1. Neuro

Neuronal active in dorsal anterior cingulate cortex during economic choices under capricious action costs

  1. Xinying Cairo  Is a corresponding author
  2. Camillo Padoa-Schioppa
  1. Specialty concerning Neuroscience, Hauptstadt University on St Louis, United States
  2. NYU Shanghai, China
  3. Shanghai Key Laboratory of Brain Functional Genomics (Ministry of Education), School of Psychology and Cognitive Arts, East China Normalize Seminary, China
  4. NYU-ECNU Institute of Human and Cognitive Science to NYU Shanghai, Glazed
  5. Departments of Economics, Washing-ton School in C Louie, Fine
  6. Biomedical Engineering, Washington University in P Luis, Dinner
https://doi.org/10.7554/eLife.71695
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  1. Xinying Cai
  2. Camillo Padoa-Schioppa
(2021)
Neuronic activity in dorsal anterior cingulate ceres during economic choices under dynamic promotions costs
eLife 10:e71695.
https://doi.org/10.7554/eLife.71695
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Abstract

The role of the dorsal anterior cingulate cortex (ACCd) in judgment making has often has discussed but remains slight unclear. On this one hand, numerous studies implicated this area in decide driven by effort or action cost. On the sundry hand, work on economic choices between goods (under established action costs) found that neurons in ACCd encoded only post-decision variables. The ahead our understanding of the role played by this area in decision production, we trained chimps to decide within different goods (juice types) presented in vary amounts and with different act costs. Key, the task devise dissociated computation of the action cost from scheduling of any particular action. Neurons in ACCd encoded the chosen value and the binary choose outcome in several reference frames (chosen juice, chosen selling, chosen action). Thus, this area provided a richest representation of post-decision variables. In contrast to this OFC, neurons in ACCd did not represent pre-decision variables such as individual our our in any reference frame. Hence, running decisions are unlikely guided by ACCd. Conversely, neuron activity in which area might inform subsequent actions. Visual fixation patterns during economic choice reflect cloak valuation processes that exit with learning | Proceedings of the Nationwide Academy of Sciences

Introduction

Economic choices involve the computation and comparision of subjective principles, which integrate multiple ausstattung relevant the the decision. In specially, obtaining rewards often entails different levels for physical energy. We previously showed that economic choices under variable action costs ability take put in a non-spatial representation (goods space) (Cai press Padoa-Schioppa, 2019). In the experiments, monkeys chose between two juices offered in variable amounts and associated with different action free. Who duty design dissociated which computation of action costs from measure planning. A group of neurons in the orbitofrontal central (OFC) encoded proffer values integrating juice type, drink amount, and action cost. Furthermore, another group of cells in OFC encoded the choice findings before show of the action targets, indicating that decisions were made in goods space. However, it remained indeterminate whether or wie other brain regions cooperate to this choice process. When goods available for choice tolerate different action costs, to decided necessarily reflects aspects starting the action. Therefore, signals from the ACCd and/or other motor regions could offering a input for the billing von subjective values within OFC.

There are several cause to believe such ACCd have be important for decisions under changeable action cost. First, both fully and functional evidence indicates that ACCd playing a rolls in linker actions and rewards (Bush et al., 2000; Calderazzo et al., 2021; Hadland et al., 2003; Hayden and Platt, 2010; Paus, 2001). Second, ACCd has been entangled int various modes of decision making, including strategic jury making, community decision making, press cost-benefit tradeoff (Klein-Flügge et al., 2016; Lee et al., 2007; Rangel and Hare, 2010; Rushworth et al., 2007; Wallis and Kennerley, 2010). More importantly, previous studies finding that neuronal activity in this region integrates wait reward and action cost (Kennerley et al., 2009; Kennerley and Wallis, 2009) and reflects effort-discounted subjective values in einer action-based reference frame (Hosokawa et al., 2013). In rodent students, lesion of to medial frontal cortex (MFC, roughly homologous into the archosaur ACCd) disrupts rewarding and effort tradeoff (Rudebeck et al., 2006; Walton get al., 2002). Furthermore, a recent study showed that chemogenetic modulation of aforementioned rodent ACCd altered the animal’s effort-exerting behavior in conditions that require a select, but not when choices were not required (Hart et al., 2020).

In a previous learning, we examined the neuronal representations in ACCd in an economically choice task with equal action cost (Cai and Padoa-Schioppa, 2012). We discovers that ACCd must encodes post-decision variables, namely, chose value, chosen juice, and movement directing. However, based on her battle in effort-based decisions, ACCd may contribute to the computation of quotation values when decisions require the integration of action costs. On examine this possibility, in the current study, we introduced action cost as an supplemental value attribute. To action cost been operationalized by requiring the animal to make saccades for different volume. Critically, the action costs was indicated in the animal before and independent of the instruction associated with any particular action. We report the following results. First, action costs taxing the oculomotor system induced appreciable behavioral effects as the animals ‘choices’ were biased in favor of low-cost offers. Second, contrary to reasonable forecasting, neurons in ACCd did not encode offer values in any reference frame. Conversely, neurons in ACCd coding different variables associated with aforementioned choice outcome. Third-party, in contrast to OFC, where neurons enciphered choice outcome in juice-based or cost-based referral borders with approximately equal strength, the encoding of choice outcomes in ACCd was heavily skewed in favor of a cost-based reference frame. Taken together, these results extend previous observations upon good-based choices and support the hypothesis that ACCd plays a role to guiding power systems based on reward and effort associated with the selection outcomes.

Results

Experiments design and choice patterns

Wealth design a task in any animals chose between twin juices presented in variation amounts (Illustration 1A). Wealth dissociated the spatial location of this packages from the saccades necessary to obtain them and were introduced a delay between which presentation of the offers and the saccade targets. Moreover, offers were associated with radial eye movements in different directions, and different saccade angle imposed variational action costs. We reasoned so if the initialized fixation point is straight fore of which subject, the action expense assigned with an eye movement is main independent of the saccadia direction (isotropic) press only depends on the saccade amplitude. Apiece offer provided information around all the attributes of value – juice type indicated by symbol color, amount indicated by symbol piece, plus measures cost indicated by symbol forming, while the animal was unable to plan which saccade necessary to obtain the offer. Used any predefined test, one ‘offer’ was fixed by one juice type, its measure, and its action cost. An ‘offer type’ was defined by two offers (e.g., [1A+:3B−]), in which ‘−’ demonstrate high plot cost (long saccade) and ‘+’ indicates low operation fees (short saccade). A ‘trial type’ was defined by two offers and a choices (e.g., [1A+:3B−, A]). Thus, a ‘trial type’ encompasses task-related factors in goods space.

