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Insect Behavior III. Control of Behavior A.
General for any behavior, animal must --
recognize relevant stimuli (stimulus
filtering) -- decide to respond --
execute appropriate motor response
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Insect Behavior III. Control of Behavior A.
General for behavior to be adaptive --
response to stimulus and decision making must be
synchronized with environment
ex apple maggot
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• Insect Behavior
• III. Control of Behavior
• A. General
• -- concept of response threshold level of
  stimulation
• necessary to trigger a response
• If threshold low heightened sensitivity
• If threshold high decreased sensitivity

-- control of behavior therefore involves
mechanisms that influence response
thresholds (ex circadian rhythms)
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Insect Behavior III. Control of Behavior B.
Nervous control 1. stimulus filtering
-- sensory tuning sign stimuli
ex bombykol and male silk moth
tarsal chemoreceptors in flies
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Insect Behavior III. Control of Behavior B.
Nervous control 2. generating response
-- CPGs (neural mechanism for FAP)
-- modified by feedback -- regulated by
supra- and subesophagael ganglia -- ex
praying mantis
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Caudal ganglion
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Insect Behavior III. Control of Behavior B.
Nervous control 2. generating response
-- CPGs (neural mechanism for FAP)
-- modified by feedback -- regulated by
supra- and subesophagael ganglia -- ex
honey bee sting
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Insect Behavior III. Control of Behavior B.
Nervous control 3. decision making
a. Feeding in blowfly
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hypocerebral ganglion
tritocerebrum
Recurrent nerve
frontal ganglion
circumesophagael connective
subesophagael ganglion
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Feeding in Blowfly
-- steps in food tarsal chemreceptors fire --
probosis extension reflex -- sensilla on
proboscis fire CPG for pharyngeal
pump crop fills inhibitory
messages stretch receptors fire via recurrent
nerve
X
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Insect Behavior III. Control of Behavior B.
Nervous control 3. decision making
b. Evasion of bats by noctuid moths
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-- hunt by echolocation -- emit pulsed sounds,
20 80 KHz
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Moth shows two responses 1. turns and flies in
opposite direction -- approaching or
receding? -- left or right? -- above or
below? 2. performs erratic, loops and dives and
eventually drops to ground
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Insect Behavior III. Control of Behavior B.
Nervous control 3. decision making
b. Evasion of bats by noctuid moths A1
-- fires only to pulsed sounds
20-80KHz -- firing rate fluctuates with
sound intensity -- sensitive to soft
sounds
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Insect Behavior III. Control of Behavior B.
Nervous control 3. decision making
b. Evasion of bats by noctuid moths A2
-- fires only to pulsed sounds
20-80KHz -- constant firing rate
-- responds to loud sounds
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Bat 100 ft away from moth -- moth can detect
bat bat cannot detect moth -- A1 fires moth
turns and fly away
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Approaching or receding? -- if approaching, rate
of A1 firing increases -- if receding, rate of AI
firing decreases
approach
recede
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Left or right?
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Above or below? -- if above, firing of A1
fluctuates with wing beat -- if below, firing of
A1 constant and not synchronized with wing
beat
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Bat 10 ft away from moth -- moth can detect bat
bat can detect moth -- A2 fires looping, erratic
flight
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation -- influences behavior on
long-term time scale -- alters response
thresholds -- helps to coordinate behavior
with long-term events such as age
maturation, season, etc.
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) -- do not release
specific behaviors -- alter response
thresholds for many stimuli -- primes
the animal to respond
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) a. biting in female
mosquito -- female takes blood meal --
stimulate ovarian development (via EDNH) --
ovaries release ecdyson -- ecdyson raises
response thresholds inhibits further
biting
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) b. division of labor
in honey bees -- exhibit polyethism division
of labor based on age
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0-2 days of age cell cleaners
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2-11 days of age brood care (nurse bees) --
hypopharyngeal and mandibular glands developed --
attend queen
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11-20 days of age middle aged bees -- food
storage and processing -- wax production and comb
building -- wax glands active -- hypopharengeal
and mandibular glands regress -- guarding alarm
pheromone glands active
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20 days of age and older foragers -- active
biological clock -- respond to stimuli outside
hive -- perform and decode waggle dances
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) b. division of labor
in honey bees -- regulated in part by JH --
as worker ages, JH increases -- changes
response thresholds
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JH titer
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2
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Age (in days)
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) b. division of labor
in honey bees 1) experimental treatments with
JH -- precocious foragers (behavioral
acceleration)
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Precocious foragers
JH treated
JH titer
untreated
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2
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Age (in days)
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) b. division of labor
in honey bees 1) experimental treatments with
JH -- precocious foragers (behavioral
acceleration) -- overaged nurses
(behavioral reversion)
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Over-aged nurse
JH titer
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2
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Age (in days)
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) b. division of labor
in honey bees 2) social regulation of JH
titers -- contact with older bees
retards rate of JH increase ( normal
development) -- ethyl oleate
forager
young bee
Workers engaged in trophallaxis
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Precocious foragers
w/o contact with older bees (accelerated increase)
JH titer
w/ contact with older bees (normal increase)
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2
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Age (in days)
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) b. division of labor
in honey bees 3) adaptive value of plasticity
of division of labor -- loss of
foragers -- JH levels of some (but not
all) younger workers increase
-- precocious foragers
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 1. Modifier effects
(primer effects) b. division of labor
in honey bees 3) adaptive value of plasticity
of division of labor -- swarming
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 2. Releaser effects
-- eclosion hormone
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Insect Behavior III. Control of Behavior B.
Hormonal Regulation 2. Releaser effects
-- Queen Retinue Pheromone (QRP)
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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene
effects sequenced genomes for
Drosophila Apis
mellifera ortholog genes in different
species that evolved from a common
ancestral gene by speciation usually
retain the same function in all species.

