Bounded Ornstein-Uhlenbeck models for two-choice time controlled tasks Jiaxiang Zhang a,b 1 , Rafal Bogacz a , a Department b School 1 of Computer Science, University of Bristol, Bristol BS8 1UB, UK of Psychology, University of Birmingham, Birmingham B15 2TT, UK Correspondence address: School of Psychology, University of Birmingham, Birmingham B15 2TT, UK. Email: [email protected] Preprint submitted to Elsevier 23 September 2009 Abstract The Ornstein-Uhlenbeck (O-U) model has been successfully applied to describe the response accuracy and response time in 2-alternative choice tasks. This paper analyses properties and performance of variants of the O-U model with absorbing and reflecting boundary conditions that limit the range of possible values of the integration variable. The paper focuses on choice tasks with pre-determined response time. The type of boundary and the growth/decay parameter of the O-U model jointly determine how the choice is influenced by the sensory input presented at different times throughout the trial. It is shown that the O-U models with two types of boundary hold close relationship and can achieve the same performance under certain parameter values. The value of the growth/decay parameter that maximizes the accuracy of the model has been identified. It is shown that when the boundaries are introduced, the O-U model may achieve higher accuracy than the diffusion model. This suggests that given the limited range of the firing rates of integrator neurons, the neural decision circuits could achieve higher accuracy employing leaky rather than linear integration in certain tasks. We also propose experiments that could distinguish between different models of choice in tasks with pre-determined response time. Key words: choice, decision, Ornstein-Uhlenbeck model, reflecting boundaries, absorbing boundaries 2 1 1 Introduction 2 Making choices on the basis of sensory information is one of the fundamental 3 cognitive functions of intelligent animals. The studies of decision process in 4 simple 2-alternative forced choice (2AFC) tasks usually employ one of two 5 paradigms. In the time controlled (TC) paradigm, subjects are required to re- 6 spond immediately after a cue presented at an experimenter-determined time 7 (Dosher, 1976, 1984; Swensson, 1972; Yellott, 1971). Alternatively, the infor- 8 mation controlled (IC) paradigm allows subjects to respond freely whenever 9 they feel confident in their choice (Luce, 1986). 10 Several mathematical models have been developed over the last half century 11 to describe the behavioural performance as well as underlying decision process 12 in 2AFC tasks (Busemeyer and Townsend, 1992, 1993; Laming, 1968; Ratcliff, 13 1978; Ratcliff et al., 1999; Stone, 1960; Usher and McClelland, 2001; Vickers, 14 1970). These sequential sampling models share a common characteristic that 15 the sensory representation of stimuli is accumulated over time to form a choice. 16 Neural correlates of such accumulation process have been observed in certain 17 cortical areas (Gold and Shadlen, 2002; Kim and Shadlen, 1999; Schall, 2001; 18 Shadlen and Newsome, 2001), which gives further support to the sequential 19 sampling framework. 20 In the IC paradigm, the decision process depends on the amount of accumu- 21 lated evidence, i.e., a choice is made as soon as sufficient evidence in favour 22 of one alternative is accumulated. In the TC paradigm, since the time of re- 23 sponse is chosen by the experimenter, a choice can be made by simply asking, 24 at the time of response, whether the accumulated evidence supporting the 3 25 first alternative exceeds that of the second. However, according to such model 26 for the TC paradigm the accumulated evidence could take arbitrarily large or 27 small values, which could lead to unrealistic predictions (Meyer et al., 1988). 28 One approach to overcome this problem is to assume that the accumulated 29 evidence is bounded within a certain range (Feller, 1968; Ratcliff, 1988). In 30 our previous study (Zhang et al., 2009), we compared the effects of two pos- 31 sible boundary mechanisms, reflecting and absorbing boundaries, on a typical 32 sequential sampling model - the diffusion model (Laming, 1968; Ratcliff, 1978; 33 Stone, 1960), and we showed that both boundary types lead to similar perfor- 34 mance in the TC paradigm and produce similar fits to experimental data. 35 This paper extends our previous findings by investigating the dynamics and 36 performance of the Ornstein-Uhlenbeck (O-U) model (Busemeyer and Diederich, 37 2002; Busemeyer and Townsend, 1992, 1993) with two types of boundaries in 38 the TC paradigm. The O-U model has been successfully applied to a vari- 39 ety of tasks (Diederich, 1995, 1997; Smith, 1995, 2000) and can approximate 40 the same computation carried out by other biologically inspired models (e.g., 41 Usher and McClelland, 2001) in 2AFC tasks. We show that the O-U model 42 with different types of boundaries could achieve the same performance under 43 certain parameter values, though the boundaries introduce differential weight- 44 ing of evidence. We also show that when the boundaries are introduced, the 45 accuracy of the O-U model is higher than that of the diffusion model. 46 The paper is organized as follows. Section 2 reviews the decision problem 47 and the O-U model as well as boundary mechanisms. Section 3 compares 48 the properties of the O-U models with two types of boundaries. Section 4 49 investigates the performance of the model and identifies the parameter values 50 that maximize accuracy. Section 5 discusses experimental predictions of the 4 51 O-U model with boundaries and Section 6 summarizes implications of our 52 study. Mathematical details are provided in Appendices. Preliminary results 53 of this study were presented in an abstract form (Zhang and Bogacz, 2006). 54 2 55 2.1 Neurobiology of decision 56 Numerous in vivo experiments have been performed to investigate the neural 57 correlates of decision processes in monkeys. In a well-known motion discrimi- 58 nation task used in this line of research, monkeys were instructed to fixate on 59 a random moving dots stimulus presented within a circular aperture (Britten 60 et al., 1993). A proportion of dots move coherently to the left or to the right 61 and the monkey has to respond to the coherent motion direction by making 62 a saccade to a left or right target. 