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Proboscis Extension Reflex
Proboscis Extension Reflex
Proboscis extension reflex (PER) is the extension by an insect with an extendable proboscis (e.g. a bee or fly) of her proboscis
(sticking out of her tongue) as areflex to antennal stimulation. It is evoked when a sugar solution is touched to a bee's antenna.
Contents
Use of PER
How the PER learning paradigm works
PER in honeybees
PER in bumblebees
PER and learning laterality
See also
References
External links
Use of PER
The proboscis extension reflex is part of an insect's feeding behavior. When the antenna is stimulated by sugar water, the proboscis
automatically sticks out to drink.[1] This reflex response can be used to study insect learning and memory in the context of foraging.
The PER paradigm is most commonly used in associative learning experiments in honeybees and bumblebees because it is easy to
use for simple Pavlovian conditioning.[2]
PER in honeybees
The PER paradigm has been successfully used to investigate olfactory learning in honeybees. It was first introduced by Kimihisa
Takeda in 1961. Experiments by Bitterman[2] used first-order classical conditioning to associate an odor with a sugar reward.
Individual bees were placed in a tube with their head sticking out. Then a stream of odorant blown towards the bee's head was
immediately followed by touching the antenna with a sugar droplet. After only three such trials, the odor alone caused the bee to
extend its proboscis approximately 90% of the time. Bees also showed second-order conditioning, learning to associate a second odor
with the original odor.[2] The PER paradigm has also been used in honeybees to study motion learning,[3] thermal learning,[4] color
learning,[5] habituation, and reversal learning.[6]
PER in bumblebees
Although the majority of PER studies are performed on honeybees, there is at least one successful study of using PER on
bumblebees. After they were exposed to a conditioning procedure like that used with honeybees (see above) they gave a conditioned
PER response to odor alone PER response 85% of the time after 10 trials.[7]
See also
Bee learning and communication
Animal behavior
References
1. Braun and Bicker. 1992. Habituation of an Appetitive Reflex in the Honeybee. Journal of Neurophysiology 67: 588-
598.
2. Bitterman et al. 1983. Classical Conditioning of Proboscis Extension in HoneybeesApis
( mellifera). J. Comp. Psych.
97: 107-119.
3. Hori et al. 2007. Associative learning and discrimination of motion cues in the harnessed honeybee,
Apis mellifera L.
J. Comp. Physiol. A 193:825-833.
4. Hammer et al. 2009. Thermal learning in the honeybee,Apis mellifera. J. Experiment. Bio. 212:3928-3934.
5. Jernigan et al. 2014. Color dependent learning in restrained africanized honey bees. J. Exp. Biol. 217:337-343.
6. Komischke et al. 2002. Successive Olfactory Reversal Learning in Honeybees. Learn. Mem. 9:122-129.
7. Riveros and Gronenberg. 2009. Olfactory learning and memory in the bumblebee,
Bombus occidentalis.
Naturwissenschaften 96:851-856.
8. Letzkus et al. 2006. Lateralization of Olfaction in the HoneybeeApis mellifera. Current Biology 16:1471-1476.
External links
Video of PER on a honeybee
Picture diagram of PER on a honeybee
Picture of a honeybee sticking out her proboscis
Article on using PER in detecting illicit drugs and explosives
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