He loves me, he loves me not:
Testosterone administration does not enhance emotional responses but rather suppresses
positive affective facial reactivity to social feedback by potential mates: an EMG study
Suzanne Hoogeveen
ABSTRACT The steroid hormone testosterone is known to play a causal role in various aspects of
social-affective behavior, and particularly regarding mating behavior. Whereas the contribution of testosterone in motivating intrasexual competition and intersexual approach tendencies has been established, its involvement in the passive affective experience of these mating-relevant behaviors, including evaluation by a potential romantic partner, has never been addressed experimentally. In this placebo-controlled, within-subjects study, the effects of testosterone administration on the affective responses to social feedback (acceptance vs rejection) of a potential mate were examined in 20 naturally-cycling single women, by means of subjective ratings and facial electromyography. Testosterone was expected to enhance positive affect in response to acceptance and to attenuate negative affect in response to rejection. Results show that, contrary to expectations, testosterone had no effect on subjective happiness ratings or corrugator supercilii activity for either acceptance or rejection feedback. However, testosterone was found to suppress positive affect elicited by acceptance of attractive males as reflected in attenuated zygomaticus major activity. These findings suggest that testosterone might dampen passive positive affective responsiveness to social evaluation, perhaps by raising confidence, thereby promoting instrumental approach behavior.Keywords: testosterone; social evaluation; facial EMG; mating behavior; affective processing
INTRODUCTION
Common conceptions about the steroid hormone testosterone (T) mainly appertain to typical male characteristics such as physical strength, aggression and dominance; prompting the sentence “Does testosterone make you…” in the Google search bar provides illustrating auto-completions: “…stronger / mean / grow hair”. Also within the scientific literature regarding social behavior, the focus appears to be primarily on effects related to social threat and dominance, for which the causal role of T has been well established (Bos et al., 2012; Eisenegger et al., 2011; van Honk et al., 2014). Another prominent line of research emphasizes the role of T in primary reproductive and sexual behavior (e.g., Alexander & Sherwin, 1993; Bancroft et al., 1983; Tuiten et al., 2000). What almost all studies regarding T effects on social behavior have in common, though, is the focus on the motivational and instrumental behavioral effects of T (e.g., aggressive decisions in economic settings, behavioral approach of members of the opposite sex), with minimal attention for T effects on passive affective responses to these kinds of behaviors provoked by T (e.g., experience of being treated unfairly in bargains, experience of being approached by a member of the opposite sex). The present study aims to fill this gap by elucidating the causal effects of T on the experience of social evaluation in a romantic partner selection setting, as this refers to an interpersonal process that is both highly relevant for everyday human social behavior and evolutionary reproduction, yet largely disregarded in the T literature.
Competition and courtship behaviors
An influential overarching theory about the role of T in social behavior is the challenge hypothesis (Wingfield et al., 1990), which postulates that T is related to the perception of social challenges and respective behavior, aimed at enhancing social status, ultimately boosting chances of reproduction. The hypothesis was originally formulated to account for the patterns of aggression displayed in male birds and formed the basis for research into dominance behavior associated with T activity in humans (Archer, 2006; Bos et al., 2012; Eisenegger et al., 2011; van Honk et al., 2001; van Wingen et al., 2009). Wingfield et al. (1990) demonstrated how temporal and interspecies variation in T among male birds is directly related to variations in mating and breeding strategies, a link that has also been established for female birds (Ketterson et al., 2005). Notably, the influence of T on mating behavior appears to be more extensive in monogamous species, for whom the stakes of finding a suitable partner are higher compared to polygamous species.
Based on this line of research, it has been postulated that T promotes investment in ‘mating effort’, which includes the tendency for intrasexual competition and aggression, as well as intersexual displays of courtship (e.g., demonstration of strength, ability and beauty, aimed at raising attractiveness in the eyes of potential mates). In rodents and primates (Schulz & Sisk, 2006; Wallen, 2001), as well as humans, T levels have indeed been related to mating success (Peters et al., 2008), sexual motivation (Bancroft et al., 1983; Forbes & Dahl, 2010) and biases towards mating-relevant information (Bobst et al., 2014; van Wingen et al., 2008b; Wang et al., 2014).