63 Neural recording data from the motion discrimination task indicate that the 64 decision process involves several cortical regions. First, the neural activity 65 in sensory areas (e.g., the middle temporal (MT) area) has been shown to 66 closely correlate with the statistics of the stimulus (i.e., the motion coherence) 67 (Britten et al., 1992, 1993, 1996). Second, neurons in the lateral interparietal 68 (LIP) area that directly receive inputs from area MT gradually build up or 69 attenuate their activity within a trial (Roitman and Shadlen, 2002; Shadlen 70 and Newsome, 2001). In particular, neural activity in LIP correlates with the 71 direction of eye movements (i.e., the decision) at the time of response. These 72 results suggest that LIP neurons accumulate sensory evidence from area MT 73 to form a choice. Sequential sampling models and boundary mechanisms 5 74 The experimental results indicate that the accumulation of sensory evidence 75 may be a general decision mechanism manifested in different brain regions 76 or species (Gold and Shadlen, 2007; Schall, 2001). Based on this assumption, 77 in this paper we investigate decision problems in 2AFC tasks formalized as 78 following (Gold and Shadlen, 2001, 2002). Two populations of sensory neu- 79 rons generate continuous noisy evidence Y1 (t) and Y2 (t) supporting the two 80 alternatives Y1 and Y2 at time t. We assume that Y1 (t) and Y2 (t) are normally 81 distributed with constant means µ1 and µ2 (µ1 , µ2 > 0). The goal of the de- 82 cision process is to identify which sensory population has higher mean based 83 on Y1 (t) and Y2 (t). 84 2.2 The Ornstein-Uhlenbeck model 85 The Ornstein-Uhlenbeck (O-U) model (Uhlenbeck and Ornstein, 1930) is a 86 stochastic process that was first introduced into psychology by Busemeyer 87 and Townsend (1992, 1993). Since then many researchers applied the O-U 88 model to a variety of studies to account for response accuracy and response 89 time in choice tasks (Busemeyer and Diederich, 2002; Diederich, 1995, 1997; 90 Heath, 1992; Smith, 1995, 1998), and the model has also been extended to 91 multi-alternative tasks (McMillan and Holmes, 2006; Usher and McClelland, 92 2001). 93 In the O-U model for 2AFC tasks a non-perfect integrator accumulates the 94 difference of sensory evidence supporting two alternatives (Y1 (t) − Y2 (t)). Let 95 X(t) denote the value of the integrator state at time t. The O-U model is 96 described by the following stochastic differential equation: dX(t) = (λX(t) + µ) dt + σdW , 6 (1) 97 where µ and σ denote the mean and standard deviation of the normally dis- 98 tributed quantity Y1 (t)−Y2 (t) (i.e., µ = µ1 −µ2 ), dX(t) denotes the increment 99 of evidence over a small unit of time dt, dW (t) is Gaussian noise with mean 100 zero and unit variance, and λ is a growth/decay parameter. According to the 101 decision problem proposed above, the sign of µ solely determines the correct 102 choice. More specifically, µ > 0 implies that the first alternative Y1 is cor- 103 rect (µ1 > µ2 ), and µ < 0 implies that the second alternative Y2 is correct 104 (µ1 < µ2 ). For simplicity, we hereafter set µ > 0 in this paper. On the other 105 hand, the magnitude of µ and σ determine the task difficulty, i.e., for |µ| close 106 to zero or large σ, it is difficult to distinguish which alternative (Y1 or Y2 ) has 107 higher mean. On each trial, the decision process starts at t = 0 with a staring 108 point X0 = X(0) = 0. In the IC paradigm, the integrator continuously accu- 109 mulates evidence according to Eq. 1 until X(t) reaches a decision threshold. 110 In the TC paradigm, the decision process terminates at a pre-determined time 111 tc , and the choice is made on the basis of the final integrator state X(tc ), i.e., 112 the model selects Y1 if X(tc ) > 0, or Y2 if X(tc ) < 0. Throughout this paper 113 we focus on the TC paradigm and quantify the model’s performance by the 114 error rate P (tc ), which is the probability of making an incorrect choice when 115 the response is required at time tc . The expression for the error rate of the 116 O-U model is given by Busemeyer and Townsend (1993): v u µ u 2 (eλtc − 1) , P (tc ) = Φ − t σ λ (eλtc + 1) where Φ(u) = Z u −∞ v2 1 √ e− 2 dv . 2π (2) (3) 117 It is worth to note that the parameter λ affects the linear drift rate λX(t) + µ, 118 and as a result the O-U model could exhibit differential weighting of evidence 7 119 within a trial. That is, for λ > 0 the evidence presented early in a trial has a 120 larger influence on the decision (a primacy effect), and for λ < 0 the choice 121 mainly depends on the evidence presented later in a trial (a recency effect) 122 (Wallsten and Barton, 1982). An explicit proof of this property is available 123 in Busemeyer and Townsend (1993). In particular, for λ = 0, the weighting 124 of evidence is independent to the time of its presentation and the O-U model 125 simplifies to a diffusion model (Laming, 1968; Ratcliff, 1978; Stone, 1960). In 126 Section 3 we will investigate how the primacy and recency effects of the O-U 127 model are affected by boundaries. 128 2.3 Boundary mechanisms 129 In earlier versions of sequential sampling models, the value of the integra- 130 tor states was unconstrained. As mentioned in the Introduction, this type 131 of model would allow the integrator state to have arbitrarily large or small 132 values. This is, however, a simplification. First, it is not realistic that any bi- 133 ological system could represent an infinite amount of evidence. Second, when 134 accumulated evidence is not limited, a sequential sampling model may not fit 135 experimental data (Ratcliff, 1988). One way to eliminate the above problems 136 is to transform the integrator state through a nonlinear activation function 137 (Brown et al., 2005; Brown and Holmes, 2001; Usher and McClelland, 2001). 138 Another approach, the focus of this paper, is to apply boundary conditions to 139 the existing linear models, which provides an approximation of the nonlinear 140 activation functions (Usher and McClelland, 2001). 