賈雁兵 楊曉麗? 孫中奎
(1.陜西師范大學(xué),數(shù)學(xué)與信息科學(xué)學(xué)院,西安 710062)(2.西北工業(yè)大學(xué),應(yīng)用數(shù)學(xué)系,西安 710072)
異質(zhì)性和時(shí)滯作用下神經(jīng)元網(wǎng)絡(luò)的共振動(dòng)力學(xué)*
賈雁兵1楊曉麗1?孫中奎2
(1.陜西師范大學(xué),數(shù)學(xué)與信息科學(xué)學(xué)院,西安 710062)(2.西北工業(yè)大學(xué),應(yīng)用數(shù)學(xué)系,西安 710072)
利用參數(shù)互異的Fitzhugh-Nagumo神經(jīng)元構(gòu)建了含耦合時(shí)滯的無標(biāo)度神經(jīng)元網(wǎng)絡(luò)模型,通過數(shù)值模擬的方法,提出研究參數(shù)異質(zhì)性和耦合時(shí)滯影響下神經(jīng)元網(wǎng)絡(luò)的共振動(dòng)力學(xué).結(jié)果發(fā)現(xiàn),當(dāng)耦合項(xiàng)中不含時(shí)滯時(shí),適中的參數(shù)異質(zhì)性能夠使得神經(jīng)元網(wǎng)絡(luò)對(duì)外界弱周期信號(hào)的響應(yīng)達(dá)到最優(yōu),即適中的參數(shù)異質(zhì)性能夠誘導(dǎo)神經(jīng)元網(wǎng)絡(luò)的共振響應(yīng),而且異質(zhì)性誘導(dǎo)共振對(duì)耦合強(qiáng)度具有魯棒性.更重要的是,耦合時(shí)滯對(duì)參數(shù)異質(zhì)性作用下神經(jīng)元網(wǎng)絡(luò)的共振特性也有著顯著性影響.當(dāng)時(shí)滯約為信號(hào)周期的整數(shù)倍時(shí),神經(jīng)元網(wǎng)絡(luò)能夠周期性地出現(xiàn)共振現(xiàn)象,即適當(dāng)?shù)鸟詈蠒r(shí)滯能夠誘導(dǎo)神經(jīng)元網(wǎng)絡(luò)的多重共振,而且這種現(xiàn)象在異質(zhì)性參數(shù)的適當(dāng)范圍內(nèi)都能明顯出現(xiàn).
共振, 異質(zhì)性, 時(shí)滯, 神經(jīng)元網(wǎng)絡(luò), 譜放大因子
2013-07-31 收到第 1 稿,2013-09-23 收到修改稿.
*國家自然科學(xué)基金(11272258,11172342)和中央高?;究蒲袠I(yè)務(wù)費(fèi)專項(xiàng)資金(GK201302001)資助項(xiàng)目
在過去的幾十年里,許多研究者致力于研究噪聲和非線性系統(tǒng)的相互作用.一些經(jīng)典的現(xiàn)象如隨機(jī)共振[1]、相干共振[2]、噪聲誘導(dǎo)或增強(qiáng)同步[3-4]、噪聲誘導(dǎo)相變[5]等見證了噪聲對(duì)非線性系統(tǒng)有序動(dòng)力學(xué)的積極影響.而且,研究對(duì)象也從簡(jiǎn)單的低維系統(tǒng),逐漸擴(kuò)展到耦合系統(tǒng)以至復(fù)雜網(wǎng)絡(luò)系統(tǒng),有興趣的讀者可參考綜述文獻(xiàn)[6-8].
對(duì)于網(wǎng)絡(luò)系統(tǒng),出于數(shù)學(xué)模型的簡(jiǎn)化,大部分研究工作認(rèn)為所有的耦合單元是完全相同的.事實(shí)上,許多真實(shí)體系如生物網(wǎng)絡(luò)和社會(huì)網(wǎng)絡(luò)等,網(wǎng)絡(luò)的個(gè)體之間往往是存在差異的.這種差異性在理論研究中通常通過模型方程中的某些關(guān)鍵參數(shù)不同來體現(xiàn),它被稱為參數(shù)異質(zhì)性[9,10].類似于噪聲,參數(shù)異質(zhì)性對(duì)網(wǎng)絡(luò)系統(tǒng)的混沌、同步、共振等集體動(dòng)力學(xué)具有深刻的影響.例如文獻(xiàn)[11]報(bào)道了最近鄰耦合的單擺振列中,擺長(zhǎng)多樣性能夠?qū)螖[振子由混沌運(yùn)動(dòng)轉(zhuǎn)變?yōu)橹芷谶\(yùn)動(dòng);在神經(jīng)元網(wǎng)絡(luò)中,參數(shù)異質(zhì)性可以將神經(jīng)元由振動(dòng)狀態(tài)轉(zhuǎn)變到可激狀態(tài)[12],也能夠增強(qiáng)網(wǎng)絡(luò)的相干共振[9,13];在 Josephson結(jié)陣列中,Braiman等發(fā)現(xiàn)參數(shù)異質(zhì)性對(duì)同步動(dòng)力學(xué)具有增強(qiáng)作用[14].
