Patients with lesions in the posterior cingulate gyrus (PCG), including the retrosplenial cortex (RSC) and posterior cingulate cortex (PCC), cannot navigate in familiar environments, nor draw routes on a 2D map of the familiar environments. This suggests that the topographical knowledge of the environments (i.e., cognitive map) to find the right route to a goal is represented in the PCG, and the patients lack such knowledge. However, theoretical backgrounds in neuronal levels for these symptoms in primates are unclear. Recent behavioral studies suggest that human spatial knowledge is constructed based on a labeled graph that consists of topological connections (edges) between places (nodes), where local metric information, such as distances between nodes (edge weights) and angles between edges (node labels), are incorporated. We hypothesize that the population neural activity in the PCG may represent such knowledge based on a labeled graph to encode routes in both 3D environments and 2D maps. Since no previous data are available to test the hypothesis, we recorded PCG neuronal activity from a monkey during performance of virtual navigation and map drawing-like tasks. The results indicated that most PCG neurons responded differentially to spatial parameters of the environments, including the place, head direction, and reward delivery at specific reward areas. The labeled graph-based analyses of the data suggest that the population activity of the PCG neurons represents the distance traveled, locations, movement direction, and navigation routes in the 3D and 2D virtual environments. These results support the hypothesis and provide a neuronal basis for the labeled graph-based representation of a familiar environment, consistent with PCG functions inferred from the human clinicopathological studies.
Citation: Yang Yu, Tsuyoshi Setogawa, Jumpei Matsumoto, Hiroshi Nishimaru, Hisao Nishijo. Neural basis of topographical disorientation in the primate posterior cingulate gyrus based on a labeled graph[J]. AIMS Neuroscience, 2022, 9(3): 373-394. doi: 10.3934/Neuroscience.2022021
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Abstract
Patients with lesions in the posterior cingulate gyrus (PCG), including the retrosplenial cortex (RSC) and posterior cingulate cortex (PCC), cannot navigate in familiar environments, nor draw routes on a 2D map of the familiar environments. This suggests that the topographical knowledge of the environments (i.e., cognitive map) to find the right route to a goal is represented in the PCG, and the patients lack such knowledge. However, theoretical backgrounds in neuronal levels for these symptoms in primates are unclear. Recent behavioral studies suggest that human spatial knowledge is constructed based on a labeled graph that consists of topological connections (edges) between places (nodes), where local metric information, such as distances between nodes (edge weights) and angles between edges (node labels), are incorporated. We hypothesize that the population neural activity in the PCG may represent such knowledge based on a labeled graph to encode routes in both 3D environments and 2D maps. Since no previous data are available to test the hypothesis, we recorded PCG neuronal activity from a monkey during performance of virtual navigation and map drawing-like tasks. The results indicated that most PCG neurons responded differentially to spatial parameters of the environments, including the place, head direction, and reward delivery at specific reward areas. The labeled graph-based analyses of the data suggest that the population activity of the PCG neurons represents the distance traveled, locations, movement direction, and navigation routes in the 3D and 2D virtual environments. These results support the hypothesis and provide a neuronal basis for the labeled graph-based representation of a familiar environment, consistent with PCG functions inferred from the human clinicopathological studies.
1.
Introduction
In recent decades, importance of fractional order models is well disclosed fact in many fields of engineering and science. Numerous fractional order partial differential equations(FPDEs) have been used by many authors to describe various important biological and physical processes like in the fields of chemistry, biology, mechanics, polymer, economics, biophysics control theory and aerodynamics. In this concern, many researchers have studied various schemes and aspects of PDEs and FPDEs as well, see [1,2,3,4,5,6,7,8,9,10]. However, the great attention has been given very recently to obtaining the solution of fractional models of the physical interest. Keeping in views, the computation complexities involved in fractional order models is very crucial and is the difficulty in solving these fractional models. Some times, the exact analytic solution of each and every FPDE can not be obtained using the traditional schemes and methods. However, there exists some schemes and methods, which have been proved to be efficient in obtaining the approximation to solution of the fractional order problems. Among them, we bring the attention of readers to these methods and schemes [11,12,13,14,15,16,17,18,19,20,21] which are used successfully. These methods and schemes have their own demerits and merits. Some of them provide a very good approximation with convenient way. For example, see the methods and schemes in the articles [22,23,24,25,26,27,28,29,30,31,32,33,34,35,36,37,38,39].