141 The first boundary mechanism is the absorbing boundary (Feller, 1968), which 142 was originally applied to choice tasks in which IC and TC paradigms were in8 143 termixed (Diederich and Busemeyer, 2003; Ratcliff, 1988; Ratcliff and Rouder, 144 2000). Once the integrator hits an absorbing boundary, the accumulation pro- 145 cess terminates and the integrator state is maintained afterwards. To intro- 146 duce the absorbing boundaries to the O-U model, we assume two symmetric 147 boundaries at ±b, and then, the O-U model with absorbing boundaries can 148 be described by: X(t + dt) = b, X(t + dt) = −b, X(t + dt) = X(t) + dX(t), if X(t) + dX(t) ≥ b or X(t) = b , if X(t) + dX(t) ≤ −b or X(t) = −b , (4) otherwise , 149 where dX(t) is given by Eq. 1. In the TC paradigm, the choice is determined 150 by which boundary the integrator hits first, or if neither boundary is reached 151 before tc , the choice is determined by the same rule as the one used for the 152 model without boundaries (see Section 2.2). 153 The second boundary mechanism is the reflecting boundary (Zhang et al., 154 2009) (also see Diederich, 1995; Diederich and Busemeyer, 2003; Ratcliff and 155 Smith, 2004), which only restricts the amount of accumulated evidence that 156 can be represented by the integrator. In other words, the value of the integrator 157 state cannot exceed a reflecting boundary, but it may continue to change (i.e., 158 move back) due to noise, as specified by: X(t + dt) = b, if X(t) + dX(t) ≥ b , X(t + dt) = −b, X(t + dt) = X(t) + dx(t), if X(t) + dX(t) ≤ −b , (5) otherwise . 159 Hereafter we refer to the O-U model with absorbing and reflecting bound- 160 aries as absorbing O-U and reflecting O-U models, and the model without a 9 161 boundary as unbounded O-U model. Fig. 1 shows sample paths of integrator 162 states in the O-U models with both types of boundaries, compared with the 163 unbounded O-U model. For the absorbing O-U model, the choice can be de- 164 termined early in a trial (after 0.35s in Fig. 1). For the reflecting O-U model, 165 the preferred choice may change throughout the decision process until the end 166 of the trial. 167 In a previous study, we compared the boundary effects on one sequential sam- 168 pling model - the diffusion model (Zhang et al., 2009). We showed that the 169 diffusion model with two types of boundaries produce exactly the same error 170 rate for any given tc . Moreover, the absorbing and reflecting boundaries yield 171 a primacy and recency effect, respectively. In the next section we will show 172 that similar properties also hold for the bounded O-U model. 173 2.4 Optimal decision making theory 174 Due to evolutionary pressure, neural systems are likely to adapt to their en- 175 vironment and optimize their behaviour to achieve desired goals (Anderson, 176 1990). This assumption has been invoked in the study of perceptual decision 177 making, and it has been proposed that the brain may implement statistically 178 optimal decision-making procedures (Bogacz, 2007, 2009; Bogacz et al., 2006; 179 Gold and Shadlen, 2001, 2002, 2007). In the sequential sampling framework, 180 an optimal decision procedure would achieve the lowest error rate and short- 181 est response time compared with all possible procedures to integrate sensory 182 evidence 1 . For the TC paradigm, the optimal procedure is the one that yields 1 The optimality criterion for the IC paradigm is to consider which decision strategy yields the fastest response time for any given error rate. The procedure that satisfies 10 183 the lowest error rate for any given response time. 184 Recall that the decision problem proposed in Section 2.1 is to identify which of 185 the two alternatives has higher mean inputs. This problem can be converted 186 to the problem of hypothesis testing in statistics (Ghosh, 1970; Lehmann, 187 1959), and the optimal procedure satisfying the above criterion is provided 188 by the Neyman and Pearson (NP) procedure. It has been proved that among 189 all hypothesis tests, the NP procedure minimizes the overall probability of 190 accepting the false hypothesis for fixed sample size (Neyman and Pearson, 191 1933), and therefore it minimizes the error rate for any response time. Bogacz 192 et al. (2006) showed that as the time intervals between samples approach 193 zero, the NP procedure becomes equivalent to the diffusion model. Hence, the 194 diffusion model is the optimal decision procedure in the TC paradigm. Since 195 the O-U model with λ = 0 can be reduced to the diffusion model, the O-U 196 model also maximizes accuracy when λ = 0. In Section 4, we will show that 197 this is no longer the case for the bounded O-U model. 198 3 199 3.1 Dynamics of bounded O-U models 200 The dynamics of the absorbing and reflecting O-U models can be illustrated by 201 the probability density p(X, t) of the integrator states X(t) at time t, which is 202 calculated in Appendix A by solving the Kolmogorov backward equation with 203 boundary conditions. The density function p(X, t) of bounded O-U models is Properties of bounded O-U models this criterion is provided by the Sequential Probability Ratio Test (Barnard, 1946; Wald, 1947). 11 204 a weighted sum of an infinite series: p(X, t) = ∞ X e−ξj t Kj (X)φj (X0 ) , (6) j=0 205 where Kj (X) is defined in Equation A.16. ξj are the non-negative eigenvalues 206 from Eqs. A.27 and A.30, and φj (X0 ) are the associated eigenfunctions defined 207 in Eqs. A.26 and A.29 as a function of the starting point X0 . The solutions of 208 the eigenvalues and eigenfunctions depend on the type of boundaries. There 209 exist infinitely many eigenvalues that can be found numerically, and an ap- 210 proximation of p(X, t) can be obtained by taking a partial sum of finite terms. 211 The more eigenvalues included in Eq. 6, the more accurate the approximation 212 of the density function p(X, t). 213 Fig. 2 compares the probability densities of the absorbing and reflecting O- 214 U models at several time instants. During the first hundred milliseconds, all 215 densities are close to a Gaussian distribution with mean close to the starting 216 point X0 and shifted towards the upper boundaries as time grows. For large t, 217 the density of the absorbing O-U model collapses to zero, while the reflecting 218 O-U model has nonzero equilibrium distributions, with a small proportion to 219 the left of the origin, corresponding to the probability of making an incorrect 220 choice (for µ > 0). 221 The difference between the stationary densities can be explained from the 222 expression of p(X, t). For any ξj > 0, the time-dependent coefficient e−ξj t in 223 Eq. 6 monotonically decreases and converges to zero as t grows. Table 1 shows 224 the first ten eigenvalues ξj . For the absorbing O-U model all eigenvalues are 225 positive, hence the density of the absorbing O-U model converges to p(X, t) = 226 0 as t → ∞. For the reflecting O-U model, one of the eigenvalues is equal to 227 zero ξ0 = 0, hence the density p(X, t) converges to K0 (X)φ0 (X0 ) as t → ∞. 