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A Two-Stage Algorithm For Origin-Destination

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A Two-Stage Algorithm For Origin-Destination

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© © All Rights Reserved
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RESEARCH ARTICLE

A Two-Stage Algorithm for Origin-Destination


Matrices Estimation Considering Dynamic
Dispersion Parameter for Route Choice
Yong Wang1,2, Xiaolei Ma3,4*, Yong Liu1, Ke Gong1, Kristian C. Henricakson2,
Maozeng Xu1, Yinhai Wang2*
1 School of Management, Chongqing Jiaotong University, Chongqing, China, 2 Department of Civil and
Environmental Engineering, University of Washington, Seattle, Washington, United States of America,
3 School of Transportation Science and Engineering, Beijing Key Laboratory for Cooperative Vehicle
Infrastructure, Systems, and Safety Control, Beihang University, Beijing, China, 4 Jiangsu Province
Collaborative Innovation Center of Modern Urban Traffic Technologies, SiPaiLou #2, Nanjing, Jiangsu, China

* [email protected] (XM); [email protected] (YH)

Abstract
OPEN ACCESS

Citation: Wang Y, Ma X, Liu Y, Gong K, Henricakson


This paper proposes a two-stage algorithm to simultaneously estimate origin-destination
KC, Xu M, et al. (2016) A Two-Stage Algorithm for (OD) matrix, link choice proportion, and dispersion parameter using partial traffic counts in a
Origin-Destination Matrices Estimation Considering congested network. A non-linear optimization model is developed which incorporates a
Dynamic Dispersion Parameter for Route Choice.
dynamic dispersion parameter, followed by a two-stage algorithm in which Generalized
PLoS ONE 11(1): e0146850. doi:10.1371/journal.
pone.0146850 Least Squares (GLS) estimation and a Stochastic User Equilibrium (SUE) assignment
model are iteratively applied until the convergence is reached. To evaluate the performance
Editor: Zhong-Ke Gao, Tianjin University, CHINA
of the algorithm, the proposed approach is implemented in a hypothetical network using
Received: August 11, 2015
input data with high error, and tested under a range of variation coefficients. The root mean
Accepted: December 21, 2015 squared error (RMSE) of the estimated OD demand and link flows are used to evaluate the
Published: January 13, 2016 model estimation results. The results indicate that the estimated dispersion parameter theta
Copyright: © 2016 Wang et al. This is an open
is insensitive to the choice of variation coefficients. The proposed approach is shown to out-
access article distributed under the terms of the perform two established OD estimation methods and produce parameter estimates that
Creative Commons Attribution License, which permits are close to the ground truth. In addition, the proposed approach is applied to an empirical
unrestricted use, distribution, and reproduction in any
network in Seattle, WA to validate the robustness and practicality of this methodology. In
medium, provided the original author and source are
credited. summary, this study proposes and evaluates an innovative computational approach to
accurately estimate OD matrices using link-level traffic flow data, and provides useful insight
Data Availability Statement: All relevant data are
within the paper and its Supporting Information files. for optimal parameter selection in modeling travelers’ route choice behavior.

Funding: This research is supported by National


Natural Science Foundation of China (Project No.
71402011, 71471024, 51408019, 71301180,
51329801), National Social Science Foundation of
Chongqing of China (No. 2013YBJJ035), and the
Scientific and Technological Research Program of Introduction
Chongqing Municipal Education Commission (No.
Urban sprawl and population growth have resulted in increasingly severe traffic congestion in
KJ1400307), and the Natural Science Foundation of
Chongqing of China (No. cstc2015jcyjA30012),
major cities around the world. City planners and decision makers have recognized the need for
National Key Technologies R&D Program of China comprehensive traffic management strategies to meet the challenges of rapidly evolving built
(2014BAG01B03), Science and Technology Project environments and population demographics. Effective transportation polices and control

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 1 / 24


Origin-Destination Matrices Estimation

on Transportation Construction by the Ministry of measures can improve traffic safety and quality of service, as well as promoting economic
Transport of China (2015318835200). development and reducing air pollution. Obtaining origin-destination (OD) traffic demand
Competing Interests: The authors have declared matrix in low-cost and high-accuracy manner not only becomes a problem transportation sci-
that no competing interests exist. ence, but also draws attentions from many scholars in various scientific fields. For example,
researchers in statistical physics and complex systems recently proposed a number of novel
methods to estimate OD matrix directly from population data [1, 2, 3, 4, 5, 6]. Reliable OD
matrix estimation can provide critical insight for traffic management, operations, and urban
planning efforts to mitigate congestion [7, 8]. Thus, a reliable OD matrix estimation method is
indispensable for both transportation planners and traffic engineers.
A number of approaches have been developed for estimating OD matrices in the past several
decades [9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20]. Compared with conventional survey-based
method, data-driven OD estimation methods relying on link-level traffic flow measurements
require less effort and offer significantly reduced time and cost for data acquisition and pro-
cessing. For such methods, observed traffic flows at key points throughout the network should
be known as prior information for OD matrix initialization.
Past research on this topic has considered range of different optimization methods, includ-
ing entropy maximizing estimators [21, 22], maximum likelihood estimation [23], Bayesian
inference estimation [24], generalized least squares (GLS) [9, 10, 25] to estimate OD demands.
Entropy maximizing estimators are used to maximize the spread of trip distributions on all
available paths (routes) where the observed traffic flows are used as the only information (i.e.
without a target trip matrix). Maximum likelihood estimation aims to maximize the likelihood
of the closeness between target OD matrix and estimated OD matrix. In the Bayesian inference
approach, the target OD matrix is a prior probability function of the estimated OD matrix on a
basis of observed traffic count data. The GLS estimator is a robust and efficient linear unbiased
estimator, which can solve the estimation of OD matrix by minimizing the Weighted Euclidean
Distances (WED) between the target data and the solution data.
User equilibrium (UE) assignment models are commonly used to obtain path choice behav-
ior based on the estimated OD demand. Deterministic UE assignment models assume that all
users have access perfect information about the generalized link travel costs, and select a route
with the lowest perceived travel cost [26]. Beckman [27] formulated the UE assignment model
by assuming that the OD demands are a function of level of service. A combined distribution
and assignment model which relies on link-level traffic flow data was presented by Fisk and
Boyce [28], and extended by Lam and Huang [29] to address multiclass-user transportation
networks. Fisk [30, 31] proposed a combined entropy maximizing model with UE constraints.
Yang et al. [11] integrated the GLS technique with a UE traffic assignment model for OD
matrix estimation, presented in the form of a convex bi-level optimization problem. Summaries
of the more recent contributions to UE-based traffic assignment are provided in Han [32], Lu
et al. [33], Inoue and Maruyama [34], Kumar and Peeta [35].
The stochastic user equilibrium (SUE) principle allows the perceived cost to vary between
individuals in a heterogeneous population, which can be seen as a more realistic approach than
deterministic UE [15, 36], in which the perceived travel costs cannot vary between travelers. The
probit SUE was first formulated as a generalization of user equilibrium by Daganzo and Sheffi
[37], and developed by Sheffi and Powell [38] as a mathematical programming problem. Liu and
Fricker [39] presented a two-stage SUE approach to estimate OD matrices and the probit disper-
sion parameter in an iterative manner. Yang et al. [15] improved on the methods described in
Liu and Fricker by incorporating link traffic flows and travel cost obtained using logit-based
SUE traffic assignment. Meng et al. [40] presented a linearly constrained model and solution
algorithm for the probit SUE problem with fixed demand and separable link travel time func-
tions. This modeling approach was extended in Meng et al. [41] using elastic demand and non-