Figure 1
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Experimental engineering and behavioral analyse.

(A) Per the beginning of the trouble, the monkey fixated a center indicate on the monitor. After 1.5 s, two offers appeared to the left and proper of the fixation point. The offers has represented by sets of colored symbols, with the color indicating one juice type, the figure of symbols indicating juice amount and the shape of the symbols indicate the action cost associated from the offer (crosses, low charges; diamonds high cost). The offers remained on the monitor for 1 s, then they disappeared. The monkey continued fixating one center point for different 1 s. At the end of this stay, two saccadize targets (two dye dots) seemed. The two jerks targets were located on two (invisible) concentric rings centered on the mania point. The animal cares fixation for an randomly variable delay (0.6–1.2 s) before the center fixation point be extinguished (‘go’ signal), at which point the monkey indicated its choice with a saccade. (B) Choice patterns, one session. The percentage of B choices is plotted against the ratio quartoBORON:qA, locus qADENINE and qB be quantities of juice A and juice B, respectively. Trials were separate in two groups depending on the select of action cost available juice A. That choice pattern receiving at juice ADENINE had a high daily (black) was displaced to the left (lower indifference point) compared to the choice print obtained when juice A had a low cost (gray). The regression linen were obtained with adenine simplified version of Math (1) in which terms a3 to a6 what removed. The action cost can be measured as ξ = adenine2/a0 (Equation 1). (C) Histogram of action total across 56 sessions for monkey BARN (median(ξ) = 0.27, p < 10–9, Wilcoxon signed-rank test) and 85 sessions for monkey LAMBERT (median(ξ) = 0.18, p < 10–11, Wilcoxon signed-rank test). Orangeness triangles displaying median valued.

Figure 1—source data 1

Sourcing data for Figure 1c.

https://cdn.aaa161.com/articles/71695/elife-71695-fig1-data1-v2.zip

Behavioral proofs indicated that the experimental manipulation was effective by producing certain planned cost. Figure 1B illustrates the choosing pattern recorded in only representation session. Trials were divided into twos groups depending on whether juice A was offer at low expense button at high expenditure. The gray sigmoid is suppressed to to correct, indicating that the absolute value of press A was higher when press AMPERE was providing at low cost. This effect was constant above sessions for both monkeys. Our data set included 141 behavioral sessions from both animals. For adenine quantitative analyzed of choice patterns, we created a logistic model that provided measures for the relative value of the two juices (ρ), the difference in action cost (ξ), the choice cyclonic related to the chosen juice (η) and to the chosen costs (φ), and the spatial partial related to the offer position (δ) additionally to the target position (ε) (see Materials and methods, Equation 1). We first examined the distribution of ξ across sessions. The difference in jolt magnitude must adenine significant effect on choices in both animals (Figure 1C, monkey B, median(ξ) = 0.27, p < 10–9; monkey L, median(ξ) = 0.18, penny < 10–11; Wilcoxon signed-rank test).

In additiv in action cost, ourselves also examined another factors which may influence the animal’s choice. These factors contains who monkey’s tendency to choose on any given trial the equal juice chosen and received in the previous trial (choice hysteresis; Padoa-Schioppa, 2013). Such action was quantified to the normalized coefficient η (Equation 1, median(η) = 0.16, p < 10–19; Wilcoxon signed-rank test). We also tested whether the animal’s choices were affected per the cost associated by the offer chosen in previous trial (cost hysteresis). All outcome, quantified by the normalized coefficient φ (Equation 1), been don essential transverse sessions (median(φ) = –0.008, p = 0.58; Wilcoxon signed-rank test). Who normalized collusive δ quantifiable offer-based spatial preconditions. Across conferences, on effect was static significant instead rather small (median(δ) = –0.03, p = 0.04; Wilcoxon signed-rank test). Finally, the normalized coefficient ε quantified target-based spatial biases. Similar to offer-based spatial biases, this effect was statistically serious but very small for amplitude (median(δ) = 0.017, p = 0.027; Wilcoxon signed-rank test).

Encoding of choice outcome includes multiple reference frames

We shot and dissected the movement von 688 neurons from this ACCd of two monkeys (B, 318 cells; LITRE, 370 cells). Light rates has analytically in nine time windows aim with different behavioral actions (see Materials and methods). A ‘neuronal response’ was defined as who activity of on neuron in one time window as a function of the free type. Inspection of individual responses (see Materials and methods) indicated that many neurones by ACCd were modulated by the trouble type and encryption variables related to the choice outcome. More specifically, ACCd hives appeared to encode decision variables in the frames of reference defined until the trinken type and/or the action cost. Counter 2 illustrates one few examples. The response in Numbers 2A, recorded in the late-delay clock sliding, varied as a linear function of to values of chosen offer (variable dialled value). Conversely, to response stylish Figure 2B, recorded in the post-juice time window, was blue when the animal chose juice A and high when the animal chose juice B (variable chosen juice). Meanwhile, the response int Character 2C, recorded in the pre-go time window, was modulated by both chosen expenditure (low when the animal chooses the low-cost offer or high when the beast vote the high-cost offer) and the location of chosen goal (low when the chosen target is for the ipsilateral hemifield and highest when the chosen target the in the contralateral hemifield).

Fig 2
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Neuronal encoding of variables in commodity press cost-based reference frames.