12 228 It is also interesting to note that the eigenvalues ξj (j > 0) of the absorbing 229 and reflecting O-U models are symmetric by flipping the sign of λ. 230 3.2 Primacy and recency effects 231 Previous studies showed that differential weighting of evidence (i.e., the pri- 232 macy and recency effects) can be introduced by either the parameter λ of 233 the unbounded O-U model (Busemeyer and Townsend, 1993) or two types 234 of boundaries (Zhang et al., 2009). Here we investigate the primacy and re- 235 cency effects in the O-U models by applying the reverse correlation method. 236 We simulate the unbounded, absorbing, and reflecting O-U models, each with 237 positive and negative λ values. For each model considered, the time course of 238 sensory evidence is recorded and averaged only from trials resulting in correct 239 choices. The averaged evidence represents how the model weights noisy inputs 240 during the decision process. For µ > 0, the model makes correct decision if the 241 integrated evidence is positive at the end of a trial. Hence a larger averaged 242 input from correct trials indicates that the sensory input at that time point 243 has, on average, a large influence on the final choice, and a smaller averaged 244 input indicates that the choice depends to a lesser extent on the input at that 245 time. 246 The simulation results are shown in Fig. 3. First, positive and negative λ values 247 introduce primacy (Fig. 3a) and recency (Fig. 3d) effects to the unbounded 248 O-U model, which is consistent with theoretical predictions by Busemeyer and 249 Townsend (1993). Second, the absorbing O-U model exhibits a strong primacy 250 effect for λ > 0 (Fig. 3b), but no clear effect was observed for λ < 0 (Fig. 251 3e). By contrast, a positive λ value does not introduce a clear effect to the 13 252 reflecting O-U model (Fig. 3c), but the reflecting O-U model has a strong 253 recency effect for λ < 0 (Fig. 3f). 254 Therefore, the primacy/recency effect in the bounded O-U models is jointly 255 determined by two independent elements: the value of λ and the type of 256 the boundary. Given different combinations of the two elements, the pri- 257 macy/recency effect in the bounded O-U model is maintained if λ and the 258 boundary provide the same effects (Fig. 3b and 3f). On the other hand, the 259 joint effect is weakened if λ and the boundary introduce opposite effects, as 260 shown in Fig. 3c and 3e. By appropriately setting the two parameters the 261 joint primacy/recency effects introduced by the boundary and λ could cancel 262 each other. In Section 4.2 we show that this condition leads to the optimal 263 performance of the bounded O-U model. 264 4 265 4.1 The error rate of bounded O-U models 266 For the reflecting O-U model, since the density of the integrator p(X, t) has 267 unit mass (see Fig. 2), the error rate is equal to the proportion of p(X, t) that 268 lies on the ’error’ side of the origin, i.e., if λ > 0, the error rate of the reflecting 269 O-U model can be calculated by integrating the density p(X, t) from −b to 0: Performance of bounded O-U models P(ref) (tc ) = Z 0 −b p(X, tc ) dX , (7) 270 where (ref) stands for reflecting boundary. Therefore by estimating p(X, t) nu- 271 merically we can obtain an approximation of the error rate. For the absorbing 272 O-U model, such calculation is not available as p(X, t) converges to zero as t 14 273 grows. 274 Now let us compare the error rates of the O-U models with two types of 275 boundaries. For the unbounded O-U model, the error rate remains the same 276 if we flip the sign of λ (see Eq. 2). Fig. 4 shows the error rates of the bounded 277 O-U models with positive and negative λ at different tc . The result suggests 278 that there exists error rate symmetry in the bounded O-U models. That is, by 279 flipping the sign of λ, the absorbing and reflecting O-U models produce the 280 same error rate for any tc . This relationship can be represented as: P(abs) (tc , −λ) = P(ref) (tc , λ) , (8) 281 where P(abs) (.) and P(ref) (.) denote the error rates of the absorbing and 282 reflecting O-U models with parameters specified in the brackets. We have not 283 succeeded in giving a complete proof of Eq. 8 since an analytical expression 284 for the error rate is not available. However we are able to prove a special case 285 of Eq. 8 when tc → ∞, which is given in Appendix B. 286 In summary, the absorbing and reflecting O-U models with opposite signs of λ 287 produce the same accuracy, and thus they would provide the same fit to data 288 from experiments in TC paradigm. 289 4.2 The optimal parameter 290 In the TC paradigm, the unbounded O-U model achieves the minimum error 291 rate when λ = 0 (i.e., when it reduces to the diffusion model) (Bogacz et al., 292 2006, and also see Section 2.4). After introducing boundaries, an intriguing 293 question arises: is λ = 0 still the optimal parameter value for the bounded 294 O-U models? If not, what is the optimal λ value that yields the lowest error 15 295 rate for the bounded O-U models? 296 To answer this question, we simulate the bounded O-U models with different 297 parameter values. The error rates of the bounded O-U models with different 298 values of λ, b and tc are shown as contour plots in Fig. 5. The result suggests 299 that the bounded O-U models with λ = 0 are not always optimal, but the 300 optimal value of λ depends on the values of other parameter such as b and 301 tc . First, if b is relatively small (b < 0.2 in Fig. 5), the value of λ does not 302 significantly affect the performance of the bounded O-U models in all panels of 303 Fig. 5. This result is expected since if the interval between the two boundaries 304 is tiny, the integrator state is restricted to be around X0 , and in this case 305 the noise would dominate the choice process. Therefore, the models produce 306 large error rates (around 40%) in this region. Second, if b is sufficiently large 307 (b > 2 in Fig. 5), the integrator hardly reaches any of the boundaries before tc 308 used in the simulation, and hence the bounded O-U models are approximately 309 equivalent to the unbounded O-U model, in which λ = 0 gives the lowest error 310 rate for any tc . 