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 2 / 24


Origin-Destination Matrices Estimation

separable link travel time functions. Time-dependent traffic assignment can be also formulated
as a multinomial logit model [42, 43, 44], and this has become one of the most common methods
for SUE-based traffic assignment [45, 46, 47]. In a fixed-point formulation, fixed target demands
or link flows are used to establish model based on UE and SUE principles [13, 14, 19].
In the multinomial logit model formulation, the link choice probability is a function of a dis-
persion parameter θ [16], which describes road users’ perception of travel costs. Though the
dispersion parameter θ is predetermined in many previous studies [14, 36, 45, 46, 47, 48], here
we assume that this value should be allowed to change with traffic conditions. In addition, Lo
and Chan [16] proposed a maximum likelihood procedure for simultaneously estimating the
OD matrix and the dispersion parameter θ, while the link choice proportions and link flows
can be further calculated based on the maximum likelihood estimators of OD matrix and θ.
Compared with the previous studies, the main contributions of this paper lie in: (1) A fixed-
point model is formulated with a dynamic dispersion parameter θ, where the estimation of link
choice proportions is integrated into the optimization procedure; (2) A GLS estimator is uti-
lized to train this model, and the link choice proportions can be simultaneously calculated
based on the OD matrix and dispersion parameter through a multinomial logit model; (3) A
two-stage iterative algorithm is presented to refine the OD matrix and dispersion parameter
estimates, and Sequential Quadratic Programming (SQP) from the extended quasi-Newton
method is applied in the two-stage algorithm process[49].
The remainder of this paper is organized as follows: In Section2, relevant notation, defini-
tions, and model formulations are presented, followed by a link choice proportion approach
to calculate the observed link flow using a true OD matrix. A two-stage algorithm is described
in Section 3, along with model implementation details. The performance of the proposed
approach is tested in a hypothetical network, and a sensitivity analysis is conducted using a
range of variation coefficients. Results are presented and compared with those obtained
through other established OD estimation methods. In section 4, results are presented for a real-
world network using loop detector data in the city of Seattle, WA to demonstrate the practical-
ity of the proposed approach. Finally, conclusions are summarized in Section 5.

Model Formulation
Related Notations and Definitions
The notation and parameter definitions used throughout the paper are as follows:
K the set of network links k 2 K, where T denotes the total number of links
L the set of observed links l 2 L, where Γ denotes the number of observed links
J the set of OD pairs j 2 J, where τ indicates the total number of OD pairs
A the set of paths connecting the OD pair j, a 2 A
ck travel cost of link k
crj travel cost of path r connecting the OD pair j
tk the free flow travel time of link k
Ck the capacity of link k
αk the performance function parameter of link k
βk the exponential value of link k’s performance function
M the set of nodes in the network

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 3 / 24


Origin-Destination Matrices Estimation

f the vector for estimated link flows, where fk is the estimated flow for link k

f^ the vector for observed link flow, where f^l is the observed link flow for link l
d the estimated OD vector matrix, where dj is the jth element of d for OD pair j
d the target OD vector matrix, where d j is the jth element of d for OD pair j

d~ the initial OD vector matrix for SQP algorithm optimization


W' the initial weight matrix in the group of all paths connecting each OD pair
w'mn the initial weight element of all paths connecting nodes m and n, m,n 2 M
W the weight matrix in the group of all paths connecting each OD pair
wmn the weight element of all paths connecting nodes m and n, m,n 2 M
E the identity matrix with the same dimension as the initial weight matrix
G the vector matrix of observed link flows, where Gi is the ith element of G
U the covariance matrix for the target OD vector and estimated OD vector
V the covariance matrix for the observed link flows and estimated link flows
P the matrix of link choice proportions, where pkj is the kjth element of P. This is equivalent to
the proportion of OD pair j traveling on the observed link k
Prj the probability of path r that connects OD pair j being chosen for a trip
xk the observed traffic flow of link k
ðsÞ
xk the estimated traffic flow of link k at the sth iteration in the traffic assignment stage
ðsÞ
yk the auxiliary mean traffic flow of link k at the sth iteration in the traffic assignment stage
θ the estimated dispersion parameter for OD estimation

y the target dispersion parameter for OD estimation

y~ the initial dispersion parameter for SQP algorithm optimization

Q the covariance matrix for θ and y


Sd the feasible solution set for OD matrix
Sθ the feasible solution set for θ parameter
s2dj the variance for OD demands

s2xk the variance for link flows

s2y the variance for dispersion parameter


λd random term for the target OD matrix
λf the random term for observed link flows
F1 the “distance” between the estimated OD vector matrix d and target demand OD matrix d

F2 the “distance” between the estimated link flow vector f and observed link flow vector f^

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 4 / 24


Origin-Destination Matrices Estimation

F3 the “distance” between the estimated dispersion parameter θ and the target dispersion
parameter y
akrj decision variable, if the link k lies on path r connecting the OD pair j, and set akrj = 1, and 0
otherwise
η the percentage of traffic flow traveling from each node to the most adjacent node
RMSE(OD) the root mean squared error between estimated and true OD matrices
RMSE(LF) the root mean squared error based on estimated and observed (true) link flows
Based on the above notations, the OD estimation model development and validation proce-
dure can be described as follows:
1. A ground truth OD matrix is used as prior information to calculate the link flows based on
the link choice proportion model. These link flows represent the measured traffic flows
obtained through fixed mechanical sensors or other means;
2. The observed link flows are chosen from those calculated link flows at fixed points through-
out the network;
3. The estimated OD matrix, link flows, and dispersion parameter are obtained via the fixed-point
model and two-stage iterative algorithm using the partial observed link flows from step (2);
4. Results are evaluated and compared with the ground truth as established in step (1).