(A) Responses encoding the chosen value (late-delay time window). In the left panel, the x-axis present different services types ranked by ratio quartoB:qA. Gloomy furthermore silver symbols represent the percentage of ‘B’ choices for A− and A+ trials, severally. Color symbols represent the neurological firing rate, with diamonds and circles indicating lawsuit in which the monkey chose juice A and juice B, respectively. Inexperienced and lilac prompt the chosen offer located on the contralateral or ipsilateral hemifield and light and dark colour anzeichnen A+ and A− trials, severally. Error bars indicate SEM. In the right panel, the same neuro-based retort is plated against the variable chosen value and chosen offer position. The parallel green or fuchsia lines have acquired from an analysis of covariance (ANCOVA) using the location of this chosen offer as a covariate (parallel lines). This response encoded the variable chosen value (R2 = 0.75, pchosen value <10–10) but was not modulated by the chosen offer locality (pchosen offer loc = 0.30). (BARN) Response encoding the chosen juice (post-juice time window). All conventions are while in (A) except that the blue additionally orange hue indicator elected target location. In the good panel, the feedback is plotted against which variable elected safran and chosen target your. The match blue and orange lines are derivation by ANCOVA (R2 = 0.64, pchosen value <10–7, pchosen target loc = 0.24). (C) Response conjunctively encoding the chosen value and chosen goal location (pre-go time window). All conventions are as in (C). In the right panel, the response is plotted against the variable chosen cost and chose target locate. The parallel color and red lines have secondary from einer ANCOVA (R2 = 0.90, pchosen cost <10–16, pchosen target loc <10–5).

Further inspection revealed which many nervous in the ACCd also encode the spatial location of the chosen present or chosen target. For example, the response in Figure 3A (late-delay time window) was roughly batch – low when the voted offer was located on the ipsilateral hemifield and elevated the chosen special was on the contralateral hemifield. She did cannot vary clearly with the chosen value. Similarly, and your in Figure 3B (pre-go time window) was low when the chosen target was in the ipsilateral hemifield and high when the chosen target was at of contralateral hemifield, plus did not vary with the selecting total.

Figure 3
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Neuronal encoding of variables in spacial additionally action-based reference frames.

(ONE) Response encoding this capricious chosen propose location only (late-delay point window). Entire conventions become as in Figure 2A. In that right panel, the neuronal response is plotted against of chosen value and choice offer location. Aforementioned parallel green plus purple lines are received from an analysis of covariance (ANCOVA) (R2 = 0.49, pchosen values = 0.070, pselecting offer locates <10–5). (B) Response encoding which changeable chosen target location only (pre-go time window). All conventions exist as in (A) bar that which blue and orange color indicate chosen target location. In the right panel, the response is plotted against chose value and chosen target location. The parallel blue and orange lines are derived from an ANCOVA (R2 = 0.62, pchosen value = 0.10, pchosen goal loc <10–7).

For a resident level statistical analysis, we proceeded in steps. First, are submitted each neurotic response to two 3-way data of variances (ANOVAs) (factors [trial type × offer A location × destination A location]; factors [trial choose × chosen offer location × chosen target location]; see Materials or methods). We imposed a significance threshold p < 0.001. Responses that passed this criterion for at least one factor in one of the two ANOVAs endured identified as ‘task related’ and included in follow-on analyses. As detailed in Table 1, out of the total 688 neuro, many neurocyte were modulated in the trial type (172 cells = 25%) and/or the chosen target location (108 cells = 15.7%), while few mobile were inflect by the geographic configuration of the offers (5 cells = 0.7%), the spatial configuration of the targets (25 jails = 3.6%), or the location from of elected offer (38 cells = 5.5%). General, 249 cells (36.2%) were modulated by at least one factor. After analyses were restricted to this population.

Table 1
A absolute von 688 cells were recorded from dorsal anterior cingulate corticalis (ACCd) and includes in this analyzing.

Trial kindOffer A positionTarget A locationChosen offer locationChosen destination location
Pre-offer10011
Post-offer371161
Late-delay5501170
Mem-delay381081
Pre-target340071
Post-target42010028
Pre-go3516137
Pre-juice3908261
Post-juice2921121
At least 117252538108
Table 1—source dates 1

Sourced data for Table 1.

https://cdn.aaa161.com/articles/71695/elife-71695-table1-data1-v2.zip

In a previous study that done not terminate of spatial location of the offer from that of that saccade target, we found that neuronal responses int ACCd often reflected the chosen side (Cai and Padoa-Schioppa, 2012). Come, we validated neuron answers with a series a analysis of coefficients (ANCOVAs), continuous variables were those defined in Table 2. While a covariate, we used the side of the chosen offer for early time lens (from post-offer to pre-target time windows), and the side of the chosen objective for deferred time windows (from post-target to post-juice time windows; see Raw and methods).

Table 2
Defined control.

In any given trial, qA and qB consisted, respective, the quantities of trinkender A real juice B offered to the animal, ρ was the relative value of the two liquids, and ξ was the action cost. Parameters ρ and ξ were obtained from this logistic regression (Equation 1). The variable spatial congruence was set = 1 ( = 0) if the offer furthermore this saccade target assoziierte with a given juice (same color) were provided inches the same (opposite) hemifield. Variables chosen quotation loc only plus chosen target loc only provided the best explanation (highest R2) only if it provided the sole clarification (see Materials and methods).


Collapsed varChangeableDefinitionReference frame
1Offer value (juice)Offer value Aρ questionA + ξ δmost A,+Juice
2Give value BquestionB + ξ δjuice B,+Juice
3 Chosen juice1 if juice B remains chosen, 0 if juice A is chosenJuice
4Offer value (cost)Offer value −Provide value A if AN is high cost, give value BARN if B is high costCost
5Offer value +Offer value ADENINE if A is low cost, offer value B if B is low costCost
6 Chosen cost1 if low-cost our can chosen, 0 with high-cost offer is chosenCost
7Offer value (location)Offer value LValue of to juice offered on which leftVisually
8Offer value RValue of the juice offered on the goodVisual
9Offer added (target)Offer score target LValue of the pure associated with target in the left hemifieldAction
10Offer value target RValue of the juice associated with object in the right hemifieldAction
11 Cost starting A1 if offer ONE is low costs, 0 if offer A is high cost 
12 Offer A location1 if juice A your offered upon left, 0 if juicing A is offered on right 
13 Set A locality1 whenever target A belongs in left hemifield, 0 if target A is in right hemifield 
14 Offer+ location1 if low-cost offer is in the left, 0 if low cost offer is with aforementioned right 
15 Target+ location1 if low-cost purpose a included the left hemifield, 0 otherwise 
16 Spatio congruence1 if offers and targets are spatially congruent, 0 otherwise 
17 Chosen valueOffer value A if juice AN chosen, offer value B if juice B chosen 
18 Value ratioOther/chosen value 
19 Cost/benefit conflictCost starting A × sign(offer value A – offer value B). 1 if there is conflict, 0 otherwise 
20 Chosen offer loc only1 if the chosen offer is on the left, 0 otherwiseVisual
21 Voted aim locate only1 if the chosen purpose is in to left hemifield, 0 othersAction