311 Except for these two extreme parameter regions, the error rate is affected by 312 the values of λ, b and tc . In general, given b and tc , for the absorbing O-U 313 model, λ < 0 gives the best performance, while for the reflecting O-U model, 314 λ > 0 yields the minimum error rate. This optimal value of λ is consistent 315 with the joint primacy/recency effects of the bounded O-U models proposed in 316 Section 3.2. Recall that the recency effect introduced by the reflecting bound- 317 ary can be weakened by setting λ > 0, and vice versa. To achieve the optimal 318 performance, the primacy/recency effects of the boundaries and λ should be 319 balanced and cancelled, so that the sensory evidence from different time points 320 contribute equally to the choice, as in the optimal NP decision procedure (i.e., 16 321 no primacy/recency effects). The simulation results in Fig. 5 accord with this 322 hypothesis. 323 Fig.5 also shows that for fixed b, the absolute value of λ that yields the optimal 324 performance increases when extending tc , i.e., the optimal λ increases for the 325 reflecting O-U model and decreases for the absorbing O-U model. To find the 326 optimal λ values, we simulate the bounded O-U model under different tc . For 327 each tc , we numerically find the value of λ that yields the minimum error 328 rate, and the results are shown in Fig. 6a. For the reflecting O-U model, the 329 optimal λ increases as tc increases, with decreasing slope. The optimal λ for the 330 absorbing O-U model is a mirror of that for the reflecting O-U model, which 331 decreases as tc grows. This result further supports the error rate symmetry of 332 the bounded O-U models (Eq. 8), because if a specific λ value is optimal for 333 the reflecting O-U model, −λ would be optimal for the absorbing O-U model. 334 To validate the optimality of λ values in Fig. 6a, we simulated the reflecting 335 O-U model with different tc . For each tc , λ is set to be the optimal value 336 shown in Fig. 6a. For comparison, we also simulated the reflecting O-U model 337 with fixed λ values λ = 3 and λ = 0. The error rates of the three models are 338 illustrated in Fig. 6b. The bounded O-U model with optimal λ values achieves 339 lower error rates than the model with other constant λ. 340 5 341 The two types of boundaries may interpret different subjects’ behaviour in 342 psychological experiments. The absorbing boundary can account for the be- 343 haviour such that the choice is made before the end of the trial and never Experimental predictions 17 344 changed. The reflecting boundary may imply that the subjects hesitate be- 345 tween the two choices even when sufficient evidence is available, and may 346 change their preference later in the trial. Nevertheless, due to the error rate 347 symmetry, it is not trivial to differentiate between the two types of boundary 348 on the basis of experimental data from simple 2AFC tasks, as the absorbing 349 and reflecting O-U models (with opposite sign of λ) would always produce the 350 same fit. 351 To investigate primacy and recency effects in the TC paradigm, stimuli could 352 be used that differ in information they contain at different times after stimulus 353 onset (Huk and Shadlen, 2005; Usher and McClelland, 2001). For example, in 354 the motion discrimination task (see Section 2.1) the coherence of motion can 355 be increased for a short period on each trial (Usher et al., 2009). The primacy 356 effect would be present, if such increase in coherence had the highest influence 357 on accuracy when occurring after stimulus onset. The recency effect would 358 be present, if the increase had the highest influence before the response. Note 359 however, that such experiment cannot differentiate between the absorbing O-U 360 model with negative λ and the reflecting O-U model with positive λ, because 361 both models do not predict strong primacy or recency effects. Our analysis 362 suggests that these two models maximize accuracy, and given evolutionary 363 pressure for accuracy of decisions, we predict that one of these two models 364 should describe choices in the TC paradigm with long tc . 365 We propose that the absorbing O-U model with negative λ and the reflecting 366 O-U model with positive λ could be distinguished in a motion discrimination 367 experiment in which the coherence is manipulated in a similar way to that 368 recently proposed by Zhou et al. (2009). In particular, the coherence could 369 be increased by a strong pulse c1 for an interval of length t1 , and then de18 370 creased by a smaller amount c2 < c1 for a longer interval t2 > t1 (Fig. 7a). If 371 the parameters of the perturbation satisfied c1 t1 = c2 t2 , the average motion 372 coherence in a trial would remain unchanged (hence the unbounded diffusion 373 model predicts that such perturbation would have no effect on accuracy). The 374 absorbing O-U model with negative λ predicts that such perturbation would 375 increase accuracy, because the first pulse could cause the correct absorbing 376 boundary to be reached. By contrast the reflecting O-U model with positive λ 377 predicts that the perturbation would decrease accuracy, because the informa- 378 tion in the first pulse may be lost if the reflecting boundary is reached. Fig. 7b 379 verifies these predictions by simulations. Note that the effects of the pulse per- 380 turbation are not due to the primacy/recency properties of the models (as we 381 intentionally selected the boundary and λ values so that the primacy/recency 382 effects were balanced), but originate from the characteristics of the two types 383 of boundaries. However, it is worth to mention that the effect of the pulse only 384 occurs if the following two conditions are satisfied. First, the pulse needs to 385 be sufficiently large to move the integrator to the boundary. Second, the pulse 386 needs to be presented early in a trial. Otherwise the pulse will not affect the 387 performance of the absorbing O-U model (as the inputs do not affect decision 388 process once the integrator hits absorbing boundaries). 389 Our analysis also shows that the accuracy in the TC paradigm is maximized if 390 parameter λ is adapted depending on the required reaction time tc . If subjects 391 indeed adapt λ depending on tc , then they should achieve higher accuracy 392 when tc can be anticipated. This predicts that the accuracy for a given tc 393 should be higher when tc is constant within a block (and hence can be antici- 394 pated) than when it varies within a block. 