The Fixed Point Model with Dynamic Dispersion Parameter. As presented in the previ-
ous subsection, the estimated OD vector matrix is expressed as d = [d1,d2,. . .,dj,. . .,dτ]', where
dj denotes the mean traffic flow of the jth element of d for OD pair j. Consider an OD pair j
connected by a link k which is associated with a link performance cost function ck(fk) equal to
the cost of using link k. The link performance cost function [50] is expressed during the traffic
assignment procedure in Eq 1:

f b
ck ðfk Þ ¼ tk ½1:0 þ ak ð k Þ k ; 8k 2 K ð1Þ
Ck

The link flow vector is defined as f = [f1,f2,. . .,fΓ]', and the matrix of link choice proportions is
denoted as P = [pkj], where 0  pkj  1. This represents the proportion of OD pair j connected
by the link k. Thus, the mathematical expectation of link flow vector f can be calculated as E[f] =
[Pd]Γ×1, where Pd is the product of the observed mean link flow vector and the matrix of link
choice proportions P. P can be adjusted by the link flows and the dispersion parameter θ.
The OD matrix can be estimated via a fixed point formulation by considering the target OD
matrix and observed link flows as follows [9, 10, 13, 14, 15, 16, 18, 19, 51]:

 þ F ðf ; ^f Þ
d ¼ arg min½F1 ðd; dÞ 2
d2Sd
ð2Þ
 T U 1 ðd  dÞ
¼ arg min½ðd  dÞ  þ ðPðdÞd  ^f ÞT V 1 ðPðdÞd  ^f Þ
d2Sd

Where:
P(d) = {pkj(dj)} is the assignment matrix, which represents the proportion of OD pair j using
the observed link k;
f = P(d)d is the estimated link flow vector. f = {fk}, where fk = Sj pkj(dj)dj.

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Origin-Destination Matrices Estimation

In this study, the dispersion parameter is integrated into the objective function (Eq 2) [16,
19] as follows:
 þ F ðf ; ^f Þ þ F ðy; yÞ
ðd; yÞ ¼ arg min½F1 ðd; dÞ  ð3Þ
2 3
d 2 Sd
y 2 St

This model can be seen as a Stochastic User Equilibrium (SUE) problem [13]. The General-
ized Least Square (GLS) estimator can be used to solve Eq 3 by minimizing the Weighted
Euclidean Distances (WED) between the target data and the solution vector, and Eq 3 can be
then reorganized as shown in Eq 4 [9, 10, 15, 48]:

ðd; yÞ
GLS  U 1 ðd  dÞ
¼ arg min½ðd  dÞ
T  þ ðPðd; yÞd  ^f Þ V 1 ðPðd; yÞd  ^f Þ þ ðy  yÞ
T  Q1  ð4Þ 2

d 2 Sd
y 2 St

Where:
P(d,θ) = {pkj(dj,θ)} is the assignment matrix, and is a function of both OD matrix and disper-
sion parameter θ.
f = P(d,θ)d is the estimated link flow vector. f = {fk}, where fk = Sj pkj(dj,θ)dj.
The matrix for link choice proportions P can be generally assumed fixed during the optimi-
zation procedure [10, 13, 14, 19]. This procedure performs well for uncongested traffic condi-
tions or an idealized traffic network with fixed link costs. However, when the network becomes
congested, users’ choices are increasingly influenced by adverse traffic condition. In this case,
link flow and cost are not independent, and the assignment matrix P should be assumed to
vary within each optimization step for link flow and OD estimation. Similarly, the GLS estima-
tors of d and θ can be also obtained by solving Eq 4.
The Link Choice Proportion Calculation Using the Dispersion Parameter. As men-
tioned in notation and definitions subsection, the link flow and cost will be updated when a
new set of values of d and θ is received. Drivers’ link choice decisions are influenced by the net-
work-wide traffic condition, and thus the link choice proportion matrix P should be allowed to
vary as well. The method of successive average (MSA) is adopted to calculate equilibrium link
flows in the traffic assignment procedure [7, 16, 45, 52].
The cost of path r connecting the OD pair j can be expressed as:
X
crj ¼ akrj ck ; 8k 2 K ð5Þ
k

The probability Prj can be then computed according to the path choice logit model [45]:
expðcrj yÞ
Prj ¼ X
expðcaj yÞ
a

1 X
¼ X expðy akrj ck Þ ð6Þ
expðcaj yÞ k
a

1
¼ X ½expðya1rj c1 Þ  expðya2rj c2 Þ  . . .  expðyaTrj cT Þ
expðcaj yÞ
a

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Origin-Destination Matrices Estimation

For a driver traveling along the path r, the weight assigned to link k is equal to exp(-ckθ). It
is worth noting that the sum of probabilities over all feasible paths for each OD pair is equal to
one.
As previously noted, W' = [w'mn] is the initial weight matrix of all possible paths connecting
each OD pair. With the initial weight is set to w'mn = exp(-ckθ), then W', W20 , and W30 represent
the weight matrix in the group of paths with one link, two links and three links respectively.
Therefore, the weight matrix for all possible paths can be formulated as:
1
W 0 þ W20 þ W30 þ    ¼ ðE  W 0 Þ E ð7Þ