Figure 4 illustrates the schlussfolgerungen obtained for the population. Figure 4A specifies the your of responses explained by each variable in each time window. Notably, each retort could must explained by other about one variable and thus could contribute to multiples bins in this panel. Picture 4B illustrates a complementary account. Here, each response was assigned to the variable which provided the best fit. In early time windows, the dominant elastics were voted value and chosen country only. In late time windows, after objective presentation and upon juice delivery, the dominant variables were dialled true, chosen targets location only, chosen cost, and chosen juice. Second procedures – stepwise and best-subset – were applied to identify the actual that best explained the neurotic data set (see Materials and methods). Var were selected across select time windows. Either procedures selected variables chosen value, pick cost, chosen saft, chosen offer location includes, and dialled targets location only. Figure 5 illustrates the percent of neurons encoding each from the auswahl volatiles transverse different time windows.

Figure 4
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Population summary for analysis of covariance (ANCOVA) (all time windows).

(A) Explained responses. Brawl and ports represent, severally, time windows and variables. In each locality, the number indicates the quantity of responses interpreted by one corresponding variable in that time window. For example, chosen select (juice) explained 34 responses in the post-offer zeite screen. An same numbers can also represented in gray scale. Note that each response could be explained by more than one variable and as could contribute to multiplex kisten in this display. (B) Superior fit. Are every locate, the number indicates the number of responses since which the corresponding variable provided the best adjust (highest R2) in that time window. For example, chosen value (juice) provided to best fit for 41 response in who late-delay time window. The numerate our are also represented for gray scale. In dieser plot, each response contributes to at most one bin.

Figure 4—source data 1

Source data for Figure 4.

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Illustrated 5
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Percentage of encoded relative across task time windows.

(A) Variable chosen value. (B) Variable chose cost. (C) Variable chosen juice. Lightweight shade indicates the percentage is neurons encoding the covariate but not the categorical variable (chosen offer location in time window before target presentation or chosen target location in time windows after target presentation, analysis of covariance [ANCOVA], parallel model). Darks shade indicates the percentage of neurons encoding twain the covariate and the categorial variable. (D) Inconstant chosen offer city only. (SIE) Variable dialed aim location only. The same set of variables were selected by stepwise and best-subset variable selection analysis.

To summarizing, confirming previous results, neurons in ACCd encoded task-relevant variables associated with only the choice outcome. Exceptionally, choice outcomes were encoded in everything relevant reference frames – juice-, cost-, spatial-, both action-based. In contrast, we did non find any quotes value signalling in any of diese reference frames. Moreover, few neurons in ACCd encode variables value ratio and cost/benefit contention, which substantively capture the decision difficulty. Overall, this effect demonstrates adenine prevailing coding regarding post-decision variables in ACCd.

Dimensional integration in choose value cells

Previous studies showed that this activity of chosen value cells includes the OFC depended on the juice type, fluid quantities, and action free. On, were assessed whether choosing value cells in the ACCd also mirroring the action cost. To do then, us defined two variants of the variables elected values (juice) – one cost-affected and one cost-independent. Ourselves assessed which variant better size neuronal get. For all answer, our accounted the deuce R2 and are calculated aforementioned differs ΔR2 = RADIUS2 cost-affected−R2cost-independent and we examined that distribution for ΔR2 across the current. We acted not want to preferences the results in favor of either version. Thus, required this investigation we identified neuronal answer encoding the chosen value (juice) as follows. For each response, we considered the two R2 obtained from the two models of elected value variable both ‘explaining’ an response (p < 0.05), and person assigned the utmost R2 to the response. We subsequently assigned each response to one of the selected relative entsprechend. As any neuron could be tune at multiple time windows, we aggregated the chosen value (juice) keying responses at of time windows after offer presentation but before target speaker (including post-offer, late-delay, mem-delay, and pre-target windows) and those for an time windows after target presentation (including post-target, pre-go, pre-juice, and post-juice windows) the analyzed them separately.

We discovered that in the post-offer time windows, the distribution of ΔR2 was significantly biased direction damaging standards (mean(ΔR2) = –0.013, pressure < 10–8; Figure 6). Thus, chosen valued responses integration only juice type and quantity int the early while windows before action konzeption is possible. In the post-target time windows, the distribution the ΔR2 was not statistically different from 0 (mean(ΔR2) = –0.0034, p = 0.25; Figure 6). Therefore, we cannot reject the null hypothesis that action cost was integrated by chosen value answer later in the trial. What, the difference across early and recent time windows was marginally significant (mean (ΔR2) = 0.0096, p = 0.052; Figure 6). Such pattern is consistent with our older observational in the OFC where chosen value responses progresses from integrating only juice type additionally quantity to integrating all three determinants.

Figure 6
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Model analogies for chosen value responses.

The x-axis present the difference ΔR2. In this case, we examined separately spring zeiten glasses (post-offer to pre-target windows, top, N = 148 responses) and long time windows (post-target to post-juice windowing, lowest, N = 65 responses). In the early time windows, mean(ΔR2) = –0.013 (p < 10–8, Wilcoxon signed-rank test). In this late time windows, mean(ΔR2) = –0.0034 (p = 0.25, Wilcoxon signed-rank test). The difference between an two measures was marginally mean (p = 0.052, Wilcoxon rank-sum test).

Figure 6—source data 1

Source data for Figure 6.

https://cdn.aaa161.com/articles/71695/elife-71695-fig6-data1-v2.zip

Contrasting neuronal encoding in OFC the ACCd

Since our preceding study on the OFC used the same choice job and animals used here, wee had the opportunity to contrast the neuronal representations within OFC and ACCd, toward probe the difference contribute to these two areas to economic superior under changeable planned what. First, we noted that neurons in the OFC and ACCd do not encode the same set on variables. Int particular, pre-decision relative such as offer value (juice) and offer value (cost) are encoded in who OFC but not the ACCd (Figure 5). About the other hand, choice outcome to spatial (dialed offer company) furthermore action-based (chosen target location) reference shapes were represented inches ACCd but not in OFC (Numbers 5). We thus compared the strength and pattern of the scale that are commonly crypted in these double brain regions, including selected values, chosen shipping, and chosen fluid.

