5. Circadian Clock and Sleep
Role of circadian clock in bees
Anticipate day–night fluctuations
Time visits to flowers
Time compensation when
referring to the sun in dance
Regulation of sleep
Molecular characterization suggests the clock of the
bee is more similar to mammals
Rothschild and Bloch (2008)5
6. “‘Hard-eyed’ creatures and insects manifestly assume the
posture of sleep; but the sleep of all such creatures is
of brief duration, so that often it might well baffle one’s
observation to decide whether they sleep or not.”
Do bees sleep?
6
(Aristotle, On Sleep and Sleeplessness 350 BC)
7. Yes they do! It seems…….
Honey bees are the first invertebrates for which a
sleep-like state was described (Kaiser, 1983)
Characteristic posture
• Flex legs
• Bringing its head to the floor
• Antennae stop moving
• Hold each others legs as they sleep
Reduced muscle tonus
Elevated response threshold (Piéron, 1913)
Sleep rebound after sleep deprivation (Sauer, 2004)7
8. Sleep stages in bees
IA-Immobile-active state FS- First sleep stage
SS-Second sleep stage TS-Third sleep stage
Rothschild and Bloch (2008)8
9. Sleep in Task- Based Castes (Age Polyethism)
Klein et al., (2008) 9
10. Bees’ Sleep Vs Mammalian Sleep
Characteristic posture
Reduced muscle tonus
Elevated response
threshold
Sleep rebound after sleep
deprivation
Persistence throughout life
Bees’ sleep+ SWS
and REM sleep
MammalianBee
10
11. Sleep Function
Energy conservation
Restoration at the cellular and network levels
Maintenance of synaptic homeostasis
Memory consolidation
Rothschild and Bloch(2008)
Research confirms, there is strong link between
learning, memory and sleep (Rasch and Born, 2013)
11
14. Well documented for humans (Born et al., 2006)
Mammals (Siegel, 2005; Capellini et al., 2009)
Birds (Jackson et al., 2008)
Sleep in Memory Processing
14
17. Navigation Memory
• RFID was used to record flights and behavior
• RFID bees behaved normally
• Bees released to unknown site
• Then those bees reached hive are sleep deprived
• And released from same site
17
19. Sleep and Navigation Memory
Bees transferred to an unknown release site
Sleep deprived: ~50% reached hive
Sleep group: 1st Release 58%, 2nd Release 83%
Menzel et al., (2012)
19
20. Sleep Duration and Navigation Task
Bees slept longer during the night following forced
navigation tasks
Not due to exhaustion
Distance didn’t affect sleep time during night
Menzel et al., (2012)
20
22. Classical Conditioning (CC) / Pavlov’s CC
Over the period of time
US CS
Extinction Learning (EL)
Continuous CS without US
CC
Unconditioned Stimulus (US)
Conditioned Stimulus (CS)
Proboscis Extension
Response (PER)
No PER
Concepts of Learning
sucrose
sucrose
22
23. Classical Conditioning (CC) / Pavlov’s CC
Over the period of time
US CS
Extinction Learning (EL)
Continuous CS without US
CC
Unconditioned Stimulus (US)
Conditioned Stimulus (CS) No PER
Concepts of Learning
sucrose
sucrose
23
Proboscis Extension
Response (PER)
24. Sleep deprivation affects extinction but
not acquisition memory in honeybees
• Sleep deprivation had no effect on retention scores
after odor acquisition
• Whereas, retention for extinction learning was
significantly reduced
• Hence, consolidation of extinction memory was
affected by sleep deprivation but not acquisition
memory
Hussaini et al., (2009)
24
32. Sleep deprivation impairs precision of
waggle dance
Slept bees (n = 7)
SD of dance angles (13.78 ± 0.91)
Sleep deprived bees (n = 6)
SD of dance angles (16.49 ± 0.98)
One angle per bee per day
Greater the SD of dance angles; lesser the directional precision
Klein et al., (2010)
32
34. Electrophysiology of Bees’ Sleep
• Neuronal activity recorded in brain regions including
the mushroom bodies
• Nocturnal resting periods - the overall spike activity
was markedly increased
• Periodical variations of spike rate were present when
no visible antennal movements occurred
• Rhythmic brain activity in sleeping bees
Schuppe, (2015 )
34
35. Conclusions
Sleep has an absolute role in memory processing
Honey bees an attractive model to study sleep,
memory and learning in a natural context
Small brain (with about 1 million neurons)
Smart learners
Learning is reinforced during sleep – humans?
BEES’ MAGIC WELL IS NOWHERE NEAR EMPTY
35
36. References
Beyaert, L., Greggers, U., Menzel, R., 2012. Honeybees consolidate
navigation memory during sleep. J. Exp. Biol. 215, 3981–3988.
Eban-Rothschild, A.D., Bloch, G., 2008. Differences in the sleep
architecture of foragerand young honeybees (Apis mellifera). J.
Exp. Biol. 211, 2408–2416.
Hussaini, S.A., Bogusch, L., Landgraf, T., Menzel, R., 2009. Sleep
deprivation affects extinction but not acquisition memory in
honeybees. Learn. Mem. 16, 698–705.
Menzel, R., 2012. The honeybee as a model for understanding the basis of
cognition. Nat. Rev. Neurosci. 13, 758–768.
Klein BA, Olzsowy KM, Klein A, Saunders KM, Seeley TD (2008) Caste-
dependent sleep of worker honey bees. J Exp Biol. 211:3028– 3040.
Eban-Rothschild, A. D., Bloch, G. (2012) Circadian Rhythms and Sleep in
Honey Bees, chapter 1.3., p. 31-46 in “Honeybee Neurobiology
and Behavior: A Tribute to Randolf Menzel”.
Kaiser, W. 1988 Busy bees need rest, too-behavioural and electromyographical
sleep signs in honeybees, J. Comp. Physiol. 163: 565–584. 36
Editor's Notes
Elevated response threshold-higher intensity stimulus is needed to produce a response
[IA] – an awake bee stays in the same place, the thorax, abdomen, and head are clearly raised above the substrate. [FS] – the abdomen and thorax are clearly raised above the substrate, and the antennae are extended at an angle of 90–180°, between the pedicle and the scape. [SS] – body is typically more adjacent to the substrate, and the antennae are extended at an angle of ~90° between the pedicle and the scape. [TS] – the muscle tonus is reduced, and the body is adjacent to the substrate. The angle between the pedicle and scape < 90°, with the antennae tips typically touching the substrate.
3 days as a cell cleaner, 4–12 of as a ‘nurse bee’, 13–20 as a ‘food storer’
(I) Encoding - up-take of the information to be stored into a neural representation. (II) Consolidation - stabilization of the memory that follows encoding and enables the retention of a memory over time. In the absence of such consolidation the information would be rapidly forgotten. Forgetting can result from a decay of the memory trace or from retroactive interference as the encoding of new information leads to an overwriting of the information encoded before. (III) Retrieval of the stored information refers to the reactivation of a stored memory in the context of more or less goal-directed behavior.
IMM-intermediate and medial mesopallium, unknown locus termed S
The bee in A was trained to a feeder close to the hive (distance 10m), The two bees in B and C were trained to a feeder 200m east of the hive. The
small triangles indicate colored tents serving as radartransparent landmarks