<p><strong>TL;DR</strong>: We don't know whether the brain really uses predictive coding or not. But neurally computing an error signal on a small scale is possible (see below).</p>

<p>Predictive coding is an <em>hypothesis</em> for a putative signal-processing mechanism used in vertebrate brains. As things stand presently (2017), mapping the hypothesis of predictive coding onto known neural structures and responses is a matter of active research (e.g. <a href="https://arxiv.org/abs/1612.05321" rel="nofollow noreferrer" title="Choi et al. 2016">Choi et al. 2016</a>, <a href="http://doi.org/10.1016/j.neuron.2016.09.057" rel="nofollow noreferrer" title="Zmarz &amp; Keller 2016">Zmarz &amp; Keller 2016</a>, <a href="https://doi.org/10.1038/nn.4197" rel="nofollow noreferrer" title="Roth et al 2016">Roth et al 2016</a>).</p>

<p>On a small-scale neural level, an error signal could be partially computed through converging an excitatory "prediction" signal and an inhibitory "evidence" signal onto a single neuron. This is only a partial error signal, since it is rectified by the spiking threshold of the post-synaptic neuron.</p>

<p>The main issue is that the forward and backwards pathways need to follow very precise wiring patterns, and be quite tightly aligned, for the simple formulations of predictive coding to work. That doesn't mesh well with intuitions of what is biologically reasonable.</p>

<p>There are some approaches to map a predictive coding framework onto other mechanisms that may be more biologically plausible (e.g. <a href="https://doi.org/10.3389/neuro.10.004.2008" rel="nofollow noreferrer" title="Spratling 2008">Spratling 2008</a>).</p>

CC BY-SA 3.0
2017-04-19T16:02:26.750Z
2017-04-19T16:02:26.750Z
Dylan Richard Muir
<p>I provided an answer to a similar question <a href="https://cogsci.stackexchange.com/questions/16400/is-neurogenesis-driven-by-prediction-error/16989#16989">here</a> that limitedly deals with the role of biological prediction errors.</p>
<p>Here's an excerpt of that answer:</p>
<blockquote>
<p>...to answer this properly, we must first make it clear that there are potentially dozens, hundreds, or an arbitrarily high number of other &quot;prediction error types&quot; in use by the brain. Here are just a few major ways, hypothetically:</p>
<ul>
<li>Lots of different neurotransmitters (e.g. dopamine)</li>
<li>The opening/closing of various ion channel species that regulate the membrane potential</li>
<li>Synaptic vescicles/receptors</li>
<li>Neuronal firing rates (as in <a href="https://en.wikipedia.org/wiki/Bursting" rel="nofollow noreferrer">bursting</a>, a rapid succession of action potentials)</li>
<li>Temporal coding (relative firing times to the firing of other neurons)</li>
<li>And I can think of 10 other more-subtle and harder to explain possibilities, but that are just as important, off the top of my head</li>
</ul>
<p>Keep in mind that each neuron also seems to have its own differentiated mechanisms for, both, interpreting and signaling prediction error. This complicates things further. For instance, one neurotransmitter may communicate prediction error to one particular neuron, but has no effect (or a different effect) on a different neuron. It may even be that neurotransmitter X must be present while temporal code Y happens for the event to be interpreted as a prediction error.</p>
</blockquote>
<p>The study of biological mechanisms for prediction error is a very complicated thing that has no simple interpretation, as opposed to what you find in artificial approaches (presumably, like your PredNet example). While we have not yet uncovered how the brain computes or uses these mechanisms to encode and communicate prediction errors across neurons, what is obvious is that the brain has to be doing some kind of prediction error coding. However, if we try to oversimplify what the brain is doing, we are likely to not have a very intelligent model. The fragility of predictions, susceptibility to error from data complications, and limited nature of current artificial intelligence implementations lends to the idea that there is a lack of good ideas about how to implement and fully utilize prediction error.</p>
<p>The work I do is of a theoretical nature so your question is right up my alley. I have some unique ideas on how various biological prediction errors may work, but it requires a lot of background to understand. Unfortunately, I also have not published my ideas so they are definitely not peer reviewed. That makes me somewhat reluctant to mention my personal ideas as an answer.</p>
<p>It's possible that a kind of prediction error is used in all the listed mechanisms to fine tune the respective properties. Each mechanism likely has a unique role that is central to intelligence- from not only prediction errors encountered in the environment, but even to intrinsic behaviors and being able to predict the outcome of its own actions.</p>