The testosterone-mating relationship in humans
With regard to human mating effort, the link between T and romantic partner selection behavior has been addressed in several ways. A proposed model by Roney and Gettler (2015) captures the role of T in intimate relationship dynamics, incorporating both T reactivity (i.e., acute changes in endogenous T levels in response to internal or external cues) effects on short term mate approach (relationship initiation) and baseline T effects on relationship long term commitment (relationship continuation). The former constitutes T as involved in promoting behavior directed at mate pursuit, including competitive drive for high social status and more direct social approach of potential mates. In this regard, several studies have shown a rapid increase in endogenous T in response to short encounters with (attractive) female strangers in men, which is absent in the case of interactions with other males (Roney, 2009; Roney & Gettler, 2015; Slatcher et al., 2011; van der Meij et al., 2008). Likewise, females watching video clips of an attractive man courting a woman, also induced an increase in T levels, which is interpreted as T release facilitating courtship interactions with potential partners (Lopez et al., 2009). Notably, even the passive viewing of facial stimuli of members of the opposite sex sufficed to significantly raise endogenous T levels compared to same-sex faces (Zilioli et al., 2014). Second, long term monogamous committed relationships are associated with lower baseline T in both men and women (Edelstein et al., 2014; Gray et al., 2004; Roney & Gettler, 2015) and T levels actually drop after marriage (Mazur & Michalek, 1998), when mating efforts are obviously less relevant. Third, more directly, a bidirectional relation between T and sexual activity has been established. That is, pre-intercourse hormone levels are characterized a rise in endogenous T, reflecting an anticipatory effect, assumed to facilitate sexual activity (van Anders et al., 2007). Furthermore, T administration has been shown to cause increased physiological arousal (i.e., genital responsiveness) and subjective feelings of arousal and lust1 (Tuiten et al., 2000).
Affective consequences and social evaluation
1With respect to the subjective feelings of arousal and lust, it has been suggested that this subjective effect occurred only
secondary, as a consequence of the interpretation of the physiological effect, as this subjective effect remained absent in a different T administration design (Tuiten et al., 2002).
In general, the effect of T is marked by behavioral disinhibition and might from an evolutionary perspective be summarized as promoting mating effort by enhancing approach behavior (Aarts & van Honk, 2009; Terburg & van Honk, 2013; van Honk et al., 2004), status-pursuit (Eisenegger et al., 2011; Mazur & Booth, 1998; Mehta & Josephs, 2006), courtship displays (Guéguen, 2009; Ronay & Hippel, 2010; Roney et al., 2003; van der Meij et al., 2012) and risk-taking (Campbell et al., 2010; Coates & Herbert, 2008; Mehta & Beer, 2009). Based on the literature, it seems solidified that a rise in T levels prepares an animal or human individual for mating behavior. However, the question of how T subsequently affects the passive emotional experience of the outcomes of these mating-relevant social approach behaviors (e.g., displays of dominance or courtship) remains remarkably unanswered. In other words, whereas many studies have emphasized how T contributes to social approach tendencies, very little research exists into the influence of T on the passive affective responses with regard to social evaluation, and none specifically in a romantic context. This seems remarkable, given the indisputable contribution of T in motivating sex, mating and reproduction, as well as the high relevance of social evaluation in general and in a dating context specifically (consider the ubiquity of dating websites and apps, e.g. Tinder, in which social evaluation by potential mates is the core component, not to mention in actual physical interactions). The current study aims to fill this gap by taking the T-mating relationship one step further, by means of examining the emotional component of the experience of being socially evaluated, i.e., individuals’ affective responses to acceptance or rejection by a potential romantic partner.