Wong [53] and Lo and Chan [16] have proven that the right side of Eq 7 is convergent for
any acyclic networks, and is equal to W = (E−W')−1 − E. Therefore, the probability of a trip
from node m to node n (OD pair j) choosing link k can be calculated as follows:
wmg expðyck Þwnn
pkj ¼ ð8Þ
wmn

Where link k connects node g and node v, and wmn expresses weight matrix of all possible
paths connecting nodes m and n, m,n 2 M. wmn is set to 1 for all nodes in the network.
Following the previous definition, the auxiliary mean traffic flow ykðsÞ of link k is defined for
each incoming d and θ via the following equation:

ykðsÞ ¼ ½Pdk
X ; 8k 2 K ð9Þ
¼ p d
j kj j

The equilibrium traffic link flows can be then obtained using the MSA method. Specifically,
the flow of link k can be calculated at the (s+1)th iteration with the following equation:
1
xkðsþ1Þ ¼ xkðsÞ þ ðykðsÞ  xkðsÞ Þ
s
1X s ; 8k 2 K ð10Þ
¼ ykðjÞ
s j¼1

As shown in Eq 10, the flow of link k at the (s+1)th iteration is equal to the mean of the aux-
iliary traffic flow of link k in the previous s iterations.
When a new set of values of d and θ is received, the matrix P of link choice proportions is
updated following the procedure described above, and is then integrated into the Eq 4 to
update the values of d and θ. This optimization procedure continues until convergence of the
OD matrix and dispersion parameter estimation is reached.

Model Solution Algorithm


To solve the Stochastic User Equilibrium (SUE) problem described above, a two-stage algo-
rithm for GLS estimation and SUE traffic assignment is proposed: First, the OD matrix d and
the dispersion parameter θ are simultaneously estimated under the condition of the fixed link
flows, link costs, and weight matrix. Second, the link flows, link costs, and link choice propor-
tions are updated according to the new values of d and θ in the SUE assignment process. The
two-stage algorithm is executed iteratively until the convergence of values of d and θ is reached.
Sequential quadratic programming (SQP) from the extended quasi-Newton method is chosen
as the solution method [49].

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 7 / 24


Origin-Destination Matrices Estimation

Two-Stage Algorithm
The initialization procedure of the two-stage algorithm can be described as follows:
 the initial dispersion
1. Initialize the counter t = 0, set the initial OD vector matrix d ð0Þ ¼ d,
ð0Þ
parameter y ¼ y,  and the initial link flow x ¼ 0, k 2 K.
ð0Þ
k

2. Calculate the initial link costs for all links in the network using Eq 1, and calculate the
weight matrix W for all paths based on the initial link costs and θ(0).
3. Calculate the link choice proportion matrix P using the weight matrix W and θ(0).
4. Calculate the initial mean auxiliary traffic flow for all the observed links with Eq 9, and
update t = t + 1.
The first stage of the algorithm is described as follows:
Step 1. The objective function (Eq 4) can be updated with the new mean auxiliary observed
link flows as follows:

ðdðtÞ ; yðtÞ Þ
GLS  U 1 ðd ðtÞ  dÞ
¼ arg min½ðdðtÞ  dÞ
T  þ ðPðtÞ d ðtÞ  ^f Þ V 1 ðPðtÞ d ðtÞ  ^f Þ þ ðyðtÞ  yÞ
T  Q1  ð11Þ
2

d0
y>0

Where:
U−1, P(t), V−1, and Q−1 can be updated using the new mean auxiliary observed link flows,
estimated OD vector matrix, dispersion parameter, and link flow vector respectively;
The feasible set for d and θ should meet the requirements d  0, θ > 0. When the value of θ
approaches zero, the path choice probabilities for all paths tend to be equal. As the value of
θ increases, the path choice probabilities tend to be deterministic.
Step 2. Use the SQP algorithm to obtain a new set of values of d(t) and θ(t) that minimizes the
objective function. The starting point for optimizing the OD vector d~ ðtÞ and dispersion param-
eter y~ðtÞ should be fixed in advance. During the iterative process of the SQP algorithm, when-
ever a new value θ is received, the link choice proportion matrix P will be updated by changing
the value of exp(−θck) in Eq 8, while the link cost and weight matrix should remain unchanged.
The second stage of the algorithm can be described as follows:
Step 3. Initialize the counter s = 1.
Step 4. Calculate the weight matrix W with the new dispersion parameter θ(t).
Step 5. Calculate the link choice proportion matrix P(t) using the weight matrix W and disper-
sion parameter θ(t).
Step 6. Calculate the mean auxiliary traffic flow for all observed links as follows:
X
ylðsÞ ¼ ½PðtÞ d ðtÞ l ¼ p d; l 2 L
j lj j

Step 7. Calculate the equilibrium traffic link flow of link k via the MSA method:
1
xlðsþ1Þ ¼ xlðsÞ þ ðylðsÞ  xlðsÞ Þ; l 2 L
s

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 8 / 24


Origin-Destination Matrices Estimation

Step 8. The maximum relative difference between current and previous mean link flows should
satisfy the following requirement:
( )
jxlðsþ1Þ  xlðsÞ j
max  ε1 ð12Þ
8l2L xlðsþ1Þ

If the above requirement is met, the algorithm proceeds directly to step 11, otherwise pro-
ceed to step 9.
ðsþ1Þ
Step 9. Calculate the new link costs according to xl , l 2 L.
Step 10. Calculate the weight matrix using the updated link costs, set s = s + 1, and return to
step 5.
Step 11. The maximum relative difference between the current and previous OD matrix esti-
mates should satisfy the following requirement:
( ðtÞ )
jdj  djðt1Þ j
max  ε2 ð13Þ
8j2J djðtÞ

If the above requirement is met, terminate the procedure and output the current estimates
of OD vector matrix d and dispersion parameter θ as d(t) and θ(t). Otherwise, set t = t + 1,
and proceed to step 12.