CC BY-SA 3.0
2017-03-24T21:23:13.203Z
2017-03-25T23:07:06.227Z
2020-06-17T09:55:31.633Z
aiwyn
<p>I am quite inspired by the results obtained by <a href="https://coxlab.github.io/prednet/" rel="noreferrer">PredNet</a>, which implements a predictive coding model using artificial neural networks. They compute the <em>prediction error</em> as a simple subtraction, and then split the positive and negative into two different feature maps.</p>

<p>Within the predictive coding framework it is believed that biological brains also uses the prediction error to communicate between different areas. <strong>How is this error signal computed in biological brains?</strong>.</p>

<p><strong>Update</strong></p>

<p>I am especially interested in the mechanical aspects, such as <em>how doe does the brain compare a prediction with the true signal?</em>. Synthetic approaches generally just use a pixel-wise error, and I am wondering if the brain does something analogously.</p>

CC BY-SA 3.0
2017-03-20T13:09:04.397Z
Hi Toke, welcome at CogSci and what an interesting question! I cannot give a full answer because I don't have the references to back things up, but perhaps I can give you a nice head start. I believe this has to do with dopamine. People have particular expectations, usually based on earlier experiences. When the expectations are met a burst of dopamine is released giving you a pleasure sensation. However, when the expectation is not met the dopamine production is decreased. The computation of expectation vs outcome happens in the ventro-lateral prefrontal cortex and the ventro-medial PFC a.o.
Robin Kramer-ten Have
CC BY-SA 3.0
2017-03-20T13:55:16.677Z
Thank you. I was especially interested in the mechanistic aspects, and have updated the question.
Faur
CC BY-SA 3.0
2017-03-20T17:59:41.740Z
@RobinKramer as far as I know a) dopaminergic neurons only correlate with _reward_ prediction errors, so they are not representing the low-level error signals (but it could be that they represent the topmost error signal which is then fed back to lower levels) and b) they correlate with the prediction _error_, so it is not true that a burst of dopamine is released when expectations are met but rather when they are exceeded (see e.g. here: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4826767/)
awakenting
CC BY-SA 3.0
2017-03-20T18:15:39.850Z
short answer to this question is: we don't know. we don't even know if the brain computes such an error signal at all, although of course there are some indications.but I think it's quite safe to say that the brain does not compute a pixel-wise error as it doesn't deal with pixels in the first place as is nicely described in this question: http://cogsci.stackexchange.com/questions/1125/where-is-the-visual-image-that-we-see-finally-assembled
awakenting
<vision><neural-network><communication><signal-processing><encoding>
How is the biological error signal in predictive coding computed?
Faur
# How is the biological error signal in predictive coding computed?

(Asked by: username_0 on 2017-03-20)

I am quite inspired by the results obtained by [PredNet](https://coxlab.github.io/prednet/), which implements a predictive coding model using artificial neural networks. They compute the *prediction error* as a simple subtraction, and then split the positive and negative into two different feature maps.

Within the predictive coding framework it is believed that biological brains also uses the prediction error to communicate between different areas. **How is this error signal computed in biological brains?**.

**Update**

I am especially interested in the mechanical aspects, such as *how doe does the brain compare a prediction with the true signal?*. Synthetic approaches generally just use a pixel-wise error, and I am wondering if the brain does something analogously.

**Tags:** vision, neural-network, communication, signal-processing, encoding

> **Comments:**
> **username_1**: Hi Toke, welcome at CogSci and what an interesting question! I cannot give a full answer because I don't have the references to back things up, but perhaps I can give you a nice head start. I believe this has to do with dopamine. People have particular expectations, usually based on earlier experiences. When the expectations are met a burst of dopamine is released giving you a pleasure sensation. However, when the expectation is not met the dopamine production is decreased. The computation of expectation vs outcome happens in the ventro-lateral prefrontal cortex and the ventro-medial PFC a.o.  
> **username_0**: Thank you. I was especially interested in the mechanistic aspects, and have updated the question.  
> **username_2**: @RobinKramer as far as I know a) dopaminergic neurons only correlate with _reward_ prediction errors, so they are not representing the low-level error signals (but it could be that they represent the topmost error signal which is then fed back to lower levels) and b) they correlate with the prediction _error_, so it is not true that a burst of dopamine is released when expectations are met but rather when they are exceeded (see e.g. here: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4826767/)  
> **username_2**: short answer to this question is: we don't know. we don't even know if the brain computes such an error signal at all, although of course there are some indications.but I think it's quite safe to say that the brain does not compute a pixel-wise error as it doesn't deal with pixels in the first place as is nicely described in this question: http://cogsci.stackexchange.com/questions/1125/where-is-the-visual-image-that-we-see-finally-assembled  

