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It is generally assumed that neurons are the basic computational units of the brain and that
all information is carried by spikes, patterns of electrochemical activity that travel along
and between neurons.

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As a first guess, we might think that the brain transmits and processes information using
strings of single neurons. However, this set up is unreliable for two reasons.

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First, neurons die throughout your life and are generally not replaced.

Second, single neurons respond somewhat randomly to their inputs.

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If we feed the same input into a neuron, say a small electrical current as symbolized here
on the left, several times, we will get slightly different electrical outputs each time. Notice
the addition or subtraction of spikes in these two responses highlighted in red.

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Either of these sources of unreliability, neuron death or response variability, can destroy
the functionality of a string of neurons. However, there are more robust ways to organize
neurons.

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One solution is to use groups of neurons arranged in a fully-connected, feed-forward chain,
that is a sequence of groups of neurons with each of the neurons in one group connected
to all of the neurons in the next group.

First, these chains reduce the variability in the response of single neurons since…

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…the more inputs to a neuron that are active…

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…and the more synchronous those inputs, the more reliably a neuron will fire.

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So if we synchronously activate the first group…

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…the activity will synchronously and reliably activate the second group…

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…which will synchronously and reliably activate the third group, and so on. This highly
reliable mode of operation gives these structures the nickname “synfire” chains
(synchronously firing chain).

However, even if a neuron does happen to die or unexpectedly fail to fire…

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…the redundancy in connections…

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…allows a synfire chain to…

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…propagate a signal anyways.

So synfire chains seem useful, but do they actually occur in real brains and if so, what do
they look like?

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My research this summer attempted to predict the likelihood and shape of synfire chains in
the brain. Now, experimentally this is quite difficult. However, as someone with a
background in mathematics, what I can do is create neural network models inspired by
biological data and ask: what are the most common synfire chains in these networks? By
doing so, I can suggest (1) which types of networks are the best environments for
producing synfire chains and (2) which types of synfire chains experimentalists should look
for in particular types of networks.

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Although I worked on a couple different models, I only have time today to present one and
I’ve chosen what I think to be the most biologically plausible model I’ve worked on.

There are two main things one needs to specify a network – the neurons and the
connections. As for the neurons, I chose a 2-dimensional circular grid.

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As for the connections, I chose to make neurons that are close together more likely to be
connected than those that are far apart, but besides this dependence on distance, the
connections were completely random. This type of connectivity is commonly referred to as
a locally connected random network.

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To be more precise about the connection probability, the probability that two neurons are
connected was given by a 2D Gaussian function with the same standard deviation in either
spatial direction. For those to whom that sentence was complete nonsense, this plot here
represents the probability of connection to a neuron in the center (at the red dot). The x
and y axes indicate x and y distances from the neuron in the center and the height of the
plot represents the probability of connection. As we move away from the neuron, we see
that the connection probability quickly becomes smaller.

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So the way I went about finding the most common chains was to start off with the
following question. Given some random subset of neurons as the prospective first group in
a synfire chain…

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…what is the most likely size and shape of the next group? To answer this question, we
need to determine the probability that neurons at various locations in the network receive
connections from all of the neurons in this first group.

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In our model, connections are independent of one another, so the probability that a neuron
is connected to all of the neurons in a group is the product of the probabilities for that
neuron being connected to each neuron in the group.

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Now, we know exactly what those connection probabilities look like – they’re the Gaussian
bumps I showed you a few slides ago. If we multiply many of them with different centers
but the same width…

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…what we get is a smaller, sharper Gaussian bump located at the average position of all the
neurons in the first group. The interpretation is that neurons near the center of our first
group are most likely to participate in the second group and the probability of a neuron
being in the second group decreases quickly as we consider neurons further and further
from the first group’s center.

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What all of this means is that the most common shape for the next group will be a smaller,
circular cluster of neurons.

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And if we apply these results iteratively and don’t allow neurons to participate in two
consecutive groups, we find out that most likely shape of a synfire chain is a series of
circular groups that become smaller and smaller and stabilize at some minimum radius.

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Now, what can we conclude from all of this?

First, “naturally occurring” synfire chains in LCRN are confined in space because
consecutive groups tend to be centered around the same point and become progressively
smaller. (I say “naturally occurring” because we didn’t have any learning in our network.)
Second, because consecutive groups of a chain tend to stay in one place, these chains are
unlikely to be useful for carrying information from place to place in the brain. Instead,
these chains seem more useful for local processing of information.
Finally, long-range chains that are useful for reliable signal propagation likely require
networks with learning in which controlled stimuli are applied in a way to encourage the
growth of long-range synfire chains.

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As for future work and improvements to the model, I’ve so far only considered single layers
of neurons but a more realistic model would include multiple layers and realistic
connectivity between those layers.

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The model I discussed today also did not include learning which we know plays a major role
in the self-organization of connectivity patterns in the brain.

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Finally, there are a few experimental investigations of activity patterns in actual brains that
may indicate synfire activity. After a few of the improvements I just mentioned, it would
be interesting to compare the model’s predictions to actual biological data.

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Finally, I’d like to thank my Kwabena Boahen and Peiran Gao for their guidance in my
research, Matt Goldstein for his guidance in putting together this presentation, Tenea
Nelson & the SSRP staff for this fantastic opportunity, and finally Nick Steinmetz, Samir
Menon, and the rest of the grad students in the Clark Center for letting me pester them
with endless streams of neuroscience questions for 8 weeks.

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