19 395 6 Discussion 396 The present study compared the properties and performance of the O-U mod- 397 els with absorbing and reflecting boundaries in the TC paradigm. The main 398 results are summarized in Table 2. First, we showed how different boundaries 399 affect the dynamics of decision process by visualizing the probability density 400 of the integrator state (Fig. 2). We then showed that the primacy and re- 401 cency effects introduced by the boundaries and the parameter λ coexist in 402 the bounded O-U models, and the joint effects depend on the two indepen- 403 dent elements (Fig. 3). Moreover, the absorbing and reflecting O-U models 404 can achieve the same error rate for any tc by flipping the sign of λ (Fig. 4). 405 Finally, we showed that the reflecting O-U model with positive λ (and the 406 absorbing O-U model with negative λ) produces lower error rates than the 407 model with λ = 0 (Fig. 5), and we numerically found the optimal λ values for 408 the bounded O-U models that yield the minimum error rate (Fig. 6). 409 6.1 Optimal integration with boundaries 410 This paper has shown that when the range of possible values of integrator 411 state is constrained, the simple integration of difference in evidence is not the 412 optimal procedure, and a better performance can be yield by the O-U model, 413 especially for long tc . This suggests that given the limited range of the firing 414 rates of integrator neurons, the neural decision circuits could achieve a higher 415 accuracy in tasks with long tc employing an integration procedure similar to 416 the O-U model rather than the linear integration. In particular, if the brain 417 employs a mechanism similar to the absorbing boundaries, as suggested by 20 418 neurophysiological data of Kiani et al. (2008), the accuracy of choices in task 419 with long tc would be maximized by employing leaky integration. 420 6.2 Relationship to other studies 421 Since the O-U model is reduced to the diffusion model when λ = 0, the results 422 presented in this paper extend our previous findings on the diffusion model 423 with boundaries (Zhang et al., 2009). In particular, Zhang et al. (2009) showed 424 that the diffusion models with absorbing and reflecting boundaries achieve the 425 same error rate. Such equality is a special case of the error rate symmetry of 426 the bounded O-U models by setting λ = −λ = 0. 427 In the TC paradigm, the O-U model produces the same simulated behaviour 428 as the leaky competing accumulator (LCA) model (Usher and McClelland, 429 2001), when λ parameter of the O-U model is set to the difference between 430 inhibition and decay parameters of the LCA model. Simulations of the LCA 431 model with attracting boundaries showed that it maximizes the accuracy in 432 the TC paradigm when the inhibition is lower than the leak (Bogacz et al., 433 2007). These simulations are consistent with the results of this paper, because 434 for such parameter values the LCA model corresponds to the O-U model with 435 λ < 0. 436 Acknowledgement 437 This work was supported by the ORS grant and the EPSRC grant EP/C514416/1. 21 j Absorbing Reflecting λ = −1 λ=1 λ = −1 λ=1 0 - - 0 0 1 1.26 2.26 2.26 1.26 2 5.08 6.08 6.08 5.08 3 11.27 12.27 12.27 11.27 4 19.90 20.90 20.90 19.90 5 31.01 32.01 32.01 31.01 6 44.58 45.58 45.58 44.58 7 60.62 61.62 61.62 60.62 8 79.12 80.12 80.12 79.12 9 100.10 101.10 101.10 100.10 10 123.54 124.54 124.54 123.54 Table 1 Eigenvalues of the bounded O-U models with positive and negative λ. The eigenvalues are numerically obtained by seeking the roots of the Eqs. A.27 and A.30. The parameters of the bounded O-U models are µ = σ = 1 s−1 , and b=1. 22 Absorbing Reflecting λ>0 uniform recency λ<0 primacy uniform λ=0 primacy recency Weighting of inputs Error rate λ 6= 0 symmetric λ=0 equality Optimality λ<0 λ>0 Table 2 Comparison of the properties of the absorbing and reflecting O-U models. 2. 5 absorbing O−U reflecting O−U O−U Integrator states 2 1. 5 1 0. 5 0 −0. 5 0 0. 2 0. 4 0. 6 Time (s) 0. 8 1 Fig. 1. Examples of trajectories of the O-U models with absorbing boundary (dashed line), reflecting boundary (solid line), and without boundary (dotted line). Models were simulated with µ = 0.5 s−1 , λ = 1, σ = 1 s−1 , dt = 0.001 s, and boundaries b = 1. 23 b a p(X,t) 1. 2 0. 8 Time (s) 0. 1 0. 3 0. 5 0. 7 1. 2 0. 8 0. 4 0 −1 0. 4 −0. 5 0 0. 5 p(X,t) 2. 5 2 0 −1 1 c 3. 5 3 Time (s) 0. 1 0. 3 0. 5 0. 7 −0. 5 0 0. 5 1 −0. 5 0 0. 5 Integrator states 1 d Time (s) 0. 1 0. 3 0. 5 0. 7 1. 2 0. 8 Time (s) 0. 1 0. 3 0. 5 0. 7 1. 5 1 0. 4 0. 5 0 −1 −0. 5 0 0. 5 Integrator states 1 0 −1 Fig. 2. Probability densities of integrator states in the bounded O-U models. State values of the integrator are shown on horizontal axes. Densities are shown for parameters µ = σ = 1 s−1 , and b=1. For each panel, different curves show densities at different time instants from 0.1s to 2s. The density functions p(X, t) are calculated in Appendix A with eigenvalues shown in Table 1. All solutions start at X0 = 0. (a) The absorbing O-U model with λ=1. (b) The absorbing O-U model with λ=-1. (c) The reflecting O-U model with λ=1. (d) The reflecting O-U model with λ=-1. 24 unbounded absorbing reflecting (a) (b) (c) average input 0.03 0.02 0.01 0 0 0. 5 1 0 0. 5 1 0 (e) (d) 0. 5 1 (f) average input 0.03 0.02 0.01 0 0 0. 5 1 0 0. 5 Time (s) 1 0 0. 5 Fig. 3. Primacy and recency effects for the bounded and unbounded O-U models with positive λ (top panels) and negative λ (bottom panels). In each panel the O-U model is simulated for 10,000 trials. For all correct trials the input to the O-U model at every time step is recorded and averaged. The data points show the mean of the inputs and the error bars show the standard error. In all panels the parameter values are µ=0.71 s−1 , σ=1 s−1 , b=0.47, tc =1s. (a) The unbounded O-U model with λ=5.5. (b) The absorbing O-U model with λ=5.5. (c) The reflecting O-U model with λ=5.5. (d) The unbounded O-U model with λ=-5.5. (e) The absorbing O-U model with λ=-5.5. (f) The reflecting O-U model with λ=-5.5. 25 1 0. 5 abs O−U ref O−U 0. 4 P(tc) 0. 3 0. 2 0. 1 0 0 1 2 3 tc(s) Fig. 4. The error rate of the bounded O-U models at different tc . The models were simulated with parameters: µ = σ = 1 s−1 and b=1. Every data point is averaged from 100,000 trials. The reflecting O-U model (solid line) is simulated with λ=2, and the absorbing O-U model (dashed line) is simulated with λ=-2. 