To our knowledge, the relation between T and social acceptance vs rejection has only been addressed in a Cyberball paradigm (involving inclusion or exclusion by other individuals in a ball tossing game), which is arguably more competitive and directly related to social status, and less to mating. These studies presented mixed results, with only one reporting a significant drop in endogenous T upon social rejection in males and females, and a rise upon acceptance only in males (Seidel et al., 2013), while the other two studies found no significant change in T levels in response to rejection or acceptance (Geniole et al., 2011; Peterson & Harmon-Jones, 2012). Moreover, the causal role of T as addressed in exogenous administration designs in the affective and physiological response to social evaluation has never been experimentally addressed.
We start from the conception that T is causally involved in primarily enhancing approach and mating behaviors – “putting oneself out there” -, thereby risking social rejection. Secondarily, as a result of this focus, T might influence the emotional response to the outcomes of these efforts. That is, in line with the challenge hypothesis, T is expected to enhance reward-sensitivity towards positive mating interest (i.e., positive emotions upon acceptance by a potential partner), as well as to attenuate negative affect upon rejection, contributing to increasing the likelihood of prospective attempts. These predictions would present new, converging evidence of T increasing sensitivity for mating stimuli specifically – faces of the opposite sex (van Wingen et al., 2008b; Wang et al., 2014) and erotic stimuli (Alexander & Sherwin, 1993; Tuiten et al., 2000) – as well as for rewarding stimuli generally (Hermans et al., 2010; Lombardo et al., 2012; Op de Macks et al., 2011; van Honk et al., 2004; Wu et al., 2016). This general reward-punishment sensitivity account of T holds that T makes individuals more sensitive to rewards and less to punishment, be it monetary or social. This, in turn, is suggested to result from the fear/anxiety- and stress-reducing properties of T (Boissy & Bouissou, 1994; Hermans et al., 2007; Hermans et al., 2006a; van Honk et al., 2005; Viau, 2002). Based on this broad research, in the current study T is hypothesized to enhance the positive emotions in response to social acceptance, while providing a buffer against the negative emotions elicited by social rejection, and thus dampening the negative affect elicited by rejection. Both these responses would by instrumental in encouraging future dating behavior.
The present study
A newly-developed dating experiment was used, comparable to apps like Tinder, in which female participants view photos of males, evaluate their attractiveness and subsequently receive positive (acceptance) or negative (rejection) feedback about the interest of the men. Based on the documented dissociation in T administration
effects on consciously reported and nonconsciously expressed affect (e.g., Schutter & van Honk, 2004; Tuiten et al., 2002; van Honk et al., 2005; van Honk et al., 2004), we included a subjective happiness rating as well as a measure for automatic, nonconscious affect by means of electromyography (EMG) over the zygomatic major and corrugator supercilii facial muscles. Zygomaticus activity (cheek muscle) typically reflects positive affect (happiness), while corrugator activity (eyebrow muscle) is indicative of negative affect such as sadness, fear or anger (Dimberg et al., 2002). Although facial EMG is predominantly used for facial mimicry effects, it has also successfully been applied to measure affective response on textual (performance) feedback (Gentsch et al., 2015; Smith, 1989). Especially relevant with respect to the current study, Wiggert et al. (2015) successfully demonstrated facial EMG activity reflecting the affective response to verbal displays of social acceptance vs rejection. Similarly, in order to purely capture the affective response to the social feedback, rather than to the male face, we measured EMG over the textual feedback, in which solely the decision of the males (i.e., acceptance or rejection) was presented.
First, some general task-effects were expected. As attractiveness has proven to veraciously signal reproductive fitness (Gangestad et al., 1994; Pflüger et al., 2012) and to be a crucial determinant in intersexual selection (Currie & Little, 2009; Riggio et al., 1991), this factor is included in the design. It was thus predicted that males judged to be highly attractive would elicit positive affect (high happiness ratings, zygomaticus activation and corrugator deactivation) when supposedly accepting the female participant and negative affect (low happiness ratings, zygomaticus deactivation and corrugator activation) when rejecting the participant. For males that were judged to be unattractive, the pattern would be reversed; acceptance by unattractive males is expected to induce negative affect, and rejection positive affect.
