Neuroticism and Conscientiousness Respectively Constrain and Facilitate Short-term Plasticity Within the Working Memory Neural Network

Transcription

1 Human Bain Mapping 36: (2015) Neuoticism and Conscientiousness Respectively Constain and Facilitate Shot-tem Plasticity Within the Woking Memoy Neual Netwok Danai Dima, 1,2 Kal J. Fiston, 3 Klaas E. Stephan, 3,4,5 and Sophia Fangou 1 * 1 Depatment of Psychiaty, Icahn School of Medicine at Mount Sinai, New Yok, New Yok 2 Institute of Psychiaty, Psychology and Neuoscience, King s College London, London, United Kingdom 3 Wellcome Tust Cente fo Neuoimaging, Univesity College London, London, United Kingdom 4 Tanslational Neuomodeling Unit, Institute of Biomedical Engineeing, Univesity of Zuich ETH, Zuich, Switzeland 5 Laboatoy fo Social and Neual Systems Reseach, Univesity of Zuich, Zuich, Switzeland Abstact: Individual diffeences in cognitive efficiency, paticulaly in elation to woking memoy (WM), have been associated both with pesonality dimensions that eflect enduing egulaities in bain configuation, and with shot-tem neual plasticity, that eflects task-elated changes in bain connectivity. To elucidate the elationship of these two divegent mechanisms, we tested the hypothesis that pesonality dimensions, which eflect enduing aspects of bain configuation, infom about the neuobiological famewok within which shot-tem, task-elated plasticity, as measued by effective connectivity, can be facilitated o constained. As WM consistently engages the dosolateal pefontal (DLPFC), paietal (PAR), and anteio cingulate cotex (ACC), we specified a WM netwok model with bidiectional, ipsilateal, and contalateal connections between these egions fom a functional magnetic esonance imaging dataset obtained fom 40 healthy adults while pefoming the 3-back WM task. Task-elated effective connectivity changes within this netwok wee estimated using Dynamic Causal Modelling. Pesonality was evaluated along the majo dimensions of Neuoticism, Extavesion, Openness to Expeience, Ageeableness, and Conscientiousness. Only two dimensions wee elevant to task-dependent effective connectivity. Neuoticism and Conscientiousness espectively constained and facilitated neuoplastic esponses within the WM netwok. These esults suggest individual diffeences in cognitive efficiency aise fom the inteplay between enduing and shot-tem plasticity in bain configuation. Hum Bain Mapp 36: , VC 2015 Wiley Peiodicals, Inc. Key wods: dynamic causal modelling; effective connectivity; pesonality; woking memoy Additional Suppoting Infomation may be found in the online vesion of this aticle. *Coespondence to: Sophia Fangou, Icahn Medical Institute, 1425 Madison Ave, New Yok, NY sophia.fangou@mssm.edu Received fo publication 25 Febuay 2015; Revised 30 June 2015; Accepted 6 July DOI: /hbm Published online 20 July 2015 in Wiley Online Libay (wileyonlinelibay.com). INTRODUCTION Woking memoy (WM) is a fundamental cognitive function that involves the maintenance of epesentations of task-elated ules and infomation ove delay peiods in the absence of extenal cues [Bach and Smith, 2008]. Numeous functional magnetic esonance imaging (fmri) studies have confimed that WM tasks ae associated with inceased blood-oxygen-level-dependent (BOLD) signal in the dosolateal pefontal (DLPFC) and paietal cotices (PAR) and the dosal anteio cingulate cotex (ACC) VC 2015 Wiley Peiodicals, Inc.