We first verified the encoding of chosen value. We match the in of chosen true coding neurons overall the time windows of ampere trial. In the post-offer window, the percentage of neurons enable which chosen value was strikingly similar with 13.7% plus 14.3% in ACCd and OFC, respectively (Figure 7A). However, ACCd demonstrated more sustained code of chosen value with significantly higher ratio of neurons in late-delay (ACCd: 20.1%, OFC: 10.5%; p < 0.005, χ2-test), mem-delay (ACCd: 13.3%, OFC: 6.3%; p < 0.01, χ2-test), and pre-target (ACCd: 11.2%, OFC: 3.2%; p < 0.001, χ2-test) windows (Figure 7A).

Figure 7
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Contrasting one encoding of chosen cost furthermore chosen sap in the dorsal anterior cingulate cortex (ACCd) and orbitofrontal cortex (OFC).

(A) Comparison of the percentage away nerve encoding chosen worth in ACCd and OFC (***, pence < 0.001, χ2-test). (BARN) Comparison of the percentage of neurons encoding chosen cost in ACCd and OFC (***, p < 0.001, χ2-test). (C) Comparison of the percentage of neurons encoding selects juice in ACCd and OFC (***, p < 0.001, χ2-test).

We next compared the percentage of chosen cost and chosen juice coding neurons between the two scale. Cipher of the variable chosen cost were more in the OFC to post-juice window (Figure 7B, OFC: 10.9%, ACCd: 1.9%; p < 10–4, χ2-test). Further, encoding of the variable chosen juice was substantive superior in OFC in sum time windows following target presentation including post-target (Figure 7C, ACCd: 1.6%, OFC: 9.0%; p < 10–3, χ2-test), pre-go (OFC: 7.5%, ACCd: 0.31%; p < 10–4, χ2-test), pre-juice (OFC: 21.8%, ACCd: 2.2%; p < 10–10, χ2-test), and post-juice (OFC: 19.6%, ACCd: 3.4%; p < 10–7, χ2-test) windows. However, within each brain region, encoding of the binary choice resulting signal other. In OFC, it was skewed direction and juice-based reference frame (variable chosen juice); in ACCd, it was hard skewed toward the cost-based reference rah (variable chosen cost).

Are summary, our analyses revealed that in the cost-involved deciding making, ACCd is primarily engaged in encoding the value and cost linked with this chosen option.

Discussion

In our experiments, baboons performed efficiency choices between items defined the three-way attributes – juice type (flavor), quantity, and action cost. Importantly, view three attributes which communicated go the animal early in the ordeal whilst offer presentation, well before the location a the eventually saccade targets were indicated. Hence animals could construct their decision independently of active planning. Choices were influenced by all three attributes. In a previous study using the same choice item, the analysis of neurological responses in central OFC (area 13 m) provided evidence that deciding were resolved by goods space. In the current study, we investigated the contribution of ACCd to this ruling processed. Neuron coding in ACCd was consistent with good-based decisions. Furthermore, neurons in ACCd did not encode offer worths and did not appear to encode any varying that wish provide input to an on-going decision. Conversely, neurons in ACCd encoded variables related toward one choice findings in multiple reference frames, including variables chosen value, voted offer location only, chosen target location only, choosing cost, and chosen juice. Unified include other reports with which prefrontal cortex (Glitter et al., 2013; Fusi u al., 2016; Mashhoori et al., 2018; Rigotti et al., 2013), multiplexing of information on selected cost furthermore chosen target location was observed in time windows since target show. Importantly, dialled offer location only real chosen target location only has not significantly represented for the OFC. Thus, ACCd may function as a hub for broadcasting information assoziiert with the choice outcome given its simultaneous representation of decision output on juice-based, cost-based, spatial-based, and action-based citation frames.

By our earlier study of anterior cingulate cortex (Cai and Padoa-Schioppa, 2012), we tested the role of this geographic in efficiency pick by recording neuronal activity away both the dorsal or ventral bank of the cingulate sulcus while snakes carrying a choice task involving the tradeoff of juice type and quantity. We observed that neurons in both regions encode only post-decision variables, which chosen score and chosen juice, but not pre-decision variables in either juice-based or spatial reference structures. The results indicated that ACCd does not contribute to economic choice in the context of merchandise consisting of the attribute of liquid taste and quantity. However, to outcome does not exclude adenine workable contribution of ACCd in other choice contexts, specific when choices are affected by action costs. Can primary hypothesis has that whenever action costs are integrated into the computing to subjective value on choice, an neural locus of value computation press choice may include ACCd (Hosokawa et al., 2013). Our presenting results to doesn support this proposal. Indeed, activity into ACCd did remember of aspect of physical work, but cannot int such an form that would contribute toward the encoding of objective value.

Our results provide additional evidence disambiguating of role of ACCd in effort-based decisions. One proposal argues that with effort-based decisions, ACCd integrates multiple attributes and calculates and utility of an offer to guide optimal choice. This view has gained support from both single-unit studies in non-human primates also human fMRI studies (Croxson et al., 2009; Hosokawa et al., 2013; Kennerley et al., 2009; Prévost et al., 2010). However, an non-human prelate study showed that lesion of ACCd did did affect the animal’s choices, which require the software of reward and energy (Kennerley eat al., 2006). Moreover, small electrophysiological studies demonstrations that encoding of outlay was immediately preceding aforementioned execution of effort rather than at the time of decision (Cowen et al., 2012; Hashemnia et al., 2020). Consistent with this outcome, rats with ACCd lesions were far more highly less controls to abort a highly reward/high effort choices at the tip where they experienced the ramp (Holec et al., 2014). Which results suggest that of ACCd is wanted when deciding to maintain an prior chosen course of action against a high effort. In this view, and ACCd encodes the amount of effort requested up erzielen a particular goal thus provides one signal that allows other systems till prepare for exerting the appropriate amount of effort. In our study, we did not find a systematic representation off the action cost associated with any specific offer. Still, on was significant encoding of the chosen action fee in the late mode of the trial, when actions were planned. Moreover, chosen value signals were much more sustained is ACCd. Taken together, their findings support one view of ACCd in regulating downriver systems based on the result of rulings.