The main aim of this work is to develop a new procedure which is easy with respect to application and more efficient as compare with existing procedures. In this concern, we introduced asymptotic homotopy perturbation method (AHPM) to obtain the solution of nonlinear fractional order models. It is a new version of perturbation techniques. In simulation section, we have testified our proposed procedure by considering the test problems of non linear fractional order Zakharov-Kuznetsov ZK(m,n,r) equations of the form [11,12]
Where a0, a1, a2 are arbitrary constants and m,n,r are non zero integers. If α=1, then equation (1.1) becomes classical Zakharov-Kuznetsov ZK(m,n,r) equation given as:
The ZK equation has been firstly modelized for depicting weakly nonlinear ion-acoustic waves in strongly magnetized lossless plasma [40]. The ZK equation governs the behavior of weakly nonlinear ion-acoustic waves in plasma comprising cold ions and hot isothermal electrons in the presence of a uniform magnetic field [41,42].
The plan of the rest paper is as follows: Section 2 provides theory of the AHPM; Section 3 provides implementation of AHPM. Finally, a brief conclusion and the further work has been listed.
2.
Basic idea of AHPM
Here, we provide that the Caputo type fractional order derivative will be used throughout this paper for the computation of derivative.
Let us consider the nonlinear problem in the form as
T(u(x,y,t))+g(x,y,t)=0,
(2.1)
B(u(x,y,t),∂u(x,y,t)∂t)=0.
(2.2)
Where T(u(x,y,t)) denotes a differential operator which may consists ordinary, partial or space- fractional or time-fractional differential derivative. T(u(x,y,t)) can be expressed for fractional model as follows:
∂αu(x,y,t)∂tα+N(u(x,y,t))+g(x,y,t)=0
(2.3)
subject to the condition
B(u(x,y,t),∂u(x,y,t)∂t)=0,
(2.4)
where the operator ∂α∂tα denotes the Caputo derivative operator, N is non linear operator and B denotes a boundary operator, u(x,y,t) is unknown exact solution of Eq. (2.1), g(x,y,t) denotes known function and x,y and t denote special and temporal variables respectively. Let us construct a homotopy Φ(x,y,t;p):Ω×[0,1]→R which satisfies
∂αΦ(x,y,t;p)∂tα+g(x,y,t)−p[N(Φ(x,y,t;p)]=0,
(2.5)
where p∈[0,1] is said to be an embedding parameter. At this phase of our work it is pertinent that our proposed deformation Eq. (2.5) is an alternate form of the deformation equations as
in HPM, HAM, OHAM proposed by Liao in [43], He in [44] and Marinca in [45] respectively.
Basically, according to homotopy definition, when p=0 and p=1 we have
Φ(x,y,t;p)=u0(x,y,t),ϕ(x,y,t;p)=u(x,y,t).
Obviously, when the embedding parameter p varies from 0 to 1, the defined homotopy ensures the convergence of ϕ(x,y,t;p) to the exact solution u(x,y,t). Consider ϕ(x,y,t;p) in the form
It is obvious that the construction of introduced auxiliary function in Eq. (2.10) is different from the auxiliary functions that are proposed in articles [43,44,45]. Hence the procedure proposed in our paper is different from the procedures proposed by Liao, He, Marinca in aforesaid papers [43,44,45] as well as Optimal Homotopy perturbation method (OHPM) in [46].
Furthermore, when we substitute Eq. (2.9) and Eq. (2.10) in Eq. (2.5) and equate like power of p, the obtained series of simpler linear problems are
We obtain the series solutions by using the integral operator Jα on both sides of the above each simple fractional differential equation. The convergence of the series solution Eq. (2.9) to the exact solution depends upon the auxiliary parameters (functions) Bi(x,y,t,ci). The choice of Bi(x,y,t,ci) is purely on the basis of terms appear in nonlinear part of the Eq. (2.1). The Eq.(2.9) converges to the exact solution of Eq. (2.1) at p=1:
˜u(x,y,t)=u0(x,y,t)+∞∑k=1uk(x,y,t;ci),i=1,2,3,….