特別地,Tessone等于2006年在PRL上報(bào)道了在全局耦合的神經(jīng)元或雙穩(wěn)振子網(wǎng)絡(luò)中,模型參數(shù)的差異性能夠增強(qiáng)網(wǎng)絡(luò)系統(tǒng)對(duì)弱周期信號(hào)的響應(yīng)并引起共振現(xiàn)象[10].相對(duì)于經(jīng)典的噪聲誘導(dǎo)隨機(jī)共振,這種現(xiàn)象被稱為異質(zhì)性誘導(dǎo)共振.隨后,異質(zhì)性誘導(dǎo)共振在轉(zhuǎn)子體系[15]、電子電路[16]、細(xì)胞鈣體系[17]等耦合體系中得到了廣泛的研究.最近,異質(zhì)性與網(wǎng)絡(luò)結(jié)構(gòu)、噪聲、時(shí)滯等因素的相互影響也引起了一些學(xué)者的關(guān)注.例如,在神經(jīng)元網(wǎng)絡(luò)中,Gassel等研究了噪聲與參數(shù)異質(zhì)性誘導(dǎo)共振的關(guān)系,發(fā)現(xiàn)噪聲的出現(xiàn)使得誘導(dǎo)共振出現(xiàn)的參數(shù)異質(zhì)性減?。?8];針對(duì)由雙穩(wěn)振子構(gòu)成的小世界網(wǎng)絡(luò)系統(tǒng),吳丹等發(fā)現(xiàn)隨機(jī)長(zhǎng)程連接能夠增強(qiáng)異質(zhì)性誘導(dǎo)的共振,而時(shí)滯卻對(duì)共振有削弱作用[19].
在神經(jīng)科學(xué)中,耦合的神經(jīng)元之間是存在差異的,這意味著神經(jīng)元網(wǎng)絡(luò)模型中參數(shù)異質(zhì)性普遍存在[9,10].同時(shí),由于動(dòng)作電位沿軸突的有限傳播速度以及突觸間隙的存在,神經(jīng)信息的傳遞存在時(shí)滯效應(yīng)[20].因此,探索參數(shù)異質(zhì)性與耦合時(shí)滯對(duì)神經(jīng)元網(wǎng)絡(luò)的集體動(dòng)力學(xué)的影響有著重要的理論意義和潛在的應(yīng)用價(jià)值.
已有的研究結(jié)果表明,參數(shù)異質(zhì)性能夠誘導(dǎo)全局耦合的神經(jīng)元網(wǎng)絡(luò)[10]和小世界神經(jīng)元網(wǎng)絡(luò)[21]的共振.然而,大腦皮層中一些區(qū)域內(nèi)神經(jīng)網(wǎng)絡(luò)的連接形式具有無標(biāo)度特性[22].那么,參數(shù)異質(zhì)性能否誘導(dǎo)無標(biāo)度神經(jīng)元網(wǎng)絡(luò)的共振?另外,在噪聲的作用下,時(shí)滯能夠誘導(dǎo)由全同神經(jīng)元耦合構(gòu)成的神經(jīng)元網(wǎng)絡(luò)的多重共振響應(yīng)[23-25].那么,在參數(shù)異質(zhì)性作用下,時(shí)滯對(duì)非全同神經(jīng)元構(gòu)成的神經(jīng)元網(wǎng)絡(luò)的共振動(dòng)力學(xué)又有著怎樣的影響呢?通過查閱文獻(xiàn),我們發(fā)現(xiàn)這些問題還沒有得到研究.本文將通過構(gòu)造節(jié)點(diǎn)上是Fitzhugh-Nagumo(FN)神經(jīng)元的無標(biāo)度神經(jīng)元網(wǎng)絡(luò),并利用數(shù)值模擬方法來探討參數(shù)異質(zhì)性和耦合時(shí)滯對(duì)神經(jīng)元網(wǎng)絡(luò)共振動(dòng)力學(xué)的關(guān)鍵影響.