2 Neual Netwok Plasticity in Behavio and Cognitive Efficiency [Rottschy et al., 2012; Wage and Smith, 2003]. Accumulating evidence povides obust suppot linking pesonality taits of neuoticism (popensity fo negative emotionality) and conscientiousness (goal-diected self-egulation) to individual diffeences in WM task-pefomance and to the maco-configuation of the WM neual cicuity. Individuals with high levels of neuoticism pefom wose on WM tasks [Smith et al., 2011], have lowe DLPFC and ACC volumes [DeYoung et al., 2010; Kapogiannis et al., 2013; Wight et al., 2006, 2007], show educed fonto-paietal anatomical connectivity [Bjønebekk et al., 2013], and attenuated DLPFC activation [Kumai et al., 2004]. Convesely, highe levels of conscientiousness have been linked to bette WM task pefomance [Smith et al., 2011] and geate DLPFC volume [DeYoung et al., 2010; Kapogiannis et al., 2013; Wight et al., 2006, 2007]. At the same time, it is well established that WM is citically dependent on neuonal computations at the micoscale level of synaptic- and neual netwok-level plasticity. Successful maintenance of task-elated epesentations is causally linked to pesistent fiing of pefontal neuons [Fuste, 2001; Goldman-Rakic, 1995; Goto et al., 2010] that ae oganized into dynamic assemblies based on Hebbian pinciples [Fujisawa et al., 2008]. These neual assemblies become sequentially active thoughout the WM task with each eceding assembly passing its epesentational content to futhe assemblies. The integated effect geneates local field potentials [Okun et al., 2010] which ae tightly linked to the fmri-bold signal [Logothetis, 2008]. At pesent thee ae no mechanistic insights as to how aspects of pesonality, that ae associated with the macooganization of the bain, might elate to neuonal computations at the mico-scale level. Dynamic Causal Modelling [DCM; Fiston et al., 2003] is a poweful tool fo the investigation of this elationship in living humans. DCM models the diected influence of one neuonal goup ove anothe (effective connectivity), which povides an indiect measue of netwok-level neuoplasticity [Stephan et al., 2007]. We have aleady demonstated that high WM load inceases inteegional effective connectivity within the WM netwok [Dima et al., 2014]. Hee, we focus on exploing the elationship between DCM-infeed netwok-level plasticity duing WM and pesonality dimensions. We use ou pevious data (Dima et al., 2014) to examine the elationship between paamete estimates of WM modulation and pesonality measues deived fom the five facto model [Costa and McCae, 1992]. Based on the available liteatue, we expected neuoticism and conscientiousness to have opposite effects, espectively negative and positive, on inte-egional connectivity within the WM cicuity. MATERIAL AND METHODS Paticipants Foty healthy paticipants wee included in the study if they (i) had no pesonal lifetime histoy of mental disode TABLE I. Chaacteistics of study sample (n 5 40) Demogaphic data Sex (male:female) 20:20 Age (yeas) 31.5 (10.4) WAIS-R IQ (15.9) NEO-PI-R scoes Ageeableness 113 (21.8) Conscientiousness (32) Extavesion (22.3) Neuoticism 84.8 (34) Openness/Intellect (13.7) Behavioual Pefomance on the n-back task Response time to 3-back (ms) 748 (224) Accuacy fo 3-back (% coect) 72.8 (16.1) Continuous vaiables ae pesented as mean and standad deviation; WAIS-R 5 Wechsle Adult Intelligence Scale-Revised; IQ 5 Intelligence Quotient; ms 5 milliseconds. o substance use as assessed following pesonal inteview using the Stuctued Inteview fo DSM-IV-TR Axis I Disodes, non-patient edition [Fist et al., 2002], (ii) had no histoy of medical disodes o head injuy, and (iii) did not take any pescibed medication. All paticipants wee ighthanded, based on self-epot. Thei intellectual quotient (IQ) was assessed using the Wechsle Adult Intelligence Scale Revised [WAIS-R; Wechsle, 1981]. Pesonality was assessed using the Neuoticism-Extavesion-Openness Pesonality Inventoy-Revised (NEO-PI-R) [Costa and McCae, 1992]. Details of the study sample ae shown in Table I. The study was appoved by the Ethics Committee of the Institute of Psychiaty and the South London and Maudsley National Health Sevice Tust. Witten infomed consent was obtained fom all subjects. Woking Memoy Functional Imaging