Second, with regard to T-effects, we hypothesized that T administration will increase the experienced positive affect upon social acceptance by a potential partner – i.e., higher happiness ratings, increased zygomaticus and decreased corrugator activity – compared to placebo. In addition, T administration is expected to attenuate the negative affective response to social rejection – i.e., higher happiness rating (lower ‘unhappiness’ rating), increased zygomaticus and decreased corrugator activity – compared to placebo. Importantly, it is expected that the influence of T on the responses to social evaluation will mostly occur for male faces of high attractiveness, as these are most relevant with respect to mating behavior; no effects of T are expected for male faces of low attractiveness.
METHODS
SubjectsTwenty healthy Caucasian females (mean age = 21.7, SD = 3.7) participated in the experiment for which they received a financial remuneration. Participants were students at the University of Cape Town and were recruited via an advertisement that was disseminated by email. Exclusion criteria were smoking, use of hormonal contraceptives and history of medical, psychiatric or endocrine illness. All women were single and heterosexual. The study was approved by the Human Research Ethics Committee (HREC) of the University of Cape Town, in accordance with the latest declaration of Helsinki.
Design and procedure
Participants were tested in a randomized, double-blind, placebo-controlled, within-subjects design. They participated on two session days within the follicular phase of their menstrual cycle (i.e., testing was done during day 1-12 from the start of the menstruation), when endogenous sex steroid levels and inter-individual variance are typically low (e.g., Armbrustera et al., 2017). An interval of exactly four hours between the sublingual administration of the drug or placebo and the experimental task was used, following Tuiten et al. (2000). T samples consisted of 0.5mg of T, 5mg of the carrier cyclodextirine, 5mg of ethanol and 5ml of water. Placebo samples
were identical except for the omission of the 0.5mg of T. The 4h interval and sublingual administration method have yielded substantial behavioral, neural and physiological effects for T in previous studies (Hermans et al., 2006a; Hermans et al., 2006b; Montoya et al., 2013; van Honk et al., 2005; van Honk et al., 2004; van Honk et al., 2001). Pre- and post-administration saliva samples were taken on both session days in order to measure baseline T-levels. Additionally, mood was measured on both days in the morning and afternoon by means of the Positive Affect Negative Affect Scale (PANAS; Watson et al., 1988). At the end of the second session, the participants had to indicate on which session they thought they received T and on which placebo, which served as a manipulation check. Moreover, they had to describe the purpose of the study as well as the effects they believed T had on their behavior. Lastly, they were asked if “there were any aspects of the dating task which they had doubts about or if they felt all the information given was real”2 and if they had some doubts, they quantified their disbelief on a 1-10
scale, with 1 = very unbelievable and 10 = very believable. After completion of all tasks and manipulation checks, participants were debriefed on the purpose of the study, as well as on the fictional nature of the dating profile setup, and received the financial remuneration (see figure 1A for study overview).
Figure 1. (A) Experimental procedure. Participants signed the informed consent document, provided the first saliva sample
and completed a checklist and mood assessment (PANAS) on the computer, followed by the sublingual intake of the drug. Blind to both participant and experimenter, they received T on one session and placebo on the other, counterbalanced across participants. After a 3.5h break they returned to the lab and again provided a saliva sample and completed the PANAS questionnaire. EMG electrodes were applied and the social feedback task was explained. The 40 experimental trials were preceded by 2 practice trials and followed by 2 questions assessing a) motivation to meet the men in the task and b) extent of experienced feelings of rejection. In the second session, 3 manipulation check questions were posed and the participants were debriefed about the fictional nature of the social evaluation. (B) Experimental task. The social feedback task (dating task) consisted of 40 trials, 20 of which were acceptance (“interested”) trials and 20 rejection (“not interested”) trials, presented in a pseudorandomized order. A trial was built up from the following components: a fixation cross (3000ms), the presentation of the male face (stimulus; 3000ms), a rating screen in which the participant rated the stimulus on attractiveness (on a 7-point scale), a fixation cross (3500ms), textual presentation of the feedback (3000ms) and a rating screen in which the participant rated her happiness with the outcome (on a 7-point scale). EMG was recorded over the stimulus presentation and the feedback presentation (blue-bordered screens in figure). (C) Electrode placement. Two ground electrodes were placed midline on the forehead as reference. Zygomaticus major activity was measured with two electrodes on the left cheek, in the middle of the line from the bone next to the ear to the corner of the mouth. Corrugator supercilii activity was measured from one electrode right above and one next to the inner corner of the left eyebrow.