我們將FN神經(jīng)元作為BA無標(biāo)度網(wǎng)絡(luò)的節(jié)點(diǎn),在外界弱周期信號(hào)激勵(lì)下,神經(jīng)元網(wǎng)絡(luò)的動(dòng)力學(xué)方程為
顯然,σ決定不同神經(jīng)元特征參數(shù)的差異性程度:當(dāng)σ=0時(shí)有ai=a0成立,即N個(gè)神經(jīng)元完全相同;當(dāng)σ>0時(shí),不同神經(jīng)元的特征參數(shù)存在差異,而且σ越大,差異性就越顯著.因此,我們也稱σ為異質(zhì)性參數(shù).另外,fsin(Ωt)表示幅值為f、頻率為Ω=2π/T的弱周期信號(hào)(T代表周期信號(hào)的周期).在以下研究中,我們固定ε=0.01,a0=1.12,f=0.05,T=5.0,并假定 ai是服從正態(tài)分布的隨機(jī)變量.
為了定量刻畫系統(tǒng)對(duì)激勵(lì)信號(hào)的響應(yīng),我們引入譜放大因子η[10],其定義為
在這一部分,我們首先討論當(dāng)方程(1)中的耦合項(xiàng)不含時(shí)滯(即τ=0時(shí)),神經(jīng)元網(wǎng)絡(luò)在參數(shù)異質(zhì)性作用下的共振響應(yīng).然后,我們?cè)隈詈享?xiàng)中引入時(shí)滯,并探究時(shí)滯對(duì)神經(jīng)元網(wǎng)絡(luò)共振特性的影響.
當(dāng)耦合項(xiàng)中的時(shí)滯τ為0時(shí),我們來研究異質(zhì)性參數(shù)σ對(duì)神經(jīng)元網(wǎng)絡(luò)集體動(dòng)力學(xué)的顯著性影響.首先通過時(shí)空?qǐng)D來刻畫神經(jīng)元網(wǎng)絡(luò)的時(shí)空動(dòng)力學(xué).當(dāng)g=0.01時(shí),圖1刻畫了不同σ取值下神經(jīng)元網(wǎng)絡(luò)的時(shí)空?qǐng)D.由圖1可以觀察到參數(shù)異質(zhì)性對(duì)耦合神經(jīng)元的放電行為產(chǎn)生了深刻的影響.當(dāng)σ=0時(shí),弱周期信號(hào)激勵(lì)仍不足以使原本處于靜息態(tài)的神經(jīng)元產(chǎn)生放電行為,此時(shí)所有耦合神經(jīng)元都沒有放電,如圖1(a)所示.隨著σ的增大,部分神經(jīng)元的特征參數(shù)減小而產(chǎn)生放電行為,放電神經(jīng)元通過耦合作用帶動(dòng)其它神經(jīng)元也放電,如圖1(b)所示.有趣的是,當(dāng)σ取值大小適中時(shí),時(shí)空?qǐng)D達(dá)到了一個(gè)最規(guī)則狀態(tài),如圖1(c)所示,此時(shí)不同神經(jīng)元的放電動(dòng)力學(xué)基本同步,而且絕大部分神經(jīng)元放電周期約為5.0,這與弱周期信號(hào)的節(jié)律一致.但是,隨著σ的進(jìn)一步增大,大的異質(zhì)性參數(shù)σ導(dǎo)致神經(jīng)元的特征參數(shù)ai的分布比較廣泛,這使得小部分神經(jīng)元的特征參數(shù)遠(yuǎn)大于1.0而不能放電,同時(shí)也使得部分放電神經(jīng)元的放電節(jié)律不能跟隨弱周期信號(hào)的節(jié)律,如圖1(d)與圖1(e)所示,神經(jīng)元網(wǎng)絡(luò)的時(shí)空?qǐng)D反而又變的不規(guī)則了,耦合神經(jīng)元的放電節(jié)律與弱周期信號(hào)節(jié)律的一致性被破壞.以上現(xiàn)象說明當(dāng)異質(zhì)性參數(shù)取值大小適中時(shí),神經(jīng)元網(wǎng)絡(luò)的集體動(dòng)力學(xué)對(duì)弱周期激勵(lì)的響應(yīng)達(dá)到了最優(yōu),即適中的參數(shù)異質(zhì)性能夠誘導(dǎo)神經(jīng)元網(wǎng)絡(luò)的共振行為.