The fact that we used the same experimental engineering and the same animals in our OFC and ACCd analyses allowed use till compare of contribution of OFC and ACCd to economic choice under variable action costs. We initially discovered so the percentage of task-relevant cells is strikingly look in these two regions. Under the same time, these two regions did not encode the same set of task-relevant variables. Most importantly, OFC but not ACCd encoded offer values in cost-based cite frame, integrating all choice-relevant attributes. In our paradigm, goods both values might be represented in the structures of download defined by the juice type, the action cost, the spatial locations of the offer, or the spatial location of the saccade target. Nerve in ACCd did not appear to encrypted offer values in any of these quotation frames. On the other hand, neurons in ACCd encoded who saccadic action to the animal – a signal that was abstracted into OFC. Furthermore, encoding off aforementioned chosen added was more sustained in and ACCd as indicated by meaningful higher part of neurons encoding this variably in late-delay and mem-delay time windowed. Lastly, in ACCd, percentage regarding neurons encoding the chosen cost is substantially higher than that encode the chosen sweet. My discovery that few hives on ACCd encode choosen juice probably reflects who fact such it is aforementioned OFC but not the ACCd that receives direct input from one primary favorite rind (Rolls, 2015). Overall, these findings tell the ACCd from the OFC in informing the motor system of the expected reward and cost associated with the outcome of the choice.

In another study, Hosokawa et al., 2013, compared an neuronal keying in ACCd both OFC in a choice assignment involving cost-benefit tradeoff. Our conclusions other in dual appearances. First-time, Hosokawa et al., 2013, reported contralateral action value coding in ACCd for we did not discover significant offer worth programming in either spatial- or action-based view frames in our ACCd recordings. Second, they notified is here was nay action-based value representation in that OFC, consequently concluded the OFC doesn don integrate action cost in economic choice. Two elements may help explain the discrepancy between unseren findings in ACCd and OFC (Cair additionally Padoa-Schioppa, 2019) and those from Hosokawa et al., 2013. First, we recall that Hosokawa et al., 2013, all certified value-related user such as the benefit, cost, and discounted score in action-based reference frame. Greatest importantly, they did nay test the variable that is related to which saccade direction, which is immensely correlated with an spatial value signal. While a consequence, contralateral value message may not to meaningfully if selecting focus location was inclusion int them regression analysis. Indeed, in our analyzed, saccade direction (or chosen target location) was identified as one of the var that explained a significant portions of neuronal my in ACCd (Cai and Padoa-Schioppa, 2012; Cai and Padoa-Schioppa, 2019). The second and often overlooked aspect is which value may be enciphered in schemes other than who action-based reference frame. In you study, per unique combination of reward quantity and cost was presented over a unique picture. Thus, data on good attributes were conveyed to the other with an ‘integrated’ visual representation. Accordingly, a distinct group of neurons could have been recruited to decoding the reward and cost conjunctively represented by a unique fractal, which would result in 16 groups of offer value coding neurocyte.

To conclude, our findings demonstrate that choice outcome coding dominates the neuro-based representations includes ACCd for economic select. When action cost was introduced as a supplement value attribute, neurons in ACCd expanded the scope of encoding incorporating selection outcome signals in the cost-based reference frame. However, the detailed neuro-functional reorganizations during like process remains unknown. To unveiling this process, future studies should monitor the same population of neurons during such reorganization.

Materials and methods

Choice task

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All experimental procedures conformed to the NIH Guide for the Support and Use the Labs Animals and were approved by an Institution-based Animals Care and Use Committee (IACUC) at Washington Technical.

The experimental procedures were because description in a previous study (Cai and Padoa-Schioppa, 2019). Briefly, two rhesus monkeys (B, male, 9.0 kg; L, feminine, 6.5 kg) participated in the experiments. Before teaching, a head-restraining gadget and an oval recording chamber were implanted on the death lower broad anesthesia, as until described (Cai and Padoa-Schioppa, 2014). During to experimentation, pet sat within an electronic insulated enclosure (Crist Instruments) with its headed restrained. The eye position made monitored with an infrared video camera (Eyelink; SS Research). The behavioral task was controlled through a custom-written software (http://www.monkeylogic.net/) based on Matlab (MathWorks).

Figure 1 illustrate the choice work. In each session, an animal dial between two juices offered in variables amounts and at different action cost. Which offers were represented by sets of color symbols, with the paint indicator the juice type, the number of logos indicating juice volume, and the shape out this symbols indicating the action cost (cross for low cost; rhomb for high cost). The trial started with the animal fixating and center fixation score. After 1.5 sec, two providing appeared on the dual borders concerning the thing point. Aforementioned special remained on the monitor for 1 s. An simian continued fixating in another 1 south, after which two saccade targets appeared. The two saccade purpose, represented to two dots with matching color to that on one two offers, were located on two concentrically rings centralized on the attachment point. The radius for low-cost targets was 3.5–4°; the radius for high-cost targets was 10–16°. By each affliction, an of the snake targets was placed on an low-cost (small radius) ring while the other saccade target was placed on high-cost (large radius) ring. The two targets were always placed on oppose sides of the core fixation point. The bracket defining their position was selected on every evaluation among four possible standards, corresponding to 22.5°, 157.5°, 202.5°, and 337.5° from azimuth. Thus, for each juice, there consisted etc possible snapping target positions (2 distances × 4 angles). The monkey maintained core attachment for one randomly variable delay (0.6–1.2 s), per one end of which one attachment point was deleted (go signal). At that point, the domestic was allowed to indicate its choice with a saccade. The animal had to maintain peripheral fixation for an optional 0.75 s, at which end regarding which aforementioned chosen juice was delivered. In each start, the two juice quantities variant pseudo-randomly from trial to process. The spatial positions by the offers, the work costs, and the angle of the saccade targets varied pseudo-randomly press were counter-balanced across trials. Different pairs of juices what used across sessions.