Figure 2. Examples of the male facial stimuli used in the social feedback task.
Social feedback task
The social feedback task is a newly developed experiment that was validated in a pilot study (n=20). The task was designed to measure subjective experience of social acceptance and rejection by a potential romantic partner. Prior to the experiment, participants filled out a (fictional) dating profile with photo and were told that this was evaluated by a pool of international men. The task consisted of 40 trials, half of which were acceptance trials (“interested”) and half rejection trials (“not interested”), which were preceded by 2 practice trials. In each trial, the participant would see a photo of a specific man, rate his attractiveness, see his evaluation of herself (i.e., social feedback; “interested” or “not interested”) and indicate how happy she is with the outcome (i.e., subjective experience), respectively. Unknowingly to the participant, the photos were randomly distributed over the acceptance (50%) and rejection (50%) condition, which were presented in a pseudo-randomized order. Attractiveness and happiness ratings were given on a 7-point scale by using the keys 1-7 on the keyboard (see figure 1B for trial overview). After 40 trials, two additional questions were asked, assessing motivation to meet the men in the task and extent to which feelings of rejection were experienced. Two versions of the task were created, containing different stimuli that were matched for race and attractiveness (see figure 2 for examples), as measured in a preliminary survey (n=241) in a Dutch sample. The two study sessions used the different versions of the task, so participants would never see the same men twice, and the versions were equally distributed over the placebo and T session.
EMG measurements
EMG activity was recorded from bipolar electrodes placed over the left zygomaticus major and the corrugator supercilii muscles, in accordance with guidelines by Fridlund and Cacioppo (1986). Two ground electrodes were placed midline on the forehead for reference (see figure 1C). The Biosemi ActiveTwo amplifier was used to record the EMG at a 2048 Hz sampling rate. Offline, the raw EMG signal was filtered between 30 – 500 Hz, rectified, segmented per feedback condition (i.e., rejection and acceptance) and baseline corrected. The data was then divided in segments of 4000ms (covering -1000 – 0ms baseline and 0-3000ms feedback presentation), allowing the extraction of the post-stimulus average activity for each individual trial (i.e., over the 0-3000ms interval). Subsequently, artifacts were removed in a completely objective manner, by evaluating the activity on each trial on the basis of being more than 3 SDs from the average for a given participant, session, feedback condition and muscle. This allowed for the removal of specific trials with artifacts, while preserving the rest of the data for that participant. The artifact-trials removal procedure was applied within-participants in order to prevent unjustified data removal due to large inter-individual differences typically observed in EMG signals. Finally, after artifact removal on the individual level, the >3 SD criterion was used at group level to reveal remaining outliers for which the signal was unreliable over the entire or a substantial part of the recording.
Because all stimuli were purposely chosen for high attractiveness3, the relative attractiveness of the stimuli and
dichotomous categorization of high vs low attractiveness is rather different for each participant, as opposed to some studies containing extremely attractive and extremely unattractive stimuli for which the allocation is unambiguous. Therefore, in the current design attractiveness is to be determined for each participant separately, which is both methodologically necessary and ecologically valid. To this end, an individual split in high and low attractiveness of stimuli is made based on the average rating per participant, per session. This average-based approach per individual will ensure a similar amount of trials per attractiveness level, in contrast to a fixed criterion or median split (on a 7-point scale).