圖1 g=0.01時(shí),不同σ取值下神經(jīng)元網(wǎng)絡(luò)的時(shí)空?qǐng)D:(a)σ =0;(b)σ =0.055;(c)σ =0.07;(d)σ =0.12;(e)σ =0.3Fig.1 The space-time plots of coupled neurons on the networks for different σ:(a)σ =0;(b)σ =0.055;(c)σ =0.07;(d)σ =0.12 and(e)σ =0.3 when g=0.01
進(jìn)一步地,下面借助于譜放大因子η(方程(4))來定量刻畫神經(jīng)元網(wǎng)絡(luò)對(duì)弱周期信號(hào)的響應(yīng).當(dāng)耦合強(qiáng)度為g=0.01時(shí),圖2刻畫了譜放大因子η隨著異質(zhì)性參數(shù)σ變化的曲線(帶實(shí)心方格的曲線).如圖所示,隨著σ的增大,η先增大后減小,并當(dāng)σ≈0.07時(shí)達(dá)到最大值.該現(xiàn)象意味著適中的參數(shù)異質(zhì)性能夠誘導(dǎo)無標(biāo)度神經(jīng)元網(wǎng)絡(luò)的共振響應(yīng),這與圖1中定性分析的結(jié)果相一致.
另外,圖2也給出了耦合強(qiáng)度取其它值(如g=0.005,0.02,0.05,0.08 時(shí)),η隨著σ的演化曲線.不難發(fā)現(xiàn),對(duì)固定的g,隨著σ的增大,η總是先增大再減小,并在適中的σ處達(dá)到最大值.這表明神經(jīng)元網(wǎng)絡(luò)中異質(zhì)性誘導(dǎo)共振對(duì)耦合強(qiáng)度具有魯棒性.同時(shí),我們也發(fā)現(xiàn)隨著耦合強(qiáng)度的增加,曲線的峰值先增大再減小,這說明存在一個(gè)適中的耦合強(qiáng)度,使得神經(jīng)元網(wǎng)絡(luò)的共振特性最佳.
2 對(duì)不同的耦合強(qiáng)度,譜放大因子η隨著異質(zhì)性參數(shù)σ變化的曲線Fig.2 The dependence of η on σ for different coupling strength
在上一節(jié)的研究基礎(chǔ)上,下面在耦合項(xiàng)中引入耦合時(shí)滯,進(jìn)一步研究時(shí)滯對(duì)異質(zhì)性誘導(dǎo)的共振動(dòng)力學(xué)的影響.不失一般性,固定異質(zhì)性參數(shù)σ=0.07,耦合強(qiáng)度g=0.01.首先通過時(shí)空?qǐng)D來刻畫時(shí)滯對(duì)神經(jīng)元網(wǎng)絡(luò)時(shí)空動(dòng)力學(xué)的影響.圖3給出不同時(shí)滯作用下神經(jīng)元網(wǎng)絡(luò)的時(shí)空?qǐng)D.
圖3 異質(zhì)性參數(shù)σ=0.07、耦合強(qiáng)度g=0.01時(shí)不同τ取值下神經(jīng)元網(wǎng)絡(luò)的時(shí)空?qǐng)D:(a)τ=0;(b)τ=2.5;(c)τ=5.0;(d)τ=7.5;(e)τ =10.0;(f)τ=11.0Fig.3 The space- time plots of coupled neurons on the networks for different τ:(a)τ=0;(b)τ=2.5;(c)τ=5.0;(d)τ=7.5;(e)τ=10.0 and(f)τ=11.0 when σ =0.07 and g=0.01
由圖3可知,隨著時(shí)滯的增加,時(shí)空?qǐng)D間歇性地呈現(xiàn)規(guī)則和不規(guī)則狀態(tài):當(dāng)τ=0、5.0和10.0時(shí),分別如圖3(a)、3(c)和3(e)所示,時(shí)空?qǐng)D呈現(xiàn)規(guī)則狀態(tài),此時(shí)耦合神經(jīng)元的放電行為基本達(dá)到同步,而且絕大部分神經(jīng)元的放電周期約為5.0,這與弱周期信號(hào)的節(jié)律一致;而當(dāng)τ=2.5、7.5和11.0時(shí),分別如圖3(b)、3(d)和3(f)所示,時(shí)空?qǐng)D呈現(xiàn)不規(guī)則狀態(tài),而且神經(jīng)元的放電節(jié)律與弱周期信號(hào)的節(jié)律不一致.