(2.12)
Particularly, we can truncate the Eq. (2.12) into finite m-terms to obtain the solution of nonlinear problem. The auxiliary convergence control constants c1,c2,c3,… can be found by solving the system
in Eq. (2.10), we obtain exactly the series problems which are obtained by OHAM after expanding and equating the like power of p in deformation equation. Furthermore, concerning the Optimal Homotopy Asymptotic Method (OHAM) mentioned in this manuscript and presented in [45], that the version of OHAM proposed in 2008 was improved in time and the most recent improvement, which also contains an auxiliary functions, are presented in the papers [47,48]. We also have improved the version of OHAM by introducing a very new auxiliary function in Eq. (2.10). Our method proposed in this paper uses a very new and more general form of auxiliary function
N(ϕ(x,y,t;p))=B1N0+∞∑i=1(i∑m=0Bi+1−mNm)pi
which depends on arbitrary parameters B1,B2,B3,… and is useful for adjusting and controlling the convergence of nonlinear part as well as linear part of the problem with simple way.
3.
Applications
In this portion, we apply AHPM to obtain solution of the following problems to show the accuracy and appropriateness of the new procedure for to solve nonlinear problems.
Problem 3.1.Let us consider FZK(2,2,2) in the form:
In similar way, we can compute the solution of the next simpler linear problems which are difficult to compute by using OHAM procedure. we choose B1=c1,B2=c2,B3=c3,B4=c4 and consider
We obtain number of optimal values of auxiliary constants by using the Eq. (2.13) and choose those optimal values whose sum is in [−1,0). Now, substituting the optimal values of auxiliary constants (from the Table 1) into the Eq. (3.12), we obtain the AHPM solutions for different values of α at k=0.001
Table 1.
The auxiliary control constants for the problem 3.1.
Tables 2 and 3 show the AHPM solution, VIM solution, exact solution and absolute error of AHPM solution. It is obvious from Tables 2 and 3 that AHPM solution results are more accurate to the exact solution results as compare with VIM [11] solution results. The AHPM solution, exact solution and absolute error of AHPM solution are plotted for different values of α, x, y and t in Figures 1 and 2. The curves of AHPM and exact solution are exactly matching as compare with homotopy perturbation transform method (HPTM)[12]. It is obvious from the Tables 2 and 3, Figures 1 and 2, that the AHPM solution of the problem 3.1 is in very good agreement with exact solution.
Table 2.
Solution of the problem 3.1 for various values of α, x, y and t at k=0.001.
We obtain number of optimal values of auxiliary constants by using the Eq. (2.13) and choose those optimal values whose sum is in [−1,0). Now, substituting the optimal values of auxiliary constants (from Table 4) into the Eq. (3.20), we obtain the AHPM solutions for different values of α at k=0.001.
Table 4.
The auxiliary control constants for the problem 3.2.
Tables 5–7 show the AHPM solution, VIM solution, exact solution and absolute error of AHPM solution. The AHPM solution, exact solution and absolute error of AHPM solution are plotted for different values of α, x, y and t in Figures 3 and 4. It is obvious from the Tables 5–7, Figures 3 and 4, that the AHPM solution of the problem 3.2 is in very good agreement with exact solution.
Table 5.
Solution of the problem 3.2 for varios values of α, x, y and t at k=0.001.
In this article, asymptotic homotopy perturbation method (AHPM) is developed to solve non-linear fractional models. It is a different procedure from the procedures of HAM, HPM and OHAM. The two special cases, ZK(2,2,2) and ZK(3,3,3) of fractional Zakharov-Kuznetsov model are considered to illustrate a very simple procedure of the homotopy methods. The numerical results in simulation section of AHPM solutions are more accurate to the exact solutions as comparing with fractional complex transform (FCT) using variational iteration method (VIM). In the field of fractional calculus, it is necessary to introduce various procedures and schemes to compute the solution of non-linear fractional models. In this concern, we expect that this new proposed procedure is a best effort. The best improvement and the application of this new procedures to the solution of advanced non-linear fractional models with computer software codes will be our further consideration.
