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mechanism is able to account for the behavioural data, based on the iring rate of a
neuronal population tuned to ITD in a similar manner as barn owl neurons.
4.5
Probabilistic Population Codes
The PV is not the only method for decoding sensory responses from a population 
of neurons. An alternative scheme is based on a probabilistic population code
(PPC; Ma et al.  2006 ). The PPC assumes that neuronal populations encode proba-
bility distributions through their joint iring rate tuning curves. As a result, the
entire tuning curve of the neuronal population and not just the preferred direction
is used in the decoding process. Let k = …
12 represent the response in a
single trial of N neurons to a ixed sound source direction θ . The posterior distribu-
tion has the form
(, ,
kk k N
,
)
p
(|)
q
k
p
( |)(),
k
q
p
q
(4.12)
where p ( θ ) is the same as in Eq. ( 4.6 ) and p ( k | θ ) is a distribution which models the
probability of a neuronal response given the stimulus.
If we assume that the neurons representing p ( k | θ ) are independent and Poisson, 
then the probabilistic population code for the distribution p ( k | θ ) has the form
r
()
!
q
k
N
n
1
r
()
q
(4.13)
p
(|
k q
)
=
n
e
,
n
k
n
=
n
where k n is the response of neuron n and r n ( θ ) is its tuning function. We model the
tuning functions similarly as in Sect.  4.4 , r n ( θ ) = r n (ITD( θ )) using Eqs. ( 4.9 ) and ( 4.3 ),
but with a uniform distribution of preferred directions over the unit circle instead of
being normally distributed. Note that the right-hand side of Eq. ( 4.13 ) is formed by
taking products of Eq. ( 4.8 ) with k i replacing k and r i ( θ ) replacing λ , since we assume
independent Poisson neurons.
Based on this probabilistic population code, an alternative decision rule to averag-
ing over unit vectors is to pick the azimuth that maximizes the posterior probability.
This rule is called the maximum a posteriori probability (MAP) rule and has the form 
ˆ ()
q
k
= arg
max(| .
p
q
k
q
The result of this estimation method based on the probabilistic population code is
given in Fig. 4.3 d as the grey starred curve. It is signiicantly different from the
population vector result and does not match the behavioural data very well. On the
other hand, a probabilistic population code has been successfully used to explain the
sensory integration of visual and vestibular cues in neurons of the monkey visual
cortex using a slightly different decoding mechanism (Fetsch et al. 2011 ).
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