圖4 異質(zhì)性參數(shù)σ=0.07、耦合強(qiáng)度g=0.01時(shí)不同時(shí)滯取值下神經(jīng)元網(wǎng)絡(luò)中耦合神經(jīng)元放電峰峰間期的統(tǒng)計(jì)直方圖:(a)τ=0;(b)τ=2.5;(c)τ=5.0;(d)τ =7.5;(e)τ=10.0;(f)τ=11.0ig.4 The histogram of interspike intervals of the networks for different τ:(a)τ=0;(b)τ=2.5;(c)τ=5.0;(d)τ=7.5;(e)τ=10.0 and(f)τ=11.0 when σ =0.07 and g=0.01
為了更形象地刻畫神經(jīng)元的放電節(jié)律,下面進(jìn)一步來描繪網(wǎng)絡(luò)中耦合神經(jīng)元的放電峰峰間期(ISI).圖4(a)、4(c)和 4(e)分別刻畫了當(dāng)τ=0、5.0和10.0時(shí)耦合神經(jīng)元ISI的統(tǒng)計(jì)直方圖.由圖可知,絕大部分神經(jīng)元的峰峰間期集中在5.0附近.圖4(b)、4(d)和 4(f)分別刻畫了τ=2.5、7.5和11.0時(shí)神經(jīng)元網(wǎng)絡(luò)ISI的統(tǒng)計(jì)直方圖,此時(shí)耦合神經(jīng)元的放電峰峰間期分布在較寬的一個(gè)區(qū)域上,神經(jīng)元的放電序列沒有明顯的規(guī)律性.根據(jù)以上分析不難得到:耦合時(shí)滯對(duì)于誘導(dǎo)神經(jīng)元網(wǎng)絡(luò)的時(shí)空有序行為起著積極的作用,而且時(shí)空有序出現(xiàn)在時(shí)滯約等于信號(hào)周期T=5.0的整數(shù)倍時(shí).
接下來,我們也借助于譜放大因子η來定量地刻畫耦合時(shí)滯對(duì)神經(jīng)元網(wǎng)絡(luò)共振動(dòng)力學(xué)的影響.當(dāng)σ=0.07,g=0.01時(shí),圖5描繪了譜放大因子η隨著時(shí)滯τ的變化曲線.由圖可知,隨著τ的增加,曲線呈現(xiàn)出多峰現(xiàn)象,而且峰值分別出現(xiàn)在τ=0、5.0和10.0處.該結(jié)果表明當(dāng)τ約為信號(hào)周期的整數(shù)倍時(shí),神經(jīng)元網(wǎng)絡(luò)對(duì)弱周期信號(hào)的響應(yīng)達(dá)到最佳,這與圖3和圖4的分析結(jié)果一致,即:適當(dāng)?shù)鸟詈蠒r(shí)滯,能夠使得參數(shù)異質(zhì)性作用下的共振響應(yīng)周期性出現(xiàn),我們也稱這種有趣現(xiàn)象為時(shí)滯誘導(dǎo)的多重共振.需要指出的是,越來越多的研究關(guān)注到耦合時(shí)滯對(duì)神經(jīng)元網(wǎng)絡(luò)動(dòng)力學(xué)中的影響,一些重要結(jié)果如時(shí)滯增強(qiáng)的同步[27,28]、時(shí)滯誘導(dǎo)的同步過渡[29,30]、時(shí)滯誘導(dǎo)的多重隨機(jī)共振[23-25]與相干共振[31]、時(shí)滯誘導(dǎo)的時(shí)空有序[32]已被揭曉.不同于已有的研究,這里我們研究的是無噪聲情形、由不同神經(jīng)元構(gòu)成的無標(biāo)度神經(jīng)元網(wǎng)絡(luò)中,耦合時(shí)滯對(duì)參數(shù)異質(zhì)性作用下網(wǎng)絡(luò)系統(tǒng)共振動(dòng)力學(xué)的影響.