Task The n-back task was employed in a block design incopoating altenating expeimental and sensoimoto contol conditions. A seies of lettes in yellow font wee displayed on a blue sceen fo 2 s each. Paticipants wee instucted to indicate by a button pess whethe the lette cuently displayed matched the lette fom the peceding n tials. In the sensoimoto contol (0-back), the lette X was the designated taget. In the expeimental conditions (1, 2, 3-back), the taget lette was defined as any lette that was identical to the one pesented in the peceding one, two, o thee tials. Thee wee 18 epochs in all, each lasting 30 s, compising 14 lettes with a atio of taget to non-taget lettes anging fom 2:12 to 4:10 pe epoch. The entie expeiment lasted 9 min and included a total of 49 taget and 203 non-taget stimuli. To avoid any systematic ode effects the conditions wee pseudo-andomised. Pefomance was evaluated in tems of eaction time to taget lettes and accuacy (% coect esponses). The task was 4159

3 Dima et al. explained to paticipants pio to scanning but thee was no taining. Image Acquisition Gadient echo plana magnetic esonance (MR) images wee acquied using a 1.5-Tesla GE Neuo-optimised Signa MR system (Geneal Electic, Milwaukee, WI) fitted with 40 mt/m highspeed gadients, at the Maudsley Hospital, London. Foam padding and a foehead stap wee used to limit head motion. A quadatue bidcage head coil was used fo adio fequency (RF) tansmission and eception. A total of 180 T2*-weighted MR bain volumes depicting blood-oxygenation level-dependent (BOLD) contast wee acquied at each of 36 nea-axial planes paallel to the inte-commissual (AC-PC) plane; epetition time (TR) 5 3,000 ms, echo time (TE) 5 40 ms, slice thickness 5 3 mm, voxel dimensions mm, inteslice gap mm, matix size , flip angle Pio to each acquisition sequence, fou dummy data acquisition scans wee pefomed to allow the scanne to each a steady state in T1 contast. Duing the same session, a high-esolution T1-weighted stuctual image was acquied in the axial plane (invesion ecovey pepaed, spoiled gadient-echo sequence; TR 5 18 ms, TE ms, TI ms, slice thickness mm, voxel dimensions mm, matix size , field of view mm, flip angle 5 208, numbe of excitations 5 1) fo subsequent co-egistation. Functional Image Pocessing Conventional and DCM analyses wee implemented using Statistical Paametic Mapping (SPM8) ( ucl.ac.uk/spm/softwae/spm8/) and the coesponding DCM analysis softwae (DCM8). fmri images wee ealigned, nomalized, and smoothed using an 8 mm fullwidth-half-maximum Gaussian kenel. The smoothed single-paticipant images wee analyzed via multiple egessions using the linea convolution model, with vectos of onset epesenting the expeimental conditions (1, 2, and 3 back) and the 0-back condition as sensoimoto contol. Six movement paametes wee also enteed as nuisance covaiates. Seial coelations wee emoved using an AR(1) model. A high pass filte (128 s) was applied to emove low-fequency noise. Contast images of each memoy load condition vesus baseline wee poduced fo each paticipant. Conventional fmri Analysis Goup-level analyses wee based on andom-effects analyses of the single-paticipant contast images using the summay statistic appoach. Regions showing significant task effect acoss all paticipants wee identified using one-sample t tests against zeo. The statistical theshold was set to P < 0.05 with familywise eo (FWE) coection on a voxelwise basis and minimum cluste size 20 voxels. Fo all analyses, esults ae epoted in Monteal Neuological Institute (MNI) space. Using the above paametes, we identified the functional netwok engaged by the n-back task (Suppoting Infomation Table S1). Dynamic Causal Modeling Selection of volumes of inteest Volumes of inteest (VOIs) wee defined bilateally in the PAR, ACC, and DLPFC based on ou esults and suppoted by pevious studies demonstating obust and consistent involvement of these egions in WM [Rottschy et al., 2012; Wage and Smith, 2003]. The coodinates fo the VOIs wee based on the goup maxima fom the contast of 1-, 2-, and 3-back minus 0-back condition afte conjunction analysis. The co-odinates of the goup maxima within the DLPFC [Left: x 5248, y 5 36, z 5 30; Right: x 5 48, y 5 38, z 5 30], PAR [Left: x 5238, y 5256, z 5 42; Right: x 5 