Step 12. Calculate the new starting points as follows: d~ ðtþ1Þ ¼ 1t d ðtÞ þ t1
t d
ðt1Þ
,
~
y ðtþ1Þ 1 ~ ðtÞ
¼ y þ y t1 ~ ðt1Þ
, and return to step 1.
t t

Model Evaluation
To evaluate the performance of the proposed method, the root mean squared errors (RMSE)
for OD matrix and link flows after convergence are defined as follows:
ðsþ1Þ
(1) The root mean squared error (RMSE) of the estimated link flows xl relative to the true
link flow xl is computed as follows:
vffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
u X
u1 G
RMSEðLFÞ ¼ t
2
ðxlðsþ1Þ  xl Þ ð14Þ
G
l¼1

Similarly, the RMSE of the observed (target) link flows ^f l relative to the true link flows xl

can be defined as RMSE ðLFÞ, where xðsþ1Þ is replaced by ^f in Eq 14.


l l
(2) The RMSE of the estimated OD matrix d(t) relative to the true OD matrix d can be defined
as RMSE (OD):
vffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi
u X t
u1
RMSEðODÞ ¼ t ðdjðtÞ  dj Þ
2
ð15Þ
t
j¼1

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Origin-Destination Matrices Estimation

Likewise, the RMSE of target OD matrix d relative to the true OD matrix d is defined as
RMSE (OD), where d(t) is replaced by d in Eq 15.
j

Numerical Experiment and Result Analysis


A Hypothetical Network Test
In this section, the performance of the proposed approach is tested in a hypothetical network.
The network and data proposed by Yang et al. [15] and Caggiani et al. [19] are adopted as the
test bed with some slight modifications. The network (presented in Fig 1), is composed of 9
nodes (3 origin centroids and 3 destination centroids), and 14 links. The true and initial OD
vector matrices d and d~ for the SQP algorithm are shown in Table 1. The initial dispersion
parameter y~ is assumed to be 4, and the true dispersion parameter y^ is fixed to 1.5. Note that
the initial OD matrix d~ and dispersion parameter y~ are quite dissimilar from those of the
ground truth data.
The following parameters in the Bureau of Public Roads (BPR) [50] link performance func-
tion are used: αk = 0.15 and βk = 4, 8k 2 K. In addition, the free flow travel time (tk) and capac-
ity (Ck) for each link are predetermined as shown in Table 2.
The ground truth link flows can be generated by allocating the true OD matrix to the traffic
network using SUE-Logit assignment method presented in Section 2.3. The true dispersion
parameter is θ = 1.5, resulting in the link flows shown in Table 3. The set of links {5, 6, 7, 11, 13}
is selected as the observed links.
In this example, we assume that the OD vector and link flow vector follow the Poisson dis-
tribution. The covariance matrices U (for OD demands) and V (for link flows) in Eq 4 can be
assumed to be diagonal matrices [9, 14, 54]. The diagonal element for U, V and Q can be com-
puted respectively through the following equations:
2 2 2
s2dj ¼ ðcvd  dj Þ ; s2xk ¼ ðcvx  xkðsþ1Þ Þ ; s2y ¼ ðcvy  yÞ

Where cvd, cvx and cvθ represent the variation coefficients for OD demands, link flows, and dis-
persion parameter respectively. Specifically, these parameters are set as cvd = 0.3, cvx = 0.05,
and cvθ = 0.1.
The target OD matrix d, observed link flow vectors ^f , and target dispersion parameter y can
be generated separately by adding random terms into the corresponding true values. The ran-
dom terms are sampled from independent normal variables with zero means. For instance, the
target OD matrix can be calculated by adding a random term with λd = 0.3 to the values of the
true OD matrix divided by two, the observed link flow vectors can be generated by adding a
random term with λf = 0.1, and the target parameter can be set as y ¼ 4. In addition, the error
tolerance threshold used in the optimization is set to ε1 = ε2 = 10−3. The convergence for theta
is plotted in Fig 2, which shows the estimate slowly falling in the first 120 iterations before rap-
idly converging to the true value at 1.5099. This is a very slight deviation with the true value of
1.5. In addition, the convergence of the objective function is presented in Fig 3, where the value
of the objective function sharply falls at the first iteration and then gradually decreases and lev-
els off at a lower value. Poor initial choices of OD input vector and dispersion parameter may
lead to the slower convergence.
In order to further evaluate the effectiveness of the proposed approach, a sensitivity analysis
is conducted with parameter cvθ(CVT) varying from 0.1 to 0.5 and cvd(CVD) changing from

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 10 / 24


Origin-Destination Matrices Estimation

Fig 1. The test network used in the numerical example.


doi:10.1371/journal.pone.0146850.g001

Table 1. The true and initial OD vector matrices.

OD pair 1–6 1–8 1–9 2–6 2–8 2–9 4–6 4–8 4–9
j 1 2 3 4 5 6 7 8 9
d 120 150 100 130 200 90 80 180 110
d~ 30 20 10 30 30 30 30 40 20

doi:10.1371/journal.pone.0146850.t001

Table 2. Free flow travel time and capacity for each link.

link 1 2 3 4 5 6 7 8 9 10 11 12 13 14
tk 2.0 1.5 3.0 1.0 1.0 2.0 2.0 1.0 1.0 1.0 2.0 1.0 1.0 1.0
Ck 280 290 280 280 600 300 500 400 500 700 250 300 350 520
doi:10.1371/journal.pone.0146850.t002

Table 3. True link flows in the hypothetical network.

link 1 2 3 4 5 6 7 8 9 10 11 12 13 14
xk 125 143 103 172 474 172 201 313 307 393 279 148 313 475
doi:10.1371/journal.pone.0146850.t003

0.1 to 1. This generates 50 different estimates for RMSE (OD), RMSE (LF) and Theta as pre-
sented in Figs 4–6.
As shown in Fig 4, RMSE (OD) increases with the variation coefficient cvθ when cvd falls
between 0.2 and 0.8. With cvθ fixed between 0.3 and 0.5, RMSE(OD) can be seen as a convex

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 11 / 24


Origin-Destination Matrices Estimation

Fig 2. Convergence of the theta estimate.