For the subjective rating data, a repeated-measures ANOVA with drug (T vs. placebo), social feedback (approval vs. rejection) and attractiveness (attractive vs. unattractive) as within-subjects factors was used for happiness ratings. Similarly, for the EMG data, two repeated-measures ANOVAs with drug (T vs. placebo), social feedback (approval vs. rejection) and attractiveness (attractive vs. unattractive) were used for both the zygomaticus and corrugator activity.
RESULTS
Confirmatory analyses
Subjective experience of social feedback
In order to examine explicit subjective evaluations, a repeated-measures ANOVA with drug, feedback and attractiveness was conducted on the happiness ratings. A clear task effect appeared, with a main effect for feedback, a main effect for attractiveness and an interaction effect for feedback x attractiveness. Specifically, feedback of acceptance (M = 4.62, SD= .64) resulted in higher happiness ratings than rejection (M = 3.98,
SD = .54), F(1,19) = 8.20, p = .010, ηp2 = .301. Independent of feedback, attractive men (M = 4.40, SD= .30)
elicited high happiness ratings than unattractive men (M = 4.20, SD = .37), F(1,19) = 8.96, p = .007, ηp2 = .320.
Crucially, in line with task hypotheses, the interaction between feedback and attractiveness indicated that participants were specifically happy with acceptance and unhappy with rejection by attractive men while this pattern was reversed and slightly attenuated for unattractive men (see figure 3A), F(1,19) = 155.52, p < .001, ηp2
= .891. Post-hoc pairwise comparisons revealed that the difference in happiness ratings between rejection and acceptance was significant for both levels of attractiveness. Specifically, for attractive men, happiness ratings were higher upon acceptance (M = 5.64, SD= .56) than rejection (M = 3.15, SD= .64), F(1,19) = 114.95, p < .001, ηp2
= .858. For unattractive men, happiness ratings were lower upon acceptance (M = 3.60, SD= .72) than rejection (M = 4.80, SD= .82), F(1,19) = 15.88, p = .001, ηp2 = .455. Finally, no main effect or interaction was found with
drug (all p’s > .05).
Zygomaticus major activity
The EMG data are analyzed separately for each muscle. Due to equipment malfunction, for one participant EMG was not recorded. In addition, based on objective application of the 3 SD rule over the total dataset (as well as visual inspection) two participants were removed from analysis for zygomaticus activity, as their data was too noisy to be reliable4. For the remaining 17 participants, our strategy for artifact removal on individual trials (see
Methods) resulted in 97.9% of the total amount of trials being preserved5.
In order to investigate facial muscular responses of positive affect elicited by feedback presentation, a
3 The pilot study indicated that the male facial stimuli need to be highly attractive, as female raters are rather critical and
generally not happy with acceptance by men that are only moderately attractive.
4 Note that for these 2 participants, the signal over the entire recording was of bad quality and contained artifacts. Removal of
individual trials with artifacts did not suffice to acquire a reliable signal for these two participants, therefore they were eliminated from the analysis.
repeated-measures ANOVA with drug, feedback and attractiveness was conducted on the zygomaticus major activity. First, no significant main effect for feedback was found, F(1,16) = 2.23, p = .115. Second, we did find an effect for attractiveness, F(1,16) = 9.06, p = .008, ηp2 = .362, with increased zygomaticus activity in response to
feedback given by attractive men (M = .202, SD= .31) compared to unattractive men (M = -.055, SD = .23), regardless of the value of the feedback6. A trend towards a significant interaction effect between feedback and
attractiveness appeared for zygomaticus activity, F(1,16) = 3.84, p = .068, ηp2 = .194, but this failed to reach
statistical significance. Specifically, analysis of pairwise comparisons showed that this trend was carried by the nearly significant difference in zygomaticus activity in response to acceptance compared rejection by attractive men (F(1,16) = 3.98, p = .063, ηp2 = .199), while there was no such difference for unattractive men
(F(1,16) < 0.01, p = .971).