---

## 2 answers

### Answer by username_3 on 2017-04-19. Score: 2

**TL;DR**: We don't know whether the brain really uses predictive coding or not. But neurally computing an error signal on a small scale is possible (see below).

Predictive coding is an *hypothesis* for a putative signal-processing mechanism used in vertebrate brains. As things stand presently (2017), mapping the hypothesis of predictive coding onto known neural structures and responses is a matter of active research (e.g. [Choi et al. 2016](https://arxiv.org/abs/1612.05321 "Choi et al. 2016"), [Zmarz & Keller 2016](http://doi.org/10.1016/j.neuron.2016.09.057 "Zmarz & Keller 2016"), [Roth et al 2016](https://doi.org/10.1038/nn.4197 "Roth et al 2016")).

On a small-scale neural level, an error signal could be partially computed through converging an excitatory "prediction" signal and an inhibitory "evidence" signal onto a single neuron. This is only a partial error signal, since it is rectified by the spiking threshold of the post-synaptic neuron.

The main issue is that the forward and backwards pathways need to follow very precise wiring patterns, and be quite tightly aligned, for the simple formulations of predictive coding to work. That doesn't mesh well with intuitions of what is biologically reasonable.

There are some approaches to map a predictive coding framework onto other mechanisms that may be more biologically plausible (e.g. [Spratling 2008](https://doi.org/10.3389/neuro.10.004.2008 "Spratling 2008")).


---

### Answer by username_4 on 2017-03-24. Score: 2

I provided an answer to a similar question [here](https://cogsci.stackexchange.com/questions/16400/is-neurogenesis-driven-by-prediction-error/16989#16989) that limitedly deals with the role of biological prediction errors.

Here's an excerpt of that answer:

> ...to answer this properly, we must first make it clear that there are potentially dozens, hundreds, or an arbitrarily high number of other "prediction error types" in use by the brain. Here are just a few major ways, hypothetically:
>
> * Lots of different neurotransmitters (e.g. dopamine)
> * The opening/closing of various ion channel species that regulate the membrane potential
> * Synaptic vescicles/receptors
> * Neuronal firing rates (as in [bursting](https://en.wikipedia.org/wiki/Bursting), a rapid succession of action potentials)
> * Temporal coding (relative firing times to the firing of other neurons)
> * And I can think of 10 other more-subtle and harder to explain possibilities, but that are just as important, off the top of my head
>
> Keep in mind that each neuron also seems to have its own differentiated mechanisms for, both, interpreting and signaling prediction error. This complicates things further. For instance, one neurotransmitter may communicate prediction error to one particular neuron, but has no effect (or a different effect) on a different neuron. It may even be that neurotransmitter X must be present while temporal code Y happens for the event to be interpreted as a prediction error.

The study of biological mechanisms for prediction error is a very complicated thing that has no simple interpretation, as opposed to what you find in artificial approaches (presumably, like your PredNet example). While we have not yet uncovered how the brain computes or uses these mechanisms to encode and communicate prediction errors across neurons, what is obvious is that the brain has to be doing some kind of prediction error coding. However, if we try to oversimplify what the brain is doing, we are likely to not have a very intelligent model. The fragility of predictions, susceptibility to error from data complications, and limited nature of current artificial intelligence implementations lends to the idea that there is a lack of good ideas about how to implement and fully utilize prediction error.

The work I do is of a theoretical nature so your question is right up my alley. I have some unique ideas on how various biological prediction errors may work, but it requires a lot of background to understand. Unfortunately, I also have not published my ideas so they are definitely not peer reviewed. That makes me somewhat reluctant to mention my personal ideas as an answer.

It's possible that a kind of prediction error is used in all the listed mechanisms to fine tune the respective properties. Each mechanism likely has a unique role that is central to intelligence- from not only prediction errors encountered in the environment, but even to intrinsic behaviors and being able to predict the outcome of its own actions.


---