doi:10.1371/journal.pone.0146850.g002

function of cvd. Alternatively, when cvθ is between 0.1 and 0.2, cvd has a negligible impact on
RMSE (OD). Thus, the maximum value (19.9022) of RMSE (OD) can be found at cvd = 0.5 and
cvθ = 0.5, and the minimum value (4.6859) is obtained at cvd = 0.3 and cvθ = 0.1. Compared
with the initial RMSE (OD) of 93.8971 calculated from Table 1, a 78.8% reduction is achieved
at the maximum RMSE (OD), and a 95% reduction is obtained at the minimum RMSE (OD).
Fig 5 shows the impact of cvd and cvθ on RMSE (LF). With the value of cvd fixed, RMSE (LF)
increases with the variation coefficient cvθ. For a fixed value of cvθ, the RMSE (LF) decreases with
an increase in cvd. Thus, we can conclude that maximum RMSE (LF) value of 19.6597 is located
at cvd = 0.1 and cvθ = 0.5, and the minimum value of 8.6696 can be found at cvd = 0.6 and cvθ =
0.1. Compared with the initial RMSE (LF) value of 30.8347, 36.2% and 71.9% reductions can be
achieved for the maximum value of RMSE (LF) and minimum value of RMSE (LF) respectively.
As shown in Fig 6, the value of theta varies negligibly with the choice of cvd and cvθ. In other
words, the estimated value of theta always converges to approximately the true value. As
shown in Fig 6, for a fixed value of cvd, the estimated θ is close to the true value for any given
cvθ. For example, the value of θ fluctuates between 1.37 and 1.51 when cvd = 0.3. Likewise, for
any fixed cvθ, the estimated θ varies minimally about the true value of θ using the proposed
method. For example, the estimated θ is between 1.35 and 1.52 for cvθ = 0.1.
 y,
The above discussion reveals a fact that the initial value of d,  and observed link flow vec-
tors ^f do not affect the theta estimation performance. This is equivalent to a convex optimiza-
tion problem, where the optimal results tend to converge near the true dispersion parameter
value. This implies that the estimate of θ is insensitive to the variation coefficients, and can be
used as a stable and accurate parameter to determine travelers’ route decisions.

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 12 / 24


Origin-Destination Matrices Estimation

Fig 3. Convergence of the objective function.


doi:10.1371/journal.pone.0146850.g003

Comparison and Analysis


To further demonstrate the advantages of the proposed methodology, two OD matrix estima-
tion methods are implemented and compared with the proposed approach. To make this com-
parison, we first implement the algorithm described in Yang et al. [15], which presents an
optimization model for OD matrix estimation in congested networks using the logit-based

Fig 4. RMSE(OD) versus cvd and cvθ.


doi:10.1371/journal.pone.0146850.g004

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Origin-Destination Matrices Estimation

Fig 5. RMSE(LF) versus cvd and cvθ.


doi:10.1371/journal.pone.0146850.g005

SUE. The method described in Lo and Chan [16] is implemented for the second comparison.
This method applies both statistical estimation and traffic assignment to simultaneously calcu-
late the OD matrix and link choice proportions based on OD survey data and traffic counts. To
maintain a fair comparison, the same test network and data set are applied in all cases.
The OD matrix estimation method proposed by Yang et al. [15] is given in section 2.2. The
objective function is shown in Eq 16.

ðd; yÞ ¼  þ F ðf ; ^f Þ
arg min½F1 ðd; dÞ 2
d2Sd

1  2 þ 1 ðPd  ^f Þ2 
¼ arg min½ ðd  dÞ ð16Þ
d0 2 2

y>0

Fig 6. Estimated Theta versus cvd and cvθ.


doi:10.1371/journal.pone.0146850.g006

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 14 / 24


Origin-Destination Matrices Estimation

Table 4. Comparison between Yang et al.’s approach and the proposed approach with OD matrix and link flow estimation.

Approach Estimated OD matrix Estimated Link flows

RMSE ðODÞ RMSE (OD) RMSE ðLF Þ RMSE (LF)

Yang et al.’s approach 24.27 20.85 26.65 19.02


Proposed approach 24.27 18.79 26.65 17.39
doi:10.1371/journal.pone.0146850.t004

In Yang et al.’s work, the weighted Euclidean distance function is used to develop a unit
weighting matrix and the value of theta is set to 1.5. The RMSE(OD), RMSE(LF), RMSE ðODÞ 
and RMSE(ðLF Þ for Yang et al.’s approach are calculated and compared with the proposed
approach in Table 4:
As shown in Table 4, the proposed method yields significantly lower RMSE (OD) and
RMSE (LF) relative to Yang et al.’s approach. Compared with the initial RMSE values, a 22.6%
reduction in RMSE (OD) is achieved using the proposed approach, while only a 14.1% reduc-
tion is achieved using the method described in Yang et al. Similarly, the proposed approach
resulted in a 34.7% reduction in RMSE(LF), while only a 28.6% reduction was achieved using
Yang et al.’s approach. One reason that the dispersion parameter is estimated and integrated
 in the proposed method, and it is able to yield a better estimate of the
into the Eq 3 by F3 ðy; yÞ
dispersion parameter than previous approaches. The other reason is that the covariance matri-
ces U (for OD demands), V (for link flows) and Q (for dispersion parameter) are not a fixed
variable during the calculation. These improvements can help the method enhance the estima-
tion performance for the OD matrix and link flow vectors.
Lo and Chan [16] present the following maximum likelihood objective function:

ðd; yÞ ¼ arg maxd0;y>0 ln Lðy; d j ^f ; dÞ


 ð17Þ

In Lo and Chan [16], it is assumed that the observed flows are equal to the true flows in the
test network. For Lo and Chan’s algorithm, we set the target dispersion parameter to y ¼ 4
(This is also equal to the initial dispersion parameter value used in Lo and Chan [16]’s work),
and the variation coefficients as follows: cvθ = 0.1, cvx = 0.05, and cvd = 0.3. In order to evaluate
the performance of the proposed approach relative to that of Lo and Chan [16]’s method,
RMSE (OD), RMSE (LF), and the estimated Theta are selected for comparison and shown in
Table 5.
Unlike Lo and Chan’s method, random terms are added to the observed link flows in the
proposed approach, thus introducing additional challenges for estimation. However, the results
presented in Table 5 demonstrate that the method proposed in this paper outperforms Lo and
Chan’s approach in terms of OD matrix, link flow, and Theta estimation accuracy.

Table 5. Comparison between Lo and Chan’s approach and the proposed approach with OD matrix, link flow and Theta estimation.