26 0 0.1 0.2 0.3 0.4 Absorbing Reflecting λ (tc=0.5s) 2 0 −2 λ (tc=1s) 2 0 −2 λ (tc=4s) 2 0 −2 1 2 3 Boundary 1 2 3 Boundary Fig. 5. The contour plots of the error rates of the bounded O-U models with different parameter values. The bounded O-U models are simulated with the following parameters: λ in [-3,3] with step 0.1, b in [0.1,3] with step 0.1, µ = σ = 1 s−1 , and tc =0.5s, 1s, and 4s. For each possible combination of the parameter values, the absorbing and reflecting O-U models are simulated for 10,000 trials to obtain the mean error rates. The error rates are then grouped by the value of tc and the type of boundaries, and illustrated as contour plots. The horizontal axes show the value of b, and the vertical axes show the value of λ. Each contour plot has 20 contour levels and the colormaps are the same for all panels. Left panels show the error rates of the absorbing O-U model, and right panels show the error rates of the reflecting O-U model. The panels in each row show the plots that have the same value of tc , as indicated on the vertical axes. 27 a b 3 2 0. 5 λ =3 λ =0 optimal λ 0. 4 reflecting O−U absorbing O−U 0 P (tc) optimal λ 1 0. 2 −1 −2 −3 0. 3 0. 1 1 2 3 tc(s) 4 5 0 1 2 3 tc(s) 4 5 Fig. 6. Optimal performance of the bounded O-U models. (a) The optimal λ values of reflecting and absorbing O-U models for different tc varying from 0.5s to 10s (in steps of 0.01s). For each tc , the absorbing and reflecting O-U models are simulated with parameters µ = σ = 1 s−1 , b=1, and λ varying from -100 to 100. For each value of λ the model is simulated for 10,000 trials, and the λ that yields minimum error rate is recorded as the optimal value at tc . (b) Error rates of the reflecting O-U model at different tc . The model is simulated with the same parameters (µ, σ, b) as in panel (a). Dotted line shows the error rate of the model with λ=3. Dashed line shows the error rate of the model with λ=0 (i.e., the diffusion model). Solid line shows the error rate of the model with optimal λ values from panel (a). Each data point is averaged over 10,000 trials. 28 a b 8 t1 P (tc) Drift rate 0.06 c1 t0 t2 2 0 0 no−pulse pulse 0.08 6 4 0.1 0. 5 1 Time (s) c2 0.04 0.02 1. 5 0 diffusion abs O−U ref O−U Fig. 7. The decision task with pulse perturbation. (a) The drift rate with pulse perturbation. The parameter values are: µ=2 s−1 , c1 =5 s−1 , c2 =1.19 s−1 , t0 =310 ms, t1 =150 ms, t2 =630 ms. (b) The error rate of the absorbing and reflecting O-U models, compared with that of the unbounded diffusion model. The models were simulated with parameter values: b=0.7, λ=±5 (-5 for the absorbing O-U model and 5 for the reflecting O-U model), c = 2 s−1 , and tc =1.8 s. Light grey bars show the error rate of the models with constant drift rate µ=2 s−1 , and dark grey bars show the error rate of the models with time-varying drift rate from (a). The error bars show the standard error from 10,000 trials. 29 438 A The probability density function for the bounded O-U model 439 This appendix calculates the probability density function p(X, t) for the O-U 440 models with absorbing and reflecting boundary conditions. Recently, a solution 441 of this problem for λ < 0 has been provided by Saphores (2005). Here we 442 reiterate the calculation and extend the solution to both positive and negative 443 λ. 444 Let us consider a general case: an O-U process defined in Equation 1 is bounded 445 over [L, R], where L and R are finite absorbing or reflecting boundaries defined 446 in Equations 4 and 5, respectively. Let p(X, X0 , t) be the probability density 447 of X(t) at time t given that X(0) = X0 . The density satisfies the Kolmogorov 448 backward equation (Karlin and Taylor, 1981): à ∂ p(X, X0 , t) ∂ p(X, X0 , t) σ2 = −(µ + λX0 ) + ∂t ∂ X0 2 ! ∂ 2 p(X, X0 , t) , ∂ X02 (A.1) 449 where the coefficients µ, λ, and σ are defined in Equation 1. Since X(t) has 450 fixed value X0 at t = 0, the initial probability distribution is a delta function: p(X, X0 , 0) = δ(X − X0 ) . (A.2) 451 Goel and Dyn (2003) showed that if boundaries are absorbing, p(X, X0 , t) has 452 to satisfy: p(X, X0 , t) = 0 , for X0 = L or R , (A.3) 453 and reflecting boundaries at L and R imply the following no-flux boundary 454 conditions: ∂ p(x, y, t) = 0 , for X0 = L or R . ∂y 455 (A.4) To solve Equation A.1 we separate variables (Boyce and DiPrima, 1997) and 30 456 seek a solution as the weighted sum of elementary solutions. p(X, X0 , t) = ∞ X aj (t) φj (X0 ) . (A.5) j=0 457 For one of the elementary solutions a(t) φ(X0 ), substituting Equation A.5 in 458 A.1 and rearranging terms, we obtain: a0 (t) σ2 µ + λX0 0 = φ00 (X0 ) + φ (X0 ) . a(t) 2φ(X0 ) φ(X0 ) (A.6) 459 Since the left side of Equation A.6 depends only on t, while the right side only 460 on X0 , they must equal to some constant, denoted by −ξ. Then solution for 461 the time-dependent coefficients a(t) is: a(t) = Ke−ξt , 462 (A.7) and the function φ(X0 ) satisfies: σ 2 00 φ (X0 ) + (µ + λX0 )φ0 (X0 ) + ξφ(X0 ) = 0 . 2 463 (A.8) Substituting A.7 into A.5, the density p(X, X0 , t) can be represented as: p(X, X0 , t) = ∞ X Kj e−ξj t φj (X0 ) . (A.9) j=0 464 The coefficients Kj in Equation A.9 are obtained from the initial probability 465 at t = 0. Firstly multiply both side of A.8 by a function of X0 : à ! 2µX0 + λX02 2 , r(X0 ) = 2 exp σ σ2 466 (A.10) then φ(X0 ) should satisfy à ! à ! 2µ + 2λX0 2µX0 + λX02 0 2µX0 + λX02 00 φ (X ) + exp φ (X0 ) + ξr(X0 )φ(X0 ) exp 0 σ2 σ2 σ2 " à ! #0 2µX0 + λX02 0 = exp φ (X0 ) + ξr(X0 )φ(X0 ) = 0 . σ2 (A.11) 31 467 Equation A.11 is the classical Sturm-Liouville equation. From the Sturm- 468 Liouville theorem (Boyce and DiPrima, 1997), there exists an infinite set 469 of non-negative eigenvalues ξj (j = 1, 2, . . . ), and associated eigenfunctions 470 φj (X0 ) that satisfy Equation A.11. Besides, the normalized eigenfunctions 471 form an orthogonal set with respect to the function r(X0 ). Hence for any two 472 eigenfunctions φi (X0 ) and φj (X0 ), there is Z R L r(X0 )φj (X0 )φi (X0 ) dX0 Z R L = δij , r(X0 )φ2j (X0 ) dX0 (A.12) 473 where δij is the Kronecker delta function. For t = 0, a single elementary 474 solution of the density p(X, X0 , t) in Equation A.5 is: p(X, X0 , t) = Kj φj (X0 ) = δ(X − X0 ) . 475 Multiplying both sides of Equation A.13 by: Z R L 476 r(X0 )φi (X0 ) (A.14) , r(X0 )φ2j (X0 ) dX0 and taking integral over the interval [L, R], we obtain: Z R Kj L Z R r(X0 )φj (X0 )φi (X0 ) dX0 Z R L 477 (A.13) = L r(X0 )φi (X0 )δ(X − X0 ) dX0 Z R r(X0 )φ2j (X0 ) dX0 L , (A.15) r(X0 )φ2j (X0 ) dX0 and the coefficients Kj is: Kj = Z r(X)φj (X) R L . (A.16) r(X0 )φ2j (X0 ) dX0 478 To get the solutions of eigenvalues ξ and eigenfunctions φ(X0 ), we need to 479 solve the differential equation A.8. First assume λ < 0, and define a new 480 variable z: z= √ −2λ X0 + µ/λ . σ 32 (A.17) 481 Substituting Equation A.17 into A.8 yields: λφ00 (z) − λzφ0 (z) − ξφ(z) = 0 . 