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Brain gene expression in Apis mellifera
Microarray analyses
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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene effects
a. per gene in honey bees young bees
arrhythmic foragers rhythmic foragers
have 3X as much PER as nurse bees
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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene effects
b. for locus -- codes for cGMP-dependent
protein kinase (PKG) -- influences foraging
activity
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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene effects
b. for locus Drosophila
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Drosophila two alleles for S sitter
lower PKG levels less active for R
rover higher PKG levels more active
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Drosophila two alleles for S favored in
habitats with food, competition for R
favored in habitats with food,
competition allelic variation affects
behavior operates over an evolutionary time
scale
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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene effects
b. for locus Apis mellifera
Amfor (for orthologue)
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Amfor mRNA
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• Honey Bees
• Amfor affects behavior, not by allelic variation,
  but by altered expression of same alleles during
  development
• Regulates PKG-induced in phototaxis move to
  hive entrance where other stimuli trigger
  foraging

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• Amfor expression influenced by social environment
• -- behavioral acceleration ? precocious
  foragers
• have increased PKG
• -- treatment with PKG ? precocious foraging
• -- behavioral reversion ? overaged nurses
• PKG levels decrease

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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene effects
c. hygienic behavior in honey bees --
uncapping and removal of diseased larvae
American foulbrood (Bacillus larvae)
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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene effects
c. hygienic behavior in honey bees 2
loci -- uncapping U (do not uncap) µ
(uncap) -- removal R (do not remove) r
(remove)
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• Insect Behavior
• III. Control of Behavior
• C. Genetic control
• 1. single gene effects
• c. hygienic behavior in honey bees
• used controlled breeding to generate
  genotypes
• U_ R_ non-hygienic
• µµ R_ would uncap but not remove
• U_ r r would not uncap, but would remove
  disease
• larvae if uncapped
• µµ r r hygienic

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Insect Behavior III. Control of Behavior C.
Genetic control 1. single gene effects
d. Gp9 locus of fire ants, Solenopsis invicta
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Solenopsis invicta
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• Two colony types
• Monogyne colonies
• -- single queen mates, founds new nest
  individually
• -- colony has only one queen
• -- workers execute all foreign queens

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• Two colony types
• Polygyne colonies
• -- newly mated queens join existing nests
• -- colony has multiple queens form huge
  extended colonies
• -- workers tolerant of foreign queens

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• Regulated by 2 alleles at Gp9 locus
• -- B always associated with monogyny
• -- b always associated with polygyny
• BB workers will accept only BB queen and only
  one of them
• kill all other queens
• Bb workers will accept multiple queen, but only
  if Bb
• bb queens lethal recessive
• BB queens are killed by workers

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• Gp9 codes for odorant binding protein
• -- involved in pheromone production and
  recognition
• -- single gene with two alleles controls
  worker response
• thresholds to queen pheromone and ultimately
  
• colony social structure

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• Insect Behavior
• III. Control of Behavior
• C. Genetic control
• 2. polygenic effects most behavior
  controlled by
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  multiple genetic loci
• a. Selection experiments and gene
  mapping
• ex pollen collection in bees
• High and low pollen hoarding lines
• Multiple loci involved that influence tendency to
  collect pollen, responsiveness to sucrose,
  learning ability, behavioral maturation, hormone
  levels, ovarian development, etc.
• But, about 50 of variation in pollen hoarding
  behavior attributed to only two loci.

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Insect Behavior III. Control of Behavior C.
Genetic control 2. polygenic effects most
behavior controlled by
multiple genetic loci
b. Hybridization experiments ex cricket
song
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Gryllus
Teleogryllus