圖5 當(dāng)σ=0.07、g=0.01時(shí)譜放大因子η隨著時(shí)滯τ變化的曲線Fig.5 The dependence of η on when σ =0.07 and g=0.01
圖6 耦合強(qiáng)度為g=0.01時(shí),譜放大因子η隨著耦合時(shí)滯τ和異質(zhì)性參數(shù)σ變化的圖像:(a)曲面圖,(b)等高線圖Fig.6 The dependence of η on τ and σ when g=0.01:(a)mesh surface,(b)contour plot
進(jìn)一步數(shù)值計(jì)算的結(jié)果表明,當(dāng)異質(zhì)性參數(shù)取其它值時(shí),時(shí)滯誘導(dǎo)的多重共振也能出現(xiàn).圖6描繪了譜放大因子η隨著耦合時(shí)滯τ和異質(zhì)性參數(shù)σ變化的曲面圖及相應(yīng)的等高線圖.由圖可知,對(duì)于適中的σ,當(dāng)τ的取值在0、5.0和10.0附近時(shí),η達(dá)到峰值.這表明對(duì)于合適的異質(zhì)性參數(shù),當(dāng)時(shí)滯約為信號(hào)周期的整數(shù)倍時(shí),即時(shí)滯和弱周期信號(hào)鎖定時(shí),時(shí)滯誘導(dǎo)的多重共振也能明顯出現(xiàn).
考慮到大腦皮層中一些區(qū)域內(nèi)的神經(jīng)元網(wǎng)絡(luò)具有無標(biāo)度特性,而且參數(shù)異質(zhì)性和時(shí)滯在神經(jīng)系統(tǒng)中普遍存在.鑒于此,本文通過構(gòu)建節(jié)點(diǎn)上動(dòng)力學(xué)不同、含有耦合時(shí)滯的無標(biāo)度神經(jīng)元網(wǎng)絡(luò)模型,首次討論了參數(shù)異質(zhì)性和耦合時(shí)滯作用下無標(biāo)度神經(jīng)元網(wǎng)絡(luò)的共振動(dòng)力學(xué),并得到了一些重要結(jié)果:當(dāng)時(shí)滯為零時(shí),適中的參數(shù)異質(zhì)性能夠誘導(dǎo)無標(biāo)度神經(jīng)元網(wǎng)絡(luò)的共振響應(yīng),而且異質(zhì)性誘導(dǎo)共振對(duì)耦合強(qiáng)度具有魯棒性;當(dāng)時(shí)滯出現(xiàn)時(shí),在一定的異質(zhì)性參數(shù)范圍內(nèi),適當(dāng)?shù)臅r(shí)滯能夠誘導(dǎo)多重共振現(xiàn)象,即當(dāng)時(shí)滯約是信號(hào)周期的整數(shù)倍時(shí),時(shí)滯能夠使得異質(zhì)性作用下的共振響應(yīng)周期性出現(xiàn).本文的結(jié)果豐富了神經(jīng)科學(xué)的理論成果,它對(duì)理解神經(jīng)系統(tǒng)中弱周期信號(hào)的探測(cè)將提供一定的幫助.
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*The project supported by the National Nature Science Foundation of China(11272258,11172342)and the Fundermental Funds Research for the Central Universities(GK201302001)
? Corresponding author E-mail:yangxiaoli@snnu.edu.cn
IMPACT OF DIVERSITY AND DELAYS ON THE RESONANCE DYNAMICS OF NEURONAL NETWORKS*
Jia Yanbing1Yang Xiaoli1?Sun Zhongkui2
(1.College of Mathematics and Information Science,Shaanxi Normal University,Xi’an710062,China)(2.Department of Applied Mathematics,Northwestern Polytechnical University,Xi’an710072,China)
A model of scale-free neuronal networks,which consists of heterogeneous Fitzhugh-Nagumo neurons and time-delayed coupling,was constructed.Then,we explored the nontrivial effects of heterogeneity and time-delayed coupling on the resonance dynamics by numerical simulation in this model.When the delays in the coupling are absent,the result has shown that the response of the neuronal networks to an external subthreshold periodic signal is optimized at an intermediate heterogeneity,namely,an appropriate tuned level of heterogeneity can induce resonance in the neuronal networks.This phenomenon was also confirmed to be robust to the changes of the coupling strength.Most importantly,we find that the delays in the coupling have significant influences on the resonance dynamics.It is revealed that proper delays can induce multiple resonances in the neuronal networks,which appears at each multiple of the oscillation period of the signal.Moreover,the performance of fine tuned delays in inducing multiple resonances can also be clearly observed when the heterogeneity is within an appropriate range.
resonance, diversity, delays, neuronal networks, spectral amplification factor
31 July 2013,
23 September 2013.
10.6052/1672-6553-2013-096
E-mail:yangxiaoli@snnu.edu.cn