FIGURE 2. The schematic procedure of converting an fMRI dataset
into a format ready for coherence analysis. (a) Player A, B, C,
and D take turns interacting as pairs at the same time. Player D is
outside the scanner. “Feedback” is defined as the time in the second
stage of a trial when the actual choices of the pairs are made, to the
end of the 2-s period in each trial. (b) A single
deconvolutional GLM is created, with 9 stick-like extensions for each of
the two regressors (“target_trial” and “other_trials”). This
method allows a least-square separation of the target trial, from which
the whole brain beta series are independently estimated subsequently for
the 9 TRs following the feedback-initiated event; (c)Concatenating all the 9 TRs from each trial, resulting in 324 volumes
[9 TRs x 6 trials (each run for each pair) x 6 runs] for each pair,
as the periodic data format ready for coherence analysis. (d)The rTPJ is chosen as the seed ROI of the current study for the
interpersonal seed-brain coherence analysis. The ROI beta series are
extracted from the previously concatenated beta series and applied to
the whole brain of the other subject. (e) The regions coupled
with the rTPJ at the target frequency bin, the 36th (1/18 s = 0.0556
Hz), are separately mapped.
Various subgroups, indices and pairing combinations for
analyses
As an alternative method to characterize individual behavioral
characteristics, k-means cluster analyses, a module in JASP with the
Silhouette optimization algorithm (Rousseeuw, 1987), was further
implemented, yielding 3 subgroups:
- reciprocal: taking turns for the bigger reward, n = 49;
- dominant: choosing the bigger reward in 2 consecutive trials,
n = 14;
- submissive: choosing the smaller reward in back-to-back
trials, n = 15.
The descriptive statistics of each subgroup were shown in the
Supplementary Table S1. These subgroup labels were instrumental in the
brain-behavior connectivity analyses.
Two kinds of behavioral indices were devised: individual-wise and
pair/dyad-wise. For the individual-based behavioral indices, each player
interacted with the other three players and had 6 consecutive
interactions with one player in a run, 6 runs in total; out of 6 runs,
there were 35 times of alternation because the first interaction/trial
was not included (35 times/trials x 3 other players = 105 trials). Three
kinds of trial could be derived out of the total 105 trial pairs for
each participant:
a) successful trials: Either Player A or B won the bigger
reward.b) failed trials: Both Player A and B chose the same (bigger or
smaller) reward, ending up both winning zero.c) successful trials with reciprocity: Player A won the bigger
reward, and then B won the bigger reward in the next trial, or vice
versa;d) successful trials without reciprocity: Either Player A or B
won the bigger reward for two trials in a row;
Three individual-based indices were calculated based on these three
kinds of trial pairs:
- the reciprocity index: defined as the number of successful
trials with the winner of the bigger reward alternated (e.g., AB or
BA), divided by the 105 calculable trials;
- the greed index: defined by the number of successful trials
without reciprocity, such as one (e.g., AA or CC) winning consecutive
bigger rewards;
- the total reward: defined as the sum of money each
participant earned in the whole experimental session.
The 2nd set of, or pair-based, indices are:
pairwise success rate: defined as the pairs successfully
winning the reward, regardless of which player won the bigger/smaller
amount;
pairwise reciprocity rate: defined as the pair taking turns
in getting the bigger reward.
Details of pairing for coherence analyses are as follows:
- Within-group Interacting Pairs (WIPs) . The WIPs are pairs
within the same group and are interacting at the same time. For
example, when Player A is interacting with Player B (i.e., A&B) and
Player C is interacting with Player D (i.e., C&D) in Block 1, A&B
and C&D are two interacting pairs. For a given experimental group (3
scanned + 1 behavioral), there are 3 players X 2 pairings X 26 groups
=156 combinations.
- Within-group Non-interacting Pairs (WNPs) . To use the
A&B/C&D pairings above for example, A&C is the non-interacting pair
(for Player D is outside the fMRI, thus with no neural data). For a
given experimental group, there are 3 players X 4 pairings X 26 groups
= 312 combinations.
- Between-group Permuted Pairs (BPPs) . BPPs are pairings of
players from different groups, e.g., A from Group 1 to B/C from Group
2, etc. (3 players X 50 pairings X 26 groups = 3,900 combinations).
Independent two-sample t-tests were carried out for various ROIs on the
three pairings (WIPs, WNPs, and BPPs) to determine whether they were
different distinct populations.
RESULTS
Behavioral results
In the individual level, the reciprocity index, the mean percentage of
successful trials with the winner of the bigger reward alternated was
56.47± 17.8 % (the reciprocity index), the mean percentage of
successful trials without reciprocity was 4.80± 4.18 % (the greed
index), and the average sum of money each participant earned in the
whole experimental session was $290.1± 44.1 NTD.