Interestingly, examining the influence of the factor drug in a three-way interaction did reveal a significant effect, F(1,16) = 9.68, p = .007, ηp2 = .377, indicating that T moderated the interaction between feedback and
attractiveness on zygomaticus activity. In order to disentangle this thee-way interaction, two ANOVAs were conducted for each drug condition separately. For the placebo condition, a clear task effect appeared (i.e., the interaction between feedback x attractiveness, which was nearly significant in the overall ANOVA),
F(1,16) = 9.00, p = .008, ηp2 = .360, indicating that under placebo, a similar pattern as for the subjective ratings
was observed. That is, for attractive men, acceptance elicited more smiling than rejection and for unattractive men, this the other way around. However, in the T condition, this interaction effect disappeared, F(1,16) < 0.01,
p = .955, indicating that under T, there was no difference in the pattern of smiling to acceptance vs rejection by
attractive or unattractive men.
Lastly, this interaction effect seems to be driven by feedback from attractive men rather than unattractive men. Paired samples t-tests showed that, in the placebo condition, zygomaticus activity was significantly higher when accepted by attractive men (M = .562, SD = .90), compared to when rejected by attractive men (M = .044,
SD = .45), t(16) = 2.365, p = .031, d = 0.621, two-tailed. Under T, there is no such effect, t(16) = 1.05, p = .308,
two-tailed. Moreover, acceptance by attractive men elicited significantly more zygomaticus activity under placebo (M = .562, SD = .90), than under T (M = .182, SD = .33), t(16) = 2.14, p = .048, d = 0.722, two-tailed. For the rejection by attractive men, there was no difference between the placebo and T condition, t(16) = 0.157, p = .887, two-tailed. These findings indicate that the increased zygomaticus activity in response to acceptance by attractive men is suppressed by T. For unattractive men, none of these differences are observed (all ps > .05).
Corrugator supercilii activity
With regard to corrugator supercilii activity, a repeated-measures ANOVA with drug, feedback and attractiveness was conducted. There were no main effects for feedback or attractiveness. However, a significant task effect (i.e., interaction between feedback and attractiveness) did appear, F(1,18) = 4.70, p = .044, ηp2 = .207, displaying the
same pattern as seen on the subjective ratings and zygomaticus activity, yet reversed (as corrugator activity reflects negative affect, instead of positive). That is, acceptance by attractive men caused corrugator relaxation, while rejection by attractive men increased corrugator activity and this pattern was reversed (or rather, any difference between acceptance and rejection was absent) for unattractive men (see figure 3C, and figure 4C and 4D for a visualization of the corrugator response over time). Indeed, pairwise comparisons revealed that the difference in corrugator activity between acceptance and rejection was nearly significant for attractive men, F(1,19) = 3.36,
p = .084, while there was no substantial difference for unattractive men, F(1,19) = 0.38, p = .545. Finally, no
significant effects for drug were found on corrugator activity (ps > .05).
A B
6 Note that EMG activity is measured over the textual feedback presentation, when the face had already disappeared from the
C D
Figure 3. (A) Subjective experience of social feedback. Happiness ratings for outcome of the dating task, displayed per level of
attractiveness of the stimuli as individually rated, per feedback (i.e., acceptance vs. rejection) condition and per drug condition (i.e., placebo and testosterone). The dashed line represents neutral affect, with positive affect above and negative affect below the line.
(B) Zygomaticus major activity, reflecting positive affect elicited by the social feedback, displayed per level of attractiveness, per
feedback condition and per drug condition. (C) Corrugator supercilii activity, reflecting negative affect elicited by the social feedback, displayed per level of attractiveness, per feedback condition and per drug condition. Error bars indicate standard errors.
(D) Correlation effect zygomaticus and ratings for acceptance by attractive men. Scatterplot of the relation between the
T-effect (i.e., difference in response under T compared to placebo) for zygomaticus activity and subjective happiness rating, demