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X
t
dt + dW
t
, X
0
= 0 (1)
where W
t
is a standard Wiener process, is a suitable time scale, and
T = inf{t > 0 : X
t
S} (2)
is the rst-passage time of X
t
through the constant threshold S. It is convenient to reformulate to the equivalent
dimensionless form
d
X
t
S
X
t
S
+
S
S
d
W
t
(3)
2
or
dY
s
= (Y
s
+) ds + dW
s
, Y
0
= 0 (4)
where
s =
t
, Y
s
=
X
t
S
, W
s
=
W
t
, =
S
, =
S
(5)
and T/ = inf{s > 0 : Y
s
1}. Therefore, without loss of generality, all considerations in the following will
be related to the dimensionless process Y
s
and its rst crossing of the level 1.
The solution process Y
s
is normally distributed with conditional moments
E[Y
s
|Y
0
= 0] = (1 e
s
) (6)
Var[Y
s
|Y
0
= 0] =
1
2
2
(1 e
2s
) (7)
The asymptotic mean is and the asymptotic variance is
2
/2 as s .
As suggested in (Wan & Tuckwell 1982) it is convenient to distinguish between two crudely dened parts of
the parameter space called the suprathreshold regime and the subthreshold regime. The suprathreshold regime
is dened as that for which 1, that is, the asymptotic mean is far above the threshold implying that the
rst-passage times are relatively regular (deterministic behavior - which means that there are crossings also in
the absence of noise). The subthreshold regime is dened as that for which 1 implying that crossings of
the threshold are caused only by random uctuations (stochastic or Poissonian behavior). The term Poissonian
behavior indicates that when 1 (measured in terms of ), the rst-passage times achieve characteristics
of a Poisson point process (Nobile, Ricciardi, & Sacerdote 1985, Wan & Tuckwell 1982). What is left over of
the parameter space is herein denoted as the threshold regime in which the asymptotic mean does not deviate
much from 1. The regimes are illustrated in Fig. 1.
0 0.5 1 1.5 2
!1
0
1
2
3
4
s
O
U
p
r
o
c
e
s
s
subthreshold regime
suprathreshold regime
threshold regime
Figure 1. Illustration of the different regimes. The dashed line indicates the asymptotic mean .
The goal of this paper is to present a method to estimate the unknown parameters and from independent
observations of T/: s
1
, . . . , s
N
, where N is the number of measurements. In neuronal modeling, the time scale
3
is an intrinsic parameter that characterizes the neuron irrespectively of the incoming signal (the time constant
of the cell membrane), whereas and are input parameters describing signals that the neuron receives from
surrounding neurons (Tuckwell & Richter 1978). The value of is often given by physiological considerations
( 520 msec), and it is only relevant as a guidance to assess a reasonable value range of the input parameters.
As it is seen from the differential equation (3), the time scale can be freely chosen whereupon the parameters
and are uniquely obtained from estimates of and . The estimates of and obtained by the method
are nally compared with the results obtained from a previously proposed method for estimating the same
parameters given that they are in the suprathreshold regime (Ditlevsen & Lansky 2005).
Let f
T/
(s) be the probability density of T/. An exact expression for f
T/
(s) is only known for = 1 and
is
f
T/
(s)
=1
=
2e
2s
(e
2s
1)
3/2
exp
2
(e
2s
1)
(8)
Moreover, an asymptotically exact expression can be obtained as and together under constant
ratio /
2
. Then the stochastic differential equation (4) asymptotically approaches the differential equation
dY
s
= ds + dW
s
for a Wiener process with corresponding rst-passage density
f
T/
(s)
,
=
1
2
2
s
3
exp
(1 s)
2
2
2
s
(9)
i.e., the inverse Gaussian distribution. If the limit passage is made by letting with , , and S kept
xed, the corresponding asymptotic density function for T is
f
T
(t )
=
S
2
2
t
3
exp
(S t )
2
2
2
t
(10)
In these two special cases the maximum likelihood estimators of and
2
are
= 1 :
2
=
1
N
N
i =1
2
e
2s
i
1
(11)
, : =
1
s
2
=
1
N
N
i =1
1
s
i
1
s
(12)
where s =
1
N
N
i =1
s
i
. Otherwise no explicit expressions for the density are known, and other ways of estimat-
ing have to be found. One approach is to use moments or functions of moments of the rst-passage time. In
(Inoue, Sato, & Ricciardi 1995, Ricciardi & Sato 1988) series expressions for the moments of T were derived.
In particular
E[T/] =
1
2
n=1
2
n
n!
(1 )
n
()
n
n
2
(13)
However, this expression is difcult to work with, especially if 1 (strongly suprathreshold) because of the
canceling effects in the alternating series. In (Ditlevsen & Lansky 2005) the moments
E[e
T/
] =
1
if > 1 (14)
E[e
2T/
] =
2
2
/2
( 1)
2
2
/2
if 1 >
2
(15)
4
are given. The rst moment is only nite in the suprathreshold regime, and for the second moment to be nite it
is further required that the asymptotic standard deviation of Y
s
is smaller than the distance between the threshold
and the asymptotic mean of Y
s
.
Estimators for the parameters and
2
from observations of rst-passage times in the suprathreshold regime
are in (Ditlevsen & Lansky 2005) proposed as
=
Z
1
Z
1
1
,
2
=
2(Z
2
Z
2
1
)
(Z
2
1)(Z
1
1)
2
(16)
where
Z
1
=
1
N
N
i =1
e
s
i
, Z
2
=
1
N
N
i =1
e
2s
i
(17)
based on the exact expressions (14) and (15). These simple estimators are only valid in suprathreshold regime,
and it remains to determine the regime from data, a problem which is not straightforward.
3 FORTET INTEGRAL EQUATION
The stationary OU process solution to the stochastic differential equation (4) has the mean value , variance
2
/2, and correlation function exp(s). By introducing the condition that Y
0
= 0, a non-stationary Gaussian
process with mean value function E[Y
s
| Y
0
= 0] = [1 exp(s)], and variance function Var[Y
s
| Y
0
= 0] =
[1 exp(2s)]
2
/2 is obtained. In the following we will denote this non-stationary process by Y(s) without
explicitly mentioning the condition Y
0
= 0.
Let f (s) be the density function for the time t / from zero to the rst crossing of the level 1 by Y. The
probability
P[Y
s
> 1] =
(1 e
s
) 1
1 e
2s
/
(18)
where () is the standard normal distribution function, can alternatively be calculated by the transition integral
P[Y
s
> 1] =
s
0
f (u)
1
/
2
1e
(su)
1e
2(su)
du (19)
because
P[Y
s
> 1 | Y
u
=1] =
E[Y
s
| Y
u
=1] 1
D[Y
s
| Y
u
]
+e
(su)
(1 ) 1
/
1e
2(su)
(20)
where D[|] indicates conditional standard deviation. Thus we have obtained an integral equation for the
unknown density function f (s). It is difcult to obtain a solution except for the case = 1, where the factor to
f (u) under the integral sign becomes the constant 1/2. Then the distribution function F(s) of the rst-passage
time simply becomes
F(s) = 2
e
s
1 e
2s
, s > 0 (21)
5
and by differentiation
f (s) =
e
s
2
2
(1 e
2s
)
3/2
e
s
1 e
2s
(22)
where () is the standard normal density function. This is the density (8).
If and are made large under constant ratio /
2
it is seen that the range of s in which there is essential
variation of the right side of (18) narrows down closer and closer to zero implying that (1 e
s
) s and
1 e
2s
/
s asymptotically as and increase. Thus the integral equation takes the limit form
s 1
s
0
f (u)
s u
du (23)
which by differentiation with respect to s gives the equation
s +1
2s
s 1
=
1
2
f (s) +
s
0
f (u)
2
s u
s u
du (24)
It is not an easy task to solve this integral equation directly. However, we may check whether the density
function (9) satises the equation. Indeed, it is the case. The calculations are shown in Appendix 1.
4 PARAMETER ESTIMATION
The integral equation
(1 e
s
) 1
1 e
2s
/
s
0
f (u)
1
/
1 e
(su)
1 +e
(su)
du (25)
obtained from (18) and (19) can be used to estimate the two parameters and as explained in the following.
The resulting estimators do not depend on the regime.
Consider the ordered sample s
1
s
2
. . . s
N
of independent observations of T/. For given values of
the parameters the right hand side of (25) can be estimated at s from the sample by the average
RHS(s)
1
N
max{n:s
n
s}
i =1
1
/
1 e
(ss
i
)
1 +e
(ss
i
)
(26)
since it is the expected value of
1
U[0,s]
1
/
1 e
(sU)
1 +e
(sU)
(27)
with respect to the distribution of U = T/, where 1
U[0,s]
is the indicator function for the event {U [0, s]}.
Upon dividing both sides of the integral equation (25) by [(1)/(/
1 2/(/
Y
s
dW
s
(30)
In case 2(/)
2
1 the process stays positive at all times (a.s.) (in Fellers terminology the boundary at zero is
a so-called entrance boundary dened as a boundary from which the process can start but not return to (Karlin
& Taylor 1981), p.234, and the Feller process becomes asymptotically stationary as s . An illustration of
a sample path is shown in Fig. 4. Feller has shown that the transition probability distribution is a non-central
2
-distribution with = 4(/)
2
degrees of freedom and with conditional mean
E[Y
s
|Y
0
= y
0
] = +(y
0
)e
s
(31)
and variance
Var[Y
s
|Y
0
= y
0
] =
2
2
(1 e
s
)
1 +
2y
0
e
s
(32)
see e.g. (Cox, Ingersoll, & Ross 1985). The standard form of the non-central
2
-distribution with degrees of
freedom and non-centrality parameter has mean + and variance 2 + 4. The conditional mean (31) and
conditional variance (32) are both obtained if the afnity factor 1/a(s) =
2
(1 e
s
)/(4) is applied to the
standardized variable and the non-centrality parameter is (s, y
0
) = (4y
0
/
2
)[e
s
/(1 e
s
)]. Thus
P(Y
s
y | Y
0
= y
0
) = F
2[a(s)y, , (s, y
0
)] (33)
where F
2[a(s), , (s, y
0
)] (34)
=
s
0
f (u){1 F
n=0
1 y
n+1
0
(n +1)
n
i =0
( +i
2
/2)
(35)
(Ditlevsen & Lansky 2006) give the moments
E[e
T/
] =
y
0
1
if > 1 (36)
E[e
2T/
] =
2( y
0
)
2
+
2
( 2y
0
)
2( 1)
2
+
2
( 2)
if (37)
1 +2(/)
2
< 1 +2( 1)/
2
(38)
that otherwise are innite. Assuming that the data are in the allowed parameter region, moment estimators of
the parameters are then obtained from equations (36) and (37) as
=
Z
1
y
0
Z
1
1
(39)
and
2
=
2(1 y
0
)
2
(Z
2
Z
2
1
)
2(Z
1
1)(Z
2
y
0
) (Z
1
y
0
)(Z
2
1)
(40)
where Z
1
and Z
2
are given by (17).
Just as for the OU process 100 times 100 simulations are run for the Feller process. The statistics of the
estimates of and are given in Table 2. The quality and distribution properties are about the same as for the
OU process.
The less good accuracy corresponding to = 11 are due to numerical difculties in connection with com-
putation of
2
-distribution values.
7 MODEL TESTING
With the parameter estimates from samples of 100 rst-passage times of the OU process simulated for the
selected values of and for =1, the left side of the integral equation (25) is in Fig. 5 compared to the right
side as estimated by the respective samples.
The diagrams in Fig. 6 show the error of t in terms of the difference between the two normalized sides
of the integral equation (34) (solid line) and of the integral equation (25) (dashed line) corresponding to the
11
Table 2. Values of and used in the simulations of Feller process realizations, and the corresponding averages of the
samples of 100 parameter estimates of and the sample standard deviation (SSD), using (39), (40) and (29), where
relevant. All rst-passage time samples contain 100 simulated observations each.
statistics of 100 estimates:
regime = 1 average SSD
=
subthr. 0.8 0.79 0.09
threshold 1 1.10 0.06 1.00 0.08
suprathr. 2 1.99 0.10 1.98 0.11
suprathr. 3 2.97 0.09 2.95 0.10
suprathr. 4 3.94 0.12 3.90 0.11
Wiener 11 10.96 0.11 9.88 0.15
s
0
f (u)
2
s u
s u
du =
1
4
s
0
1
u
u(su)
exp
(1u)
2
+
2
(su)u
2
2
u
du
=
4
2
exp
(2s)s
2
2
s
s
0
exp
1/(2
2
u)
u(s u)
du
=
4
2
s
exp
(2s)s
2
2
s
1
0
e
/w
w
w(1w)
dw
13
where = 1/(2
2
s) and where the substitution w = u/s is used. The denite integral is manipulated further
as follows:
1
0
e
/w
w
w(1 w)
dw =
1
e
v
v 1
dv =
2e
0
e
u
d
u =
2e
0
e
1
2
(
2 u)
2
d
u
=
s exp
1
2
2
s
s
0
f (u)
2
su
su
du =
s
2s
s1
Adding f (s)/2 we obtain the left side of the integral equation (24). This concludes the proof.
REFERENCES
Aalen, O. & H. Gjessing (2004). Survival models based on the Ornstein-Uhlenbeck process. Lifetime data analysis 10:407423.
Alili, L., P. Patie, & J. Pedersen (2005). Representations of the rst hitting time density of an Ornstein-Uhlenbeck process. Stochastic
Models 21:967980.
Cox, J., J. Ingersoll, & S. Ross (1985). A theory of the term structure of interest rates. Econometrica 53:385407.
Ditlevsen, S. & P. Lansky (2005). Estimation of the input parameters in the Ornstein-Uhlenbeck neuronal model. Phys. Rev.
E 71(011907).
Ditlevsen, S. & P. Lansky (2006). Estimation of the input parameters in the Feller neuronal model. Phys. Rev. E 73(061910).
Feller, W. (1951). Diffusion processes in genetics. In J. Neyman (ed.), Proceedings of the second Berkeley symposium on mathematical
statistics and probability, pp. 227246. University of California Press, Berkeley.
Giorno, V., P. Lansky, A. Nobile, & L. Ricciardi (1988). Diffusion approximation and rst-passage-time problem for a model neuron.
Biol. Cybern. 58:387404.
Inoue, J., S. Sato, & L. Ricciardi (1995). On the parameter estimation for diffusion models of single neurons activity. Biol. Cy-
bern. 73:209221.
Karlin, S. & H. Taylor (1981). A second course in stochastic processes. Academic Press, San Diego.
Lansky, P., L. Sacerdote, & F. Tomasetti (1995). On the comparison of Feller and Ornstein-Uhlenbeck models for neural activity. Biol.
Cybern. 73:457465.
Linetsky, V. (2004). Computing hitting time densities for CIR and OU diffusions: applications to mean-reverting models. J. Comput.
Finance 7:122.
Nobile, A., L. Ricciardi, & L. Sacerdote (1985). Exponential trends of Ornstein-Uhlenbeck 1st-passage-time densities. J. Appl.
Prob. 22:360369.
Ricciardi, L. & S. Sato (1988). First-passage-time density and moments of the Ornstein-Uhlenbeck process. J. Appl. Prob. 25:4357.
Tuckwell, H. & W. Richter (1978). Neuronal interspike time distributions and the estimation of neurophysiological and neuroanatom-
ical parameters. J. Theor. Biol. 71:167180.
Wan, F. & H. Tuckwell (1982). Neuronal ring and input variability. J. Theoret. Neurobiol. 1:197218.
14
!3 !2 !1 0 1 2
!0.06
!0.04
!0.02
0
0.02
0.04
0.06
!=1
log s
e
r
r
o
r
!3 !2.5 !2 !1.5 !1 !0.5 0 0.5
!0.03
!0.02
!0.01
0
0.01
0.02
0.03
!=2
log s
e
r
r
o
r
!2.5 !2 !1.5 !1 !0.5
!0.03
!0.02
!0.01
0
0.01
0.02
0.03
!=4
log s
e
r
r
o
r
Figure 6. The tting error LHS-RHS (LHS, RHS are normalized to the range [0, 1]) when estimating the parameters of the
Feller process by use of (34) (solid line) or the parameters of the OU process by use of (25) (dashed line) from a simulated
sample of 100 rst-passage times of the level 1 by the Feller process with the -value written on top of each diagram and
all for = 1 and y
0
= 0.5. The errors are close to be the same for the two models.
15
0 0.5 1 1.5 2
0
0.2
0.4
0.6
0.8
1
s
d
i
s
t
r
i
b
u
t
i
o
n
f
u
n
c
t
i
o
n
uniformly distributed first!passage data
0 0.5 1 1.5 2
0
0.2
0.4
0.6
0.8
1
s
d
i
s
t
r
i
b
u
t
i
o
n
f
u
n
c
t
i
o
n
truncated normal first!passage times
0 0.5 1 1.5 2 2.5 3
0
0.2
0.4
0.6
0.8
1
s
d
i
s
t
r
i
b
u
t
i
o
n
f
u
n
c
t
i
o
n
Rayleigh distributed first!passage data
Figure 7. Indication of mist of the OUmodel when the rst-passage data are uniformly distributed (top),truncated normally
distributed (density (s), s > 0) (middle), and Rayleigh distributed (density s(s/), s > 0,
2
= 0.5.1, 2 from left
to right). The test uses the integral equation (25).
16
0.4 0.6 0.8 1 1.2 1.4
0.7
0.8
0.9
1
1.1
1.2
1.3
!
"
0.5 1 1.5
0.7
0.8
0.9
1
1.1
1.2
1.3
!
"
1.6 1.8 2 2.2 2.4
0.7
0.8
0.9
1
1.1
1.2
1.3
!
"
Figure 8. This gure continues in next gure, see caption in Fig. 9.
17
2.6 2.8 3 3.2 3.4 3.6
0.7
0.8
0.9
1
1.1
1.2
1.3
!
"
3.4 3.6 3.8 4 4.2 4.4 4.6
0.7
0.8
0.9
1
1.1
1.2
1.3
!
"
10 10.5 11 11.5
0.7
0.8
0.9
1
1.1
1.2
1.3
!
"
Figure 9. Scatterplots of the 100 pairs of estimates of (, ), each estimated from a sample of 100 simulated rst-passage
times for the OU process. From the top the simulated data correspond to the true values = 0.8, 1, 2, 3, 4, 11 and = 1,
respectively. The shown axis parallel lines cross at the point of true parameter values. The two other lines are the estimated
linear regressions of on and on , illustrating the degree of correlation between the two estimators. The estimated
correlation coefcients are -0.643, -0.618, -0.452, -0.369, -0.039, 0.032, respectively.
18