Approach Estimated OD matrix Estimated Theta

RMSE (OD) RMSE (LF) Theta target Theta estimated


Lo and Chan’s approach 5.34 12.08 4 1.572
Proposed approach 4.69 9.77 4 1.509
doi:10.1371/journal.pone.0146850.t005

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Origin-Destination Matrices Estimation

Fig 7. Square network in Seattle. Double circle nodes represent zone centroids (origins and destinations).
doi:10.1371/journal.pone.0146850.g007

Application to A Square Network in Seattle


A square network in Seattle is used as a congested network case study to demonstrate the
applicability and transferability of the proposed approach in a real-world traffic network
(Shown in Fig 7). Empirical data was collected from loop detectors located along one freeway
section in Seattle area, and obtained for this research through the Strategic Highway Research
Program 2 (SHRP 2 program) supported by Washington State Department of Transportation
(WSDOT) [55].
The square test network used in this case study consists of 4 nodes and 8 links, where all
nodes are centroids (origins and destinations). The topology of the test network is outlined in
Fig 7. We assume that the study network is acyclic, such that the traffic flow starting from one
node will leave the network before returning to the original node. Specifically, Links 1 and 2
represent the SR 520 Bridge connecting I-5 in Seattle and SR 202 in Redmond. Interstate 90 (I-
90) is represented by Links 3 and 4, and Interstate 5 (I-5) is represented by Links 5 and 6. Links
7 and 8 represent Interstate 405 (I-405), which intersects I-90 in the south and SR 520 in the
north.
Traffic flows were obtained from loop detectors installed at nodes 1, 2, 3 and 4, illustrated in
Fig 8. The parameters for the BPR link performance cost function (Eq 18) were estimated
based on the empirical data and are presented in Table 6.
f b
ck ðfk Þ ¼ tk ½1:0 þ ak ð k Þ k ; 8k 2 K ð18Þ
Ck

Table 7 indicates the external traffic flow recorded for each node during peak hour, where
1-Link 1 represents the external traffic flow on Link 1 from node 1, and 2-Link 7 represents the
external traffic flow on Link 7 from node 2, and so forth. To convert true link flows into a
ground truth OD matrix, the flow proportion for each node η = 0.6 is assumed based on exten-
sive video records and filed surveys. This implies that, for the traffic leaving each node, 60%
exits the network from an adjacent node while 40% exits from the other nodes. In order to
avoid circular flow in the OD calculation process, it is assumed that the final remaining traffic

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 16 / 24


Origin-Destination Matrices Estimation

Fig 8. Traffic flow fluctuation by time of day.


doi:10.1371/journal.pone.0146850.g008

flow will leave from the last node before returning to the original node. Based on these assump-
tions, the ground truth OD matrix is calculated and shown in Table 8. In addition, the initial
OD matrix d~ can be computed by rounding the last digit of the true OD matrix as shown in
Table 8.
The true OD matrix in Table 8 is then used to assign the corresponding traffic flow into
each link according to Eq 5 through Eq 10. The calculated traffic flows can be assumed to rep-
resent the true link flows, where link 1, 3, 5, 6, and 8 are selected as the observed links to esti-
mate OD matrix shown in Table 9.
Similar to the hypothetical network, we assume that the OD demands and observed link
flows follow the Poisson distribution, and the covariance matrices U and V can be assumed to
be diagonal. The initial value of the dispersion parameter y~ is set to 40.5. The remaining input
parameters are set identically to the hypothetical network. In addition, a sensitivity analysis
with 50 different combinations of variation coefficients cvd and cvθ was conducted to investi-
gate the optimal parameter initialization for the proposed approach. The results of this sensitiv-
ity analysis are shown in Figs 9–11.

Table 6. BPR link performance cost function parameter calibration.

Links Road Name Length(km) tk(h) Ck(pcu/h) αk βk


1,2 SR 520 7.5 0.1162 4149 0.1450 3.5
3,4 I-90 3.6 0.0667 8685 0.1035 2.7
5,6 I-5 3.5 0.1016 9683 0.0988 2.7
7,8 I405 7.8 0.1332 7961 0.1242 3.5
doi:10.1371/journal.pone.0146850.t006

Table 7. The external true traffic flow for each node at peak hour.

Link direction 1-Link 1 2-Link 2 3-Link 3 4-Link 4 4-Link 5 1-Link 6 2-Link 7 3-Link 8
Traffic flow 3199 2480 5499 5535 6018 7169 5628 8153
doi:10.1371/journal.pone.0146850.t007

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Origin-Destination Matrices Estimation

Table 8. True OD matrix and initial OD matrix at peak hour for each OD pair.

OD pair 1–2 1–3 1–4 2–1 2–3 2–4 3–1 3–2 3–4 4–1 4–2 4–3
j 1 2 3 4 5 6 7 8 9 10 11 12
d 3067 2489 4814 2389 3774 1946 3277 5772 4604 4497 2773 4284
d~ 307 249 482 239 378 195 328 578 461 450 278 429

doi:10.1371/journal.pone.0146850.t008

Table 9. Observed link flows at peak hour.

Link No. 1 2 3 4 5 6 7 8
xk 3711 - 8282 - 8439 7857 - 7591
doi:10.1371/journal.pone.0146850.t009

Fig 9. RMSE(OD) versus cvd and cvθ in the actual network.


doi:10.1371/journal.pone.0146850.g009

Fig 9 shows the value of RSME (OD) versus cvd and cvθ. For a fixed value of cvd between 0.1
and 0.7, RMSE(OD) increases with cvθ. RMSE(OD) is a concave function of cvθ when cvd is
fixed between 0.8 and 1.0, and a convex function of cvd for a fixed value of cvθ between 0.1 and
0.5. Thus, the maximum RMSE (OD) of 85.6123 can be obtained at cvd = 0.7 and cvθ = 0.5, and
the minimum value of 23.5917 can be obtained at cvd = 0.1 and cvθ = 0.1.
As shown in Fig 10, the value of RMSE (LF) increases with cvθ for a fixed value cvd. For a
fixed value of cvθ, the value of RMSE (LF) decreases with an increase of cvd. The maximum
RMSE (LF) of 82.5113 is found at cvd = 0.1 and cvθ = 0.5, and the minimum value of 7.3277 at
cvd = 1.0 and cvθ = 0.1.
As noted in the hypothetical case, the choice of cvd and cvθ has very little impact on the esti-
mation of Theta. As shown in Fig 11, the estimated dispersion parameter θ is between 20.8327
(cvd = 0.5 and cvθ = 0.5) and 22.7165 (cvd = 0.7 and cvθ = 0.2) in all cases. The best estimate of
dispersion parameter θ can be found between 20.8327 and 22.7165.
Finally, using the BPR link performance cost function parameters described in Table 6, differ-
ent combinations of variation coefficient cvd = 0.3 and cvθ = 0.1; cvd = 0.5 and cvθ = 0.5;cvd = 0.7
and cvθ = 0.2 are used to estimate theta for the actual network.