482 (A.18) We seek the solution of φ(z) of the form: φ(z) = ez 2 /4 ω(z) . (A.19) 483 Substituting Equation A.19 in A.18, the exponential term is canceled and the 484 new function is: à ! 1 ξ 1 d2 ω(z) + − − z 2 ω(z) = 0 . 2 dz 2 λ 4 (A.20) 485 Equation A.20 is the classical parabolic cylinder function, and there exists two 486 independent solutions, given by (Abramowitz and Stegun, 1965): ! ¶ à µ 2 ξ 1 z 1 2 ω1 (z) = exp z F ; ; , 4 2λ 2 2 ! µ ¶ à 1 2 ξ 3 z2 1 , z F + ; ; ω2 (z) = z exp 4 2 2λ 2 (A.21) 2 487 where F(.) is the confluent hypergeometric function. Representing Equation 488 A.21 as a function of X0 yields: ! à 1 λ ξ 2 ; ; − 2 (X0 + µ/λ) , E1 (X0 ) = F 2λ 2 σ à ! √ ξ 3 λ X0 + µ/λ 1 2 F + ; ; − 2 (X0 + µ/λ) . E2 (X0 ) = −2λ σ 2 2λ 2 σ 489 For the case of λ > 0, select the substitution of X0 as: z= 490 (A.22) √ 2λ X0 + µ/λ . σ (A.23) Then by following the same approach, we can obtain the elementary solutions 33 491 for φ(X0 ): à ! à ! (µ + λX0 )2 1 ξ 1 λ 2 E1 (X0 ) = exp − F (X0 + µ/λ) , − ; ; λσ 2 2 2λ 2 σ 2 à ! à ! 2 (µ + λX ) ξ 3 λ 0 F 1 − ; ; 2 (X0 + µ/λ)2 . E2 (X0 ) = exp − 2 λσ 2λ 2 σ (A.24) 492 The solution of Equation A.8 is the linear combination of the two elementary 493 solutions given by A.22 or A.24 (depending on the sign of λ). For absorbing 494 boundaries, φ(X0 ) must satisfy the following boundary condition (cf. Equa- 495 tions A.3 and A.5): φ(L) = φ(R) = 0 . (A.25) 496 By appropriately selecting the weighting value, the solution of φ(X0 ) for ab- 497 sorbing boundary condition is: φ(X0 ) = E1 (X0 )E2 (L) − E1 (L)E2 (X0 ) , (A.26) 498 and the eigenvalues ξ can be obtained numerically by seeking the roots of the 499 equation: E1 (L)E2 (R) − E1 (R)E2 (L) = 0 . 500 If the boundaries are reflecting, the boundary condition changes to: ∂ φ(L) ∂ φ(R) = = 0. ∂ X0 ∂ X0 501 (A.27) (A.28) The above condition is satisfied when: φ(X0 ) = E1 (X0 )E20 (L) − E10 (L)E2 (X0 ) , 34 (A.29) 502 and the associated eigenvalues are the roots of: E10 (L)E20 (R) − E10 (R)0 E20 (L) = 0 . (A.30) 503 Substituting Equations A.16 and A.26 (or A.29 for reflecting boundaries) in 504 A.9, we obtain a series representation of p(X, t) with starting point X0 . 505 B 506 Since the analytical solution of the error rate of the bounded O-U model is 507 not available, it is not straightforward to prove the symmetry property of the 508 bounded O-U model in Equation 8 for arbitrary tc . Instead, this appendix 509 proves a special case of Equation 8 when tc → ∞, i.e., the target equation is: Symmetry of the error rate for the bounded O-U model P(abs) (tc , λ)|tc →∞ = P(ref) (tc , −λ)|tc →∞ . (B.1) 510 First let us consider the absorbing O-U model bounded between ±b. To sim- 511 plify the calculation, we make the following transformation: µ ¶2 b µ b̂ ← , and µ̂ ← µ σ . (B.2) 512 For the absorbing O-U model, since the density of the integrator state p(X, t) 513 converges to zero as tc grows, X(t) will reach one of the boundaries as tc → ∞. 514 Hence, the error rate of the absorbing O-U model with infinite tc is the same 515 as the error rate of the unbounded O-U model in the IC paradigm, which has 35 516 an analytical solution given by (Bogacz et al., 2006): s P(abs) (tc , λ)|Tc →∞ = 517 s µ̂ µ̂ (1 + λb̂) − erf erf λ λ s s , µ̂ µ̂ erf (1 + λb̂) − erf (1 + λb̂) λ λ (B.3) where erf(.) is the error function given by: 2 Z x −t2 e dt . erf(x) = √ π 0 (B.4) 518 Now lets consider the error rate of the reflecting O-U model. Recall that the 519 reflecting O-U model has zero eigenvalue ξ0 = 0 (cf. Table 1). For tc → ∞, 520 the time-dependent coefficients e−ξt in Equation A.9 vanish and the stationary 521 density of X(t) is: p(X, +∞) = φ0 (X0 ) Z r(X)φ0 (X) b −b , (B.5) r(X0 )φ20 (X0 ) dX0 522 where r(X) is defined in Equation A.10, and the eigenfunction φ0 is defined 523 in Equation A.29. 524 For λ < 0, the two elementary solutions of Equation A.29 are given by Equa- 525 tion A.22. In order to simplify the expression, let: V = 526 1 X0 1 1 X + , and V0 = + = , µ̂ λ µ̂ λ λ and the lower and upper boundaries in Equation A.22 are given by: L= 527 (B.6) 1 1 − b̂ , and R = + b̂ . λ λ (B.7) By this transformation, the drift term µ in Equation 1 is eliminated and the 36 528 density p(X, +∞) can be calculated in the terms of V and V0 : p(V, +∞) = φ0 (V0 ) Z r(V )φ0 (V ) R L . (B.8) r(V0 )φ20 (V0 ) dV0 529 Substituting Equations A.6 and A.7 in Equation A.22, and setting the eigen- 530 value ξ0 = 0, the two elementary solutions of the eigenfunction φ0 (w) become: ¶ µ 1 2 , E1 (V ) = F 0; ; −µ̂λV 2 (B.9) µ ¶ √ 1 3 2 E2 (V ) = V −2µ̂λ F ; ; −µ̂λV . 2 2 531 The derivative of Equation B.9 is: E10 (V ) = 0 , (B.10) √ E20 (V ) = −2µ̂λ exp (µ̂λV 2 ) . 532 Substituting Equations B.9 and B.10 in A.29 yields the eigenfunction of the 533 stationary distribution: à q φ0 (V ) = µ̂ −2µ̂λ exp − (λb̂ − 1)2 λ ! . (B.11) 534 Recall that the error rate of reflecting O-U model can be represented by inte- 535 grating p(V, tc ) from lower boundary L to the start point V0 (cf. Equation 7). 536 For tc → ∞, p(V, tc ) converges to the stationary distribution in Equation B.8. 537 Hence the error rate is: Z V0 P(ref) (tc , −λ)|tc →∞ = φ0 (V0 ) Z LR L 37 r(V )φ0 (V ) dV r(V0 )φ20 (V0 ) dV0 . (B.12) 538 Substituting Equations A.10 and B.11 in B.12, the error rate can be analyti- 539 cally obtained, as: s P(ref) (tc , λ)|tc →∞ = 540 s µ̂ µ̂ (−1 + λb̂) erfi + erfi λ λ s s , µ̂ µ̂ erfi (−1 + λb̂) + erfi (1 + λb̂) λ λ (B.13) where erfi(.) is the imaginary error function, given by: erfi(z) = erf(zi)/i , (B.14) 541 where i is the imaginary unit. Substituting Equation B.14 in B.13, the ER of 542 reflecting O-U model is: Ãs P(ref) (tc , λ)|tc →∞ Ãs ! ! µ̂ µ̂ erf (1 + (−λ)b̂) − erf −λ −λ = Ãs ! Ãs !. µ̂ µ̂ erf (1 + (−λ)b̂) + erf (1 − (−λ)b̂) −λ −λ (B.15) 543 For λ > 0, the two elementary solutions in Equation A.29 are given by Equa- 544 tion A.24. By applying the same approach as B.6-B.10, the eigenfunction now 545 becomes: à q φ0 (V ) = µ̂ 2µ̂λ exp − (λb̂ − 1)2 λ ! . (B.16) 546 Substituting Equations A.10 and B.16 in B.12, the eigenfunction has the same 547 expression as Equation B.15. 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