Synchronisations of neural oscillations

"Synchronization of rhythmic activities is likely to underlie functional

interactions between neurons within a defined brain structure, or between
disparate populations in distinct structures. Equally, abnormal synchronization
may impair functional interactions and contribute to complex cognitive
disorders such as schizophrenia and attention deficit/hyperactivity disorder."
Jones & Wilson 2005

Targetted recordings of genetically labelled neurons allow greater resolution
in understanding the cellular mechanisms behind synchrony.
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Membrane Potential Dynamics and Synchrony
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Temporal Binding Hypothesis
This hypothesis was proposed in 1981 by Von der Malsburg and refined
further by Eckhorn et al. in 1988 (Singer 2007). The hypothesis was an
answer to the concern that the number of higher cardinal cells, cells
representing combinations of lower-order neurons, would not be sufficient to
account for the permutations of possible representations. Synchronization is
believed to allow one cell to belong to multiple assemblies. (Shadlen &
Movshon, 1999). One problem to be addressed was how to discriminate
'visually elicited synchrony' from synchrony due to binding.

Feature Binding
Herculano-Houzel, et al. (1999) stimulated mesencephalic reticular formation
to evaluate spontaneous state chances and transitions. Oscillatory patterning
in the 70-105 HZ range is attributable to sensory input requiring a minimal
level of activation. Contrasting to this, synchronization at the 20-65 Hz range
occurs from intra-cortical synchronizing mechanisms. They found binding
occurred more as central activation increased. From this they concluded that
feature binding depends on central states.

Acquisition and recall of memory can engage multiple brain regions
The Medial Temporal Lobe has been well established in the encoding of
memories.
The Hippocampus has projections into multiple regions within the MTL. It is
generally attributed to spatial memory tasks and behavioural learning.
The Amygdala can play a critical role in heightened emotional memory
encoding.

How multiple brain regions contribute to a common behavioural
outcome

Emotional memory formation
During the waking state, theta is observed in the amygdala during periods of
heightened arousal and this seemed to be associated with the coherence to
the frontal cortex (Par & Gaudreau, 1996).

Recall
Whilst it had already been shown in marsupials, Anderson et al (2009)
recorded cortical electrical activity from the medial temporal lobe, the
prefrontal cortex and the lateral temporal lobe in human epilepsy patients.
They found significantly higher MTL-PFC pairing during free recall and further
analysis revealed this was resulting from a higher bi-directional information
flow between the two areas.

Sleep
Pare and Gaudreau (1996) performed intra cellular recordings in the slow-
wave, sleeplike rhythms of ketamine-xylazine anesthetised cats. During sleep,
the Hippocampus and entorhinal cortex display heightened theta oscillations.
The perirhinal and BL amygdala also display these oscillations however this
seems to be phase locked to the hippocampal theta oscillations (Par &
Gaudreau, 1996). This supports the idea that sleep rhythm is shaped by
corticothalamic feedback, rather than exclusively thalamic signalling.

Conditioned freezing in rats is a response elicited by rats particularly when
trapped and subjected to an aversive unconditioned stimulus such as a foot
shock. (Fanselow, 1984)

The Amydala
Early electro-convulsive therapy showed that the Amygdala was responsible
for positively and negatively motivated tasks: socially motivated tasks,
uncontrolled fear responses and emotional content (Pare, Collins, Pelletier,
2002).
Paz et al (2006) showed that the Basolateral Amygdala increased
transmission from perirhinal to entorhinal neurons after unexpected rewards.
This seemed to enhance emotional memory formation by increasing
sensitivity to emotional cues.
The Hippocampus
After Scoville and Milner (1957) implicated the MTL in anterograde amnesia,
subsequent mice studies sought to highlight the specific function of the
Hippocampus. Clark & Isaacson (1965) showed that in bilateral hippocampal
lesions in rats failed to suppress behaviours that were no longer reinforced, ie.
no longer appropriate.


Spatial memory tasks have also been attributed to the dorsal hippocampus
via a number of lesion studies. (Bannerman et al, 2002). Because the ventral
hippocampus has a number of projections into both the hypothalamus and
the amygdala it was theorized that the hippocampus might play a role in
contextual behaviours: such as contextually conditioned
freezing. (Bannerman et al, 2004).


Richmond et al. (1999) had shown that cell loss in the dorsal hippocampus did
not elicit this effect.
The Medial Temporal Lobe
Scoville and Milner (1957) published a case where bilateral MTL lesions led to
anterograde amnesia. (Anderson et al 2009).
Fell et al (2001) used EEG recordings in epilepsy patients to implicate the
MTL in successful memory formation. They found that successful declarative
memory formation is determined by a direct and temporary rhinal-
hippocampal gamma synchronization.


References
Anderson, KL et. al. (2009) Theta Oscillations Mediate Interaction between
Prefrontal Cortex and Medial Temporal Lobe in Human Memory. Cerebral
Cortex. 20(7). p 1604-1612.

Bannerman DM, et. al. (2002) A double dissociation of function within the
hippocampus: spatial memory and hyponeophagia. Behavioural Neuroscience.
116(5). p 884901.

Bannermana DM, et. al. (2004) Regional dissociations within the hippocampus
memory and anxiety. Neuroscience & Biobehavioral Reviews. 28(3). p 273-283.

Clark CVH, Isaacson RL (1965) Effect of bilateral hippocampal ablation on DRL
performance. Journal of Computational Physiology & Psychology. 59. p 137140.

Fanselow MS (1984) What is conditioned fear? Trends in Neuroscience. 7(12). p
460462.

Fell et al (2001) Human memory formation is accompanied by
rhinalhippocampal coupling and decoupling. Nature Neuroscience. 4(12) p
1259 1264.

Herculano-Houzel, et. al. (1999) Precisely synchronized oscillatory firing
patterns require electroencephalographic activation. Journal of Neuroscience.19
p 3992-4010.

Jones MW, Wilson MA (2005) Theta Rhythms Coordinate Hippocampal
Prefrontal Interactions in a Spatial Memory Task. PLoS Biology. 3(12): e402.
doi:10.1371

Par D, Collins DR, Pelletier JG (2002) Amygdala oscillations and the
consolidation of emotional memories. Trends in Cognitive Sciences. 6(7). p 306-
314

Par D, Gaudreau H, (1996) Projection cells and interneurons of the lateral and
basolateral amygdala: distinct firing patterns and differential relation to theta
and delta rhythms in conscious cats. Journal of Neuroscience. 16(1996). p 334-
335.

Paz R et. al. (2006) Emotional enhancement of memory via amygdala-driven
facilitation of rhinal interactions. Nature Neuroscience. 9, p 1321 - 1329 (2006)
Published online: 10 September 2006 | doi:10.1038/nn1771

Richmond MA, et. al. (1999) Dissociating context and space within the
hippocampus: effects of complete, dorsal and ventral excitotoxic lesions on
conditioned freezing and spatial learning. Behavioural Neuroscience. 113(6). p
11891203.

Scoville WB, Milner B (1957) Loss of recent memory after bilateral hippocampal
lesion. Journal Neurology, Neurosurgery & Psychiatry. 20(1). p 1121.

Shadlen MN, Movshon JA (1999) Synchrony Unbound: A Critical Evaluation of
the Temporal Binding Hypothesis. Neuron 24(1), p 67-77.

Singer W (2007) Binding by synchrony. Scholarpedia. 2(12). E.1657. revision
#124403, http://www.scholarpedia.org/article/Binding_by_synchrony