Acknowledgments
The authors would like to thank anonymous referees for their careful corrections to and valuable comments on the original version of this paper.
Conflict of interest
The authors declare no conflict of interest.
Acknowledgments
This research was supported by the Takeda Science Foundation and a research grant from the University of Toyama.
Conflict of interest
The authors declare no conflict of interest.
Author contributions
HisN and HirN conceived the study and designed the experiment. YY, TS, and HisN performed the experiment. YY, TS, HisN and JM analyzed data and wrote the paper. HisN, HirN, JM, and AB revised the paper. All the authors discussed the results and commented on the manuscript, and read and approved the final manuscript.
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Figure 1. Virtual navigation tasks in monkeys. (A) Setup of the experimental system. (B) Spatial arrangement in and around the mobility area. The monkey was allowed to navigate within the mobility area. In the left route, the monkey visited the reward areas in the order of a series of blue arrows: sequence of reward areas 1→2→3→4→5. In the right route, the monkey visited the reward areas in the order of a series of red arrows: sequence of reward areas 5→6→7→8→9. The common central path segment (location sequence 2-3-4 and 6-7-8) was shared by the left and right routes. (C) Examples of spatial images in the first-person virtual navigation (FP-VN, a), third-person VN (TP-VN, b), and aerial-VN (c) tasks used in this study
Figure 2. An example of a PCG neuron with place-related (A) and heading direction-differential (B) responses. (A) Firing-rate maps in the FP-VN (a), TP-VN (b), and aerial-VN (c) tasks. Left, left route; Right, right route; Composite, composite routes. In the FP-VN task, this neuron showed place-related responses in the right (b) and composite (c) routes. Thin lines on the firing-rate maps indicate the monkey's movement trajectories. Areas marked by thick lines indicate the place fields. Calibration bars on the right indicate the mean firing rates (spikes/s) in the firing-rate maps. (B) Polar plots of neuronal activity in terms of the head direction (1° of angular bin width) in the FP-VN (a), TP-VN (b), and aerial-VN (c) tasks. Each number on each circle indicates firing activity (spikes/s). This neuron showed differential sensitivity to moving directions in the TP-VN task (Rayleigh test, p < 0.0001)
Figure 3. An example of neural activity in the common central path segment with route modulation (a route-modulated zone-related neuron). (A-C) Firing-rate maps in the left (left panel, a) and right (right panel, c) routes in the FP-VN (A), TP-VN (B), and aerial-VN (C) tasks. Thin lines on the firing-rate maps indicate the trajectories of the monkey. The middle panels (b) indicate the mean firing rates in zones 1–3 in the left (dark gray columns) and right (light gray columns) routes. ***, significant main effect of route (p < 0.001)
Figure 4. An example of a path segment-differential neuron. (A-C) Firing-rate maps (a) and mean firing rates in the five segments (b) in the FP-VN (A), TP-VN (B), and aerial-VN (C) tasks. ***, significant difference from the path segments 2/5, 4 and 6 (Tukey test, p < 0.001); #, significant difference from the path segment 4 (Tukey test, p < 0.01)
Figure 5. An example of reward-related responses with route modulation (a route-modulated reward-related neuron). This neuron showed a significant interaction between the route and reward area (p = 0.0021). Each histogram indicates the mean firing rates before and after reward delivery. Zero on the abscissa indicates the time point of reward delivery. Each p-value indicates the results of a one-way ANOVA in each reward area
Figure 6. Decoding of the spatial parameters from the data in the FP-VN task. (A) Ideal location (Aa) and moving direction (Ab) of the monkey and avatar during the tasks based on the labeled theory. (B) Predicted (black dots) and actual ideal (blue and red lines) spatial parameters [distance traveled (a), moving direction (b), X-coordinates (c), and Y-coordinates (d)] in the FP-VN task. AU, arbitrary unit
Figure 7. Decoding of the spatial parameters [distance traveled (a), moving direction (b), X-coordinates (c), and Y-coordinates (d)] from the data in the TP-VN (A) and aerial-VN (B) tasks. AU, arbitrary unit. Other descriptions as for Fig. 6