36, y 5252, z 5 44], and ACC [Left: x 5210, y 5 26, z 5 28; Right: x 5 12, y 5 24, z 5 28]. Fo each paticipant, VOIs of 5 mm adius wee defined centeed on paticipant-specific maxima in these egions that wee (i) within 4 mm fom the goup maxima, (ii) within the same anatomical aea, as defined by the PickAtlas toolbox ( and (iii) adjusted using the effect of inteest F-contast. Regional time seies wee summaized with the fist eigenvaiate of all activated (at P < 0.01) voxels within the paticipant-specific VOIs. Model specification As we have peviously descibed [Dima et al., 2014], we used the VOIs defined above in the left and ight PAR, left and ight ACC, and left and ight DLPFC to specify a sixaea DCM in all paticipants fo each expeimental condition (1, 2, and 3 back). Within each hemisphee, we defined bidiectional connections between these egions. Bidiectional connections wee also specified between homologous egions in each hemisphee. Fo each expeimental condition, 18 endogenous connections wee specified in total with the main effect of memoy as the diving input enteing the PAR bilateally. Fo each expeimental condition, this base model was elaboated systematically to poduce altenative vaiants to test how WM load could modulate the 18 connections. Fom this point on, all analyses wee esticted to the high memoy load condition (3-back) vesus sensoimoto contol (0-back) contast because diffeences in cognitive and neual efficiency ae moe appaent at high WM load [Gevins and Smith, 2000]. (Analyses elevant to the 2-back vesus 0-back contast may be found in the Suppoting Infomation Mateial). 4160

4 Neual Netwok Plasticity in Behavio and Cognitive Efficiency (a) Neuoticism was negatively coelated with the WM modulation of the fowad connection fom the R-PAR to the R-DLPFC; (b) Random-effects Bayesian Model Selection identified one model with an exceedance pobability of 96%. Post hoc t-tests on Bayesian Model Aveages of connectivity confimed that WM load significantly modulated the fowad connection Figue 1. fom the R-PAR to the R-DLPFC in all paticipants; (c) Conscientiousness was positively coelated with WM modulation of the fowad connection fom the R-PAR to the R-DLPFC. Abbeviations: L 5 left; R 5 ight; ACC 5 anteio cingulate cotex; DLPFC 5 dosolateal pefontal cotex; PAR 5 paietal cotex; WM 5 woking memoy. Model compaison As we have peviously descibed [Dima et al., 2014], we used andom-effects Bayesian Model Selection (BMS) [Stephan et al., 2009] in DCM8 in ode to compae all plausible models of effective connectivity. Fo the 3-back vs. 0-back condition, 18 models wee compaed in tems of thei exceedance and posteio pobabilities in ode to identify the optimal model. Connectivity paamete estimation We then used andom effects Bayesian Model Aveaging (BMA) to obtain aveage connectivity estimates acoss all models fo each paticipant [Penny et al., 2010] as BMA accommodates uncetainty about models when estimating the consistency and stength of connections. Relationship Between Connectivity Estimates and NEO-PI-R Domains Scoes BMA connectivity estimates of the connections modulated by WM within the optimal DCM model wee examined in tems of thei elationship with all NEO-PI-R domain scoes using a foced egession model into which age [Stanley et al., in pess], sex [Hill et al., 2014], and IQ [Nisbett et al., 2012] wee also consideed as these factos ae known to influence WM. RESULTS NEO-PI-R Domains As shown in Table I, the means of the NEO-PI-R scoes of the study sample wee within thei espective nomative ange. Conbach s alpha coefficients fo the five pesonality dimensions in ou sample wee high (Neuoticism , Extavesion , Ageeableness , Openness/ Intellect , and Conscientiousness ). Identification of Optimal DCM DCM analysis identified a ight sided, fowad WM model with an exceedance pobability of 96%. In this optimal model, the WM load at 3-back significantly inceased effective connectivity in the fowad connection fom the ight PAR to the ight DLPFC (P < 0.002). The stength of the WM modulation of the effective connectivity fom the ight PAR to the ight DLPFC was associated with impoved esponse time ( , P ), without change in pefomance accuacy. Regession Analysis The BMA connectivity estimate of the fowad connection fom the ight PAR to the ight DLPFC was enteed as a dependent vaiable in a foced egession model with age, sex, IQ, and NEO-PI-R scoes of all five pesonality domains as factos. We found that highe neuoticism scoes wee associated with educed WM modulation of the connection fom the ight PAR to the ight DLPFC (b , P ), while the opposite was the case fo conscientiousness (b , P ). Ageeableness (b , P > 0.2), Extavesion (b , P > 0.2), Openness to Expeience (B , P > 0.2), age (b , P > 0.2), sex (b , P > 0.2), and IQ (b , P > 0.2), 4161

