophageal connective(s) Subesophageal ganglion Grasshopper brain (Orthoptera) Same as 520my-old 3-part brain of earliest arthropods, e.g. Fuxianhuia. 3 11/12/2015 The CNS: anterior components corpus cardiacum ocellar nerve PROTOCEREBRUM hypocerebral ganglion antennal nerve AORTA ocellar nerve corpus allatum DEUTOCEREBRUM inner esophageal nerve position of molt gland TRITOCEREBRUM frontal ganglion lateral connective ventricular (ingluvial) ganglion` recurrent nerve circumesophageal connective labral nerve mandibular nerve SUBESOPHAGEAL GANGLION PROTHORACIC GANGLION hypopharyngeal nerve nerve to neck maxillary nerve labial nerve ocelli frontal nerve (to cibarium) compound eye PROTOCEREBRUM Ganglia posterior to the brain lateral connectives neurosecretory cells DEUTOCEREBRUM FRONTAL GANGLION TRITOCEREBRUM frontal connective with tritocerebrum labral nerve recurrent nerve postesophageal (paraesophageal) connectives CORPUS CARDIACUM circumesophageal connective to mandible (to hypopharynx) to maxilla to labium (& hypopharynx?) HYPOCEREBRAL GANGLION CORPUS ALLATUM SUBESOPHAGEAL GANGLION VENTRICULAR (INGLUVIAL) GANGLION ventral nerve cord PROTHORACIC GANGLION to anterior body wall to leg to posterior body wall to spiracle & alary muscles MESOTHORACIC GANGLION to anterior body wall to wing to leg to posterior body wall to spiracle & alary muscles CAUDAL GANGLION (METATHORACIC & ABDOMINAL ganglia not shown) caudal nerve 4 11/12/2015 Brain sizes (as fraction of body weight) Dytiscidae (Coleoptera): 1/4000 Termitinae (Isoptera): 1/280 Pompilidae (Hymenoptera): 1/400 Apidae (Hymenoptera): 1/75 Mushroom bodies or corpora pedunculata (Ex. = worker ant brain) …and in Drosophila yellow = visual data red = olfactory data blue = higher associative centers 5 11/12/2015 Giant fibers (axons): No synapses, large diameter ventral nerve cord of Syrphidae (Diptera) 20-60 µ ? Deuterostomia Lophotrochozoa <5µ Insecta Nematodes Ecdysozoa Myelinated (red) (100x faster) Non-myelinated (blue) Evolution of myelinated axons (red) The VNS: Stomatogastric components corpus cardiacum ocellar nerve protocerebrum hypocerebral ganglion antennal nerve AORTA ocellar nerve corpus allatum deutocerebrum inner esophageal nerve position of molt gland tritocerebrum frontal ganglion lateral connective ventricular (ingluvial) ganglion recurrent nerve circumesophageal connective labral nerve subesophageal ganglion prothoracic ganglion mandibular nerve hypopharyngeal nerve nerve to neck maxillary nerve labial nerve 6 11/12/2015 ocelli frontal nerve (to cibarium) compound eye PROTOCEREBRUM Visceral Nervous System (in blue) lateral connectives neurosecretory cells DEUTOCEREBRUM FRONTAL GANGLION TRITOCEREBRUM frontal connective with tritocerebrum labral nerve recurrent nerve postesophageal (paraesophageal) connectives CORPUS CARDIACUM circumesophageal connective to mandible to maxilla HYPOCEREBRAL GANGLION CORPUS ALLATUM to labium & hypopharynx SUBESOPHAGEAL GANGLION VENTRICULAR (INGLUVIAL) GANGLION ventral nerve cord PROTHORACIC GANGLION to anterior body wall to leg to posterior body wall to spiracle & alary muscles MESOTHORACIC GANGLION to anterior body wall to wing to leg to posterior body wall to spiracle & alary muscles CAUDAL GANGLION caudal nerve Stomatogastric VNS Function 7 11/12/2015 CNS function: Reflex behavior 1. Reflex excitability – e.g. light, through dorsal ocelli 2. Reflex immobilization – total inhibition of CNS • • • Thanatosis Warning attitudes (e.g. caterpillars) Camouflage (e.g. stick insects) 3. General immobilization (insect “sleeps”) • • • Remove antennae Remove halteres in Diptera Block off all ocelli 4. Note: Brain is the inhibitory center • Example: mantis male with head eaten Nymphalis antiopa – the mourning cloak butterfly (experiments on neural integration) (also: brain ruling ganglia) 8 11/12/2015 Reflex immobilization (Thanatosis) Thanatosis in a gryllid “Death-feigning beetle,” Crytoglossa variolosa (Tenebrionidae) Learning 1. Habituation (learning not to respond) • • cedarwood oil or some other challenge constant prodding of a cockroach 2. Conditioning (responding to something previously ineffectual) • floral odors; stimulus assoc. w/ feeding 3. Imprinting (special type of conditioning) • • critical developmental stage (plasticity) host shifts, host fidelity, and speciation 4. Trial and Error • beetle courtship – “long axis” 5. Latent Learning (recognition of landmarks) • antlion Note: Percent of brain made up of corpora pedunculata: workers = 13.5% queen = 9.2% drones = 5.6% Pompilidae (Hymenoptera) solitary wasps, bee foraging, ant mazelearning 6. Time sense (cuing to 24 hour cycle) 9 11/12/2015 “Sophisticated” Learning Latent learning (landmark orientation) (Digger wasp – Hymenoptera: Sphecidae) (not immediately expressed as an overt response) Conditioning [to nectar sources] (Honey bee – Hymenoptera: Apidae) Learning through Imprinting (host fidelity and speciation – Diptera: Tephritidae: Rhagoletis pomonella) hawthorns apples overlap 10