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 18 / 24


Origin-Destination Matrices Estimation

Fig 10. RMSE(LF) versus cvd and cvθ in the actual network.
doi:10.1371/journal.pone.0146850.g010

It is interesting to observe that the estimated RMSE(OD), RMSE(LF), and Theta for both
hypothetical and actual networks exhibit a similar trend yet have obvious differences. Two pri-
mary reasons may explain these differences: First, the network topology is quite different for
the two scenarios. The hypothetical network is unidirectional, where each node can be either
origin or destination. In contrast, the actual network is bidirectional, where each node is both
origin and destination, and thus multiple paths may exist between each OD pair. For example,
the traffic flows on both 1-Link 1 and 1-Link 6 contribute to the OD demands from node 1 to
node 2. Second, compared with the hypothetical network with equal cost parameters for all
links, a more realistic BPR link performance cost function is adopted for the actual network. In
the real-world network, the parameters (e.g. free-flow travel time and link capacity) are

Fig 11. Theta estimated versus cvd and cvθ in the actual network.
doi:10.1371/journal.pone.0146850.g011

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 19 / 24


Origin-Destination Matrices Estimation

Fig 12. Estimated Theta values in the actual network by time of day.
doi:10.1371/journal.pone.0146850.g012

calibrated for each link based on empirical data. That said, the sensitivity analysis for Theta
produced similar results for both the hypothetical and actual networks, indicating that this
parameter is not sensitive to the choice of variation coefficients. In addition, the theta estimates
obtained using a range of different parameter settings exhibits a similar and regular trend over
time of day as shown in Fig 12. These findings provide guidance for initial parameter selection,
and offer useful insight for interpreting modeling results.

Conclusions
This paper proposes a two-stage algorithm to simultaneously estimate origin-destination
matrices and link choice proportions by incorporating a dynamic dispersion parameter into
the route choice model. The dispersion parameter θ is of practical significance in describing
travelers’ route choice decisions, but has typically been assumed constant in previous studies.
Finding the optimal dispersion parameter is not a straightforward task. To address this issue,
this paper presents a model calibration procedure to simultaneously estimate the dispersion
parameter θ, link choice proportions, and OD matrix. In order to obtain the Generalized Least
Square (GLS) estimators of the above listed parameters, a two-stage algorithm is proposed
which integrates GLS estimation into the SUE traffic assignment procedure. The first and sec-
ond stages of the algorithm are applied iteratively until the maximum relative difference pre-
sented in Step 11 is achieved, after which the estimated OD matrix, link choice proportion, and
dispersion parameter θ can be obtained. The SQP approach based on the extended quasi-New-
ton method is used to search for the optimal solution in the first stage of the algorithm. The
SUE traffic assignment procedure is applied to incorporate both OD matrix and link choice
proportion estimation into the second stage of the algorithm, and MSA is used to obtain the
equilibrium link flows.
A hypothetical network was constructed to test the performance of the proposed approach,
followed by a comprehensive sensitivity analysis with 50 combinations of variation coefficient

PLOS ONE | DOI:10.1371/journal.pone.0146850 January 13, 2016 20 / 24


Origin-Destination Matrices Estimation

combinations cvd (CVD) and cvθ (CVT) to investigate the stability of the estimated OD matrix,
link flows, and Theta. A comparison with two different methods described in Yang et al. [15]
and Lo and Chan [16] suggests that the proposed approach can achieve superior performance
in terms of RMSE (OD), RMSE (LF), and accuracy of the estimated Theta parameter. More-
over, a case study is presented using a real-world congested square network in Seattle, WA to
demonstrate the practicality of the proposed approach, in which the true OD matrix and
observed link flows are calculated via ground-truth traffic count data collected by loop detec-
tors. The proposed method is shown to be robust under a range of initial parameter values. The
RMSE (OD) can be reduced from 3426.9 to 23.6 at cvd = 0.1 and cvθ = 0.1 when traffic flows are
observed on five out of eight links. In addition, the estimated dispersion parameter exhibits a
consistent and regular trend by time of day for all combinations of initial parameters. For
future research, the proposed approach should be tested on a network of greater complexity
and size, and the impact of input data inaccuracy should be considered. Additionally, further
work is needed to determine the number and location of observed links required for accurate
OD estimation using the proposed approach.

Supporting Information
S1 Dataset. The dataset includes the Link Speed Data and Link Volume data, and the data
were collected from loop detectors located along the freeway section (I-5, I-90, I-405 and
SR 520) in Seattle area, and are retrieved via the Strategic Highway Research Program 2
(SHRP 2 program). The file named as “S1 Link Speed Data” records the average speed for all
links every 20-second time interval, and the other file named as “S1 Link Volume data” records
volume for all links every 20-second time interval.
(RAR)

Acknowledgments
The helpful comments from the two anonymous reviewers are gratefully acknowledged. This
research is supported by National Natural Science Foundation of China (Project No. 71402011,
71471024, 51408019, 71301180, 51329801), National Social Science Foundation of Chongqing
of China (No. 2013YBJJ035), and the Scientific and Technological Research Program of
Chongqing Municipal Education Commission (No. KJ1400307), and the Natural Science
Foundation of Chongqing of China(No. cstc2015jcyjA30012), National Key Technologies
R&D Program of China (2014BAG01B03), Science and Technology Project on Transportation
Construction by the Ministry of Transport of China (2015318835200).

Author Contributions
Conceived and designed the experiments: Yong W. XM Yinhai W. Performed the experiments:
Yong W. XM YL KG. Analyzed the data: Yong W. XM YL KG MX. Contributed reagents/
materials/analysis tools: KCH Yinhai W. Wrote the paper: Yong W.

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