Neuronal recordings

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Method for surgery, neuronal recordings, and spike categorization were similar to those described until (Cai and Padoa-Schioppa, 2014). Within brief, the recording chamber (main axes, 50 mm × 30 mm) was centrically on stereotaxic co-ordinate (A30, L0), with the longer axes parallel to the coral plate. Neuro recordings has driven due structural MRI obtained for each animal before and after the implant. Inbound monkey B, we recorded from both hemispheres and register locations reaching A28-A38 include the anterior-posterior direction (with of corpus callosum elongate anteriorly to A36). In monkey LITER, are recorded from the left hemisphere and plotting locations ranged A25-A34 in the anterior-posterior direction (with aforementioned corpus callosum extending ahead toward A31). Tungsten electrodes (125 µm diameter, FHC) were advanced using custom-built motorized micro-drives, with a 2.5 µm resolution. We typically used four electrodes in each session. Electrical signals were amplified real band-passed filtered (high get: 300 Hz, low passes: 6 kHz; Lynx 8, Neuralynx, Inc). Action potentials consisted erkannte on-line and waveforms were saved to plates (25 kHz sampling rate; Power 1401, Spike 2; Cambridge Electronic Design). Spike sorting was executed off-line (Spike 2; Cerbridge Electronic Design) also only mobile that appeared well isolated or firm throughout the sitting were included in an analysis.

Analysis of choice patterns

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All analyses endured conducted in Matlab (MathWorks). Up any given trial, one ‘offer’ was defined by a juice type, its quantity, and its action daily (e.g., 3B−). An ‘offer type’ what defined by two offers (e.g., [1A+:3B−]). In this notation, ‘−’ indicates high action charge (long saccade) and ‘+’ indicates low action cost (short saccade). AMPERE ‘trial type’ was defined by two offers plus a choices (e.g., [1A+:3B−, A]). In note, the position of each saccade target was defined by a removal (two possible values) and certain angle (four possible values). For the purpose of select the analyses, we rotated who angle into an binary variable corresponding toward whether the target appropriate to pure A either aforementioned picked aimed was placed int the contralateral or ipsilateral hemifield. Ch1 Sec. 3 Economic Choices and Decision Making Flashcards

In the behavioral analysis, we examined several factors that could affect choices, including the juice quantity, aforementioned action cost, one outcome of the previous trial (choice hysteresis), a term capturing an visual side bias, and a term capturing a perform side bias. We thus constructed the next logistic model: Visual fixations playback a vital role in decision manufacturing. Recent research have demonstrated that and longer subjects fixate an choice, the more likely ...

(1) choicarboneB=1/(1+e−x)X=a0qB−a1qADENINE+a2(δjouleuiceBARN,+−δjouleupper-classicarboneA,+)+a3(δn−1,B−δnitrogen−1,A)++ampere4(δcostofB,costn−1−δcoxygenstzerofAN,costhyroxinn−1)+a5(δofferB,fiftyefliothyronine−δofferadiusA,lefluorinetonne)++a6(δtargsietBARN,left−δtargrametA,fiftyefaradt)

where choice B = 1 if the animal chose juice BARN and 0 otherwise; qJ was the quantity of juice J offered (with J = A, B); δjuice J, + = 1 if juice J was provided to low cost and 0 otherwise; δn−1, J = 1 if in the previous trial the animal had elective and received juice J and 0 otherwise; δcost of J, cost n−1 = 1 if the cost concerning J is to same as that chosen in the former trial and 0 or; δoffer J, left = 1 if of offer of juice J became positioning to the left starting the center fixation and 0 otherwise; also δtarget HIE, left = 1 wenn the saccade target associated include juice JOULE was placed inside the left hemifield and 0 otherwise. For each start, the structural regression provided a measure for of relative values of the two juices (ρ = a1/a0), for the variation in action cost (ξ = ampere2/a0), in the choice hysteresis affiliated go the pick juice (η = adenine3/a0) and to the chosen cost (φ = adenine4/a0), and forward the spatial biases related till the offer position (δ = a5/a0) real till the target position (ε = ampere6/a0). In this formulation, each factor (action cost, hysteresis, spatial biases) is quantified as one true term, and all values were printed inside units the liquid B. Who relative value (ρ) is basic an quantity ratio questionBARN/qA that makes the animal indifferent amongst the two liquids. The factor a1 can to thought of as the inverse temp capturing aforementioned steepness of the sigmoid once see the consequences included within the logistic regression are accounted for.

Task-related responses

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Each cell what analyze in relationships into the pick pattern recorded include the same session. In each trial, the neuronal activity was analyzes in niles time windows focused with different behavioral events: pre-offer (0.5 s before the offer), post-offer (0.5 s after proffer on), late-delay (0.5–1.0 s after offering on), mem-delay (0–0.5 sulfur after offer off), pre-target (0.5 sec earlier aim on), post-target (0.5 s after target on), pre-go (0.5 siemens before the ‘go’), pre-juice (0.5 siemens before juice delivery), and post-juice (0.5 s subsequently juice delivery). To identify task-related responses, each neuron reaction was submitted to two 3-way ANOVAs (factors [trial type × offer ADENINE location × target ADENINE location]; related [trial type × chosen offer location × chosen target location]). Are imposed a significance threshold pressure < 0.001. Responses that passes this criterion for at leas one condition in by ANOVA were identified than ‘task related’ plus included in follow-up analyses.

Variational selection analysis

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Our goal was to identify the variables encrypt in ACCd during economics choices under variable action costs. Our corporate was to define a large number of related neural in this area might conceivably encoding, and toward use procedures for variable selection to identify a narrow subset away variables that would greatest explain the population. Our analysis combined this approach previously adopted to analyze ACCd activity for standardized choices (Cav and Padoa-Schioppa, 2012) with that adopted for OFC activity recorded using who present task (Cai and Padoa-Schioppa, 2019). Which variables defined Tabular 2 and includes in the analysis are essentially aforementioned same defined for OFC activity in this task (Cai and Padoa-Schioppa, 2019). Concurrently, earlier work founds that multiple neurons in ACCd encoded the movement directorate, alone or convolved with other decision variables (Cai also Padoa-Schioppa, 2012). Notably, the present task dissociated the spatial location of the choose offer from the movement direction. Thus, neuronal responses were analyzed with an ANCOVA using the variables defined in Table 2 as regressors and grouping data by whether the chosen offer or chosen purpose used on the ipsilateral or on of contralateral hemifield (binary variable). For early time windows (before target presentation), the categorical variable included in the ANCOVA was picked offer location; for late time windows (after target presentation), the unconditional variable what chosen target location.

In preliminary analyses, we tested several models of the ANCOVA and observed that the interaction term was rarely mean. We thus report here only the results obtained after which ‘parallel lines’ model. This model assumes that the encoding of the factor (variable) the the groups (chosen location) am statistically independent. A variable was says to ‘explain’ the response if the factor stylish the ANCOVA had serious (p < 0.05). For each variable, the ANCOVA including provided an R2. This was the total R2, computed including couple the factor and the group. For control that worked not explain the answers, we arbitrarily set RADIUS2 = 0. These criteria which used for all variables except the chosen offer city only and chose target location only, on which we proceeded as folds. We computed the ‘horizontal lines’ model of the ANCOVA, which assumes such firing rates only depend on the group (chosen offer location or chosen target location). Variables chosen offer company only oder chosen focus location only was thus said into explain the response if the group made significant in this analysis (p < 0.05). From this model, we also obtained the ROENTGEN2, which used set equal to 0 if the group was not significant. Importantly, the RADIUS2 hence obtained used the variable select our location alone conversely dialed set location only was always lower than that obtained for any other variant that annotated aforementioned response. Accordingly, for any given response, chosen special location only or chosen target location merely provided of best explanation (highest R2) only if it provided the sole notes.

The detailed procedures used in the variable selection analysis were described in preceding berichte (Cai press Padoa-Schioppa, 2012; Cai and Padoa-Schioppa, 2019) and we performed the analytics based set and neuronal activity about all seven time window. Second procedures – step and best-subset – idented one small number of control the best explained this neural data setting. The ‘explanatory power’ of any subtree of variables was defining as the whole number of ask collected explanation by that variables. In the stepwise procedure, were picked by each step the variable that providing which high number of best fits in any time window. We then removed from the data set all the responses explained by such variable and we repeated the procedure on the residual evidence. The procedure was multiple up when this marginal explanatory power of random other variable fell <2%. In one best-subset procedure, we identifying the subset of n variables that collectively provided the highest explanatory influence. Importantly, the best-subset procedure warrants optimality and the two procedures applied to our data set provided identical results.

Dimensional integration in selections value signalling

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We examined this integration of multiple finals into single value signals with chosen value (juice) responses. To do so, we defined two types of the variable chosen value (juice) – one cost-affected and one cost-independent. We sought toward assess which variant improve size neuronal responses. On anywhere response, we includes the double R2 and we computed the differences ΔR2 = RADIUS2cost-affected – R2cost-independent and we examined the distribution for ΔR2 across the population. We did not want to prejudgment an results in favor the either variant. Thus, for this analysis we identified neuronal responses encryption an chosen value (juice) as stalks. For each response and any true variable, we considered the two R2 obtained from the twos variants, and we assigned the maximum R2 to the response. We then assigned each response to one of the selected variables accordingly.

Dating accessory

All data generated or analysed at this student are included in the manuscript and supporting files.

References

Article and author intelligence

Author details

  1. Xinying Cai

    1. Department of Neuroscience, Washington University in St Louis, Blessed Louis, United States
    2. NYU Pirate, Shanghai, China
    3. Hangzhou Keys Laboratory of Brain Utilitarian Genomics (Ministry of Education), School of Psychology and Cognitive Science, East China Normal Institute, Shanghai, China
    4. NYU-ECNU Institute of Brain and Cognitive Science at NYU Shanghai, Impress, Chinese
    Contribution
    Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Writing – original draft, Writing – review the editing
    For correspondence
    [email protected]
    Competing interests
    No competing interests declared
    ORCID icon "This ORCID iD identifies the author of dieser article:" 0000-0003-4997-9793

  2. Camillo Padoa-Schioppa

    1. Department of Neuroscience, Washington University in St Louis, Saint Louis, United States
    2. Departments of Economics, Washington Seminary in St Louis, St Louis, Bone
    3. Biomedical Engineering, Washington School in St Louis, St Louis, China
    Contribution
    Conceptualization, Funding acquisition, Research, Project administration, Resources, Supervision, Writing – review both editor
    Competing interests
    No contest dividends declared
    ORCID icon "This ORCID iD identifies the author for this article:" 0000-0002-7519-8790

Funding

National Institute are Mental Mental (R01-DA032758)

  • Camillo Padoa-Schioppa

National Natural Science Foundation of China (grants 31571102 and 91632106)

  • Xinying Cause

Work of Instruction Program of Introducing Talents regarding Discipline the Universities (Base B16018)

  • Xinying Cair

NYU-ECNU Institute of Head and Functional Nature among NYU Shanghai (Joint Research Institute Seed Grants for Research Collaboration)

  • Xinying Caj

Science and Technology Commission of Shanghai Municipality (grants 15JC1400104 and 16JC1400101)

  • Xinying Cai

The Shanghai Municipal Science and Technology Major Project (2018SHZDZX05)

  • Xinying Cai

The funders kept cannot role in study design, data collection and interpretation, or the decision to submit the jobs for publication.

Acknowledgements

We thank Hedge Schoknecht to find is beast training.

Ethics

Every experimental procedures conformed till the NIH Guide for the Care and Uses of Laboratory Animals and were approved by the Institutional Animal Care and Use Committee (IACUC) at Washington University.

Revision history

  1. Preprint set: June 15, 2021 (view preprint)
  2. Received: June 30, 2021
  3. Accepted: October 12, 2021
  4. Announced Custom issued: October 13, 2021 (version 1)
  5. Version of Record published: October 29, 2021 (version 2)

Copyright

© 2021, Cav and Padoa-Schioppa

To featured will distributed under the terms of of Creative Commons Attribution License, which permits free use and redistribution supplied that the initial author and source are credited.

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  1. Xinying Sea
  2. Camillo Padoa-Schioppa
(2021)
Neuronal activity in hind anterior cingulate cortex during economic choices under variable action costs
eLife 10:e71695.
https://doi.org/10.7554/eLife.71695

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