5 Dima et al. wee not significant. The oveall model fit was R (Fig. 1). DISCUSSION In healthy individuals, neuoticism, chaacteized by negative affect, has been associated with disadvantageous changes in cognition, bain stuctue, anatomical connectivity [Bjønebekk et al., 2013; DeYoung et al., 2010; Kapogiannis et al., 2013; Kumai et al., 2004; Wight et al., 2006, 2007], and esting state connectivity [Kuschwitz et al., 2015]. Convesely, conscientiousness, chaacteized by adheence to ules and goals, seems to have a potective ole [DeYoung et al., 2010; Fobes et al., 2014; Kapogiannis et al., 2013; Wight et al., 2006, 2007]. The novel finding in this study is the identification of a mechanistic link between pesonality dimensions and cognitive efficiency duing a WM task though changes in ight fontopaietal effective connectivity. Effective connectivity is consideed an index of shot-tem neual-netwok level plasticity [Fiston, 1994; Fiston et al., 2003; Stephan et al., 2010]. This shot-tem plasticity epesents a fundamental mechanism by which the bain altes o contextualizes its connectivity and function in esponse to extenal o intenal cues [Salinas and Sejnowski, 2001]. Two independent lines of evidence suppot the cucial ole of fowad connections fom the paietal to the fontal cotex and thei elationship to WM. Electophysiological studies have found that tasks with high WM load place geate demands on paietal egions, expessed as pesistently high EEG alpha hythm [Gevins and Smith, 2000] and inceased paietal involvement impoves esponse time without affecting accuacy. Similaly, epeated tanscanial magnetic stimulation has been shown to incease paietofontal functional connectivity and impove esponse time in WM tasks [Esslinge et al., 2014]. The paieto-fontal changes in connectivity wee context sensitive (taskelated) and not pesent in esting state data [Esslinge et al., 2014]. Togethe, this evidence suggests that duing WM, high-load conditions tansiently incease paietofontal coupling. We have shown hee that WM modulation of the ight paieto-fontal effective connectivity seems to be influenced by pesonality taits of neuoticism and conscientiousness. Highe neuoticism scoes wee associated with educed WM modulation while highe conscientiousness scoes wee associated with inceased WM modulation. This study pesents a poweful appoach fo identifying mechanistic links between cognitive neuoscience constucts and domains of psychopathology. Deficits in WM of medium to lage effect size ae pesent in most if not all neuopsychiatic disodes [ecently eviewed by Snyde et al., 2015] although thei diagnostic and pognostic significance has been studied most extensively in psychotic disodes, namely schizophenia and bipola disode [Fangou, 2014; Schmidt et al., 2015]. It is also the case that abnomalities in fonto-paietal connectivity in psychotic disodes have also been obustly associated both with WM dysfunction and the clinical symptom expession [Fangou, 2014; Schmidt et al., 2015]. Thee ae seveal methodological consideations in intepeting ou esults. The NEO-PI-R is a self-epot instument. Although thee is much evidence in suppot of its validity and eliability, paticulaly in healthy individuals [Young and Schinka, 2001], self-epot measues inevitably eflect subjective judgement. Futhemoe, ou sample is elatively small fo the study of individual diffeences. Although the association between pesonality dimensions and effective connectivity is obust, confimation of ou findings in a lage, independent sample would be desiable. Last, DCM is not a diect measue of shot-tem neuoplasticity. In fmri studies, neuoplastic esponses can only be infeed fom task-elated changes in BOLD signal. 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