The results of the pair-based indices indicate\sout averagely 76.4%
(=27.5/ out of 36, or 6 in 6 runs) of the trials the pairs successfully
won the reward (the pairwise success rate), and \soutaveragely 73.8%
(=25.8/35 possible alternations from each dyad) the pair took turns in
getting the bigger reward (pairwise reciprocity rate). The pairwise
success rate was significantly correlated with the pairwise alternation
rate (r = 0.921, p < 0.001), which suggests that the
majority of players soon learned to coordinate with three others,
without being given explicit instructions and being strangers throughout
(i.e., participants were drawn from three different universities). These
also reflect the effectiveness of the study design, which enabled the
participants to follow their natural tendencies.
The homogeneity checks of tri-fMRI data
As a sanity check, the triad-fMRI data from the three sites (NCKU with
GE MR750, NTU with Siemens Prisma, and NCCU with MAGNETOM Skyra), after
the same preprocessing steps and coherence preparations \sout(Figure
2), were compared in the seed-rTPJ betas and the rTPJ-rTPJ coherence
values (Supplementary Figure S1). The results show that these data were
highly overlapping on the major dependent measures. In other words,
despite different raw image values and scanning parameters from each
vendor, the three sites’ fMRI data remained homogenous and comparable,
ready for the interpersonal coherence analyses.
GLM contrasts and PPI analyses
As the first pass of most fMRI studies, the GLM contrast between the
successful and failed reciprocal trials, on the 2nd feedback phase, were
identified and compared. Here, only the stage of revealing the final
result (i.e., Stage 2 Feedback) was chosen as the dyads shared this
period to see whether the collaboration/coordination was (reciprocally)
successful for reward. This feedback stage helped the dyads with
trust-building for the following trials. Two additional reasons are as
follows: a) collinearity and b) the extension and replication of our
previous study (Wang et al., 2023). In our experiment design, the events
of the parameters estimated were fixed. These highly correlated events
(i.e., Stage 1 Decision is always followed by Stage 1 Feedback, then
Stage 2 Decision, and finally Stage Feedback 2 Feedback) might result in
being biased. Consequently, the GLMs were separate for each stage (only
‘Stage 2 feedback’ was adopted in the following analysis), instead of a
single GLM run with all 4 onsets. Second, we adopted Stage 2 Feedback in
our previous analysis for coherence. This study was aimed to extend and
replicate our previous one; thus, the same period of time was employed
for further analysis.
Figure 3a represents the brain regions with significant activations in
the [Successful > Failed trials] contrast. As shown in
the Supplementary Table S2, the contrast [Successful trials
> Failed trials] in the second stage of feedback seven
activated clusters are identified, including the right Precentral Gyrus
(rPG), the right Posterior Cingulate, the right Caudate, the left Middle
Temporal Gyrus, the left Middle Frontal Gyrus, the left Medial Frontal
Gyrus, and the left Inferior Frontal Gyrus. To examine the
brain-behavior relationship, we calculated the correlation between the
three indices (i.e., greed, and total reward) and the [Successful
trials > Failed trials] contrast. No ROI was found
correlated.
In addition to GLM contrast
analysis, the rTPJseed downloaded from Neurosynth.com
was used as the seed region to examine the target regions where their
functional connectivity was significantly stronger in the successful vs.
failed reciprocity trials. In the second feedback phase, the bilateral
Precentral Gyri, bilateral Inferior Frontal Gyrus, bilateral Insula,
right Medial Frontal Gyrus, left Inferior Parietal Lobule, right
Cingulate Gyrus, left Middle Frontal Gyrus, right Postcentral Gyrus and
left Precuneus were identified as more negatively functionally connected
with rTPJ in the successful trials than in the failed trials (see the
Supplementary Table S3). The target regions were also mapped in Figure
3b. To examine the brain-behavior relationship, we calculated the
correlation between the three indices (i.e., reciprocity, greed, and
total reward) and the [Successful trials > Failed
trials] contrast. Only the right Sub-gyral was found correlated with
reciprocity (see the supplementary Table S4).
We further employed connectivity-based correlations with individual
total reward. To investigate the underlying network differences with
their individual total reward in the successful vs. in the failed
trials, the rTPJseed connectivity was further examined.
The results showed negative functional connectivity in the right
postcentral gyrus, right Angular Gyrus, and right Inferior Parietal
Lobule when the participants successfully reciprocated with their
pairmates (see Figure 3c and the Supplementary Table S5).These results
indicated that the lower the functional connectivities between the hub
areas of different sub-networks, such as attention/memory and self-other
awareness areas between rTPJ-rIPL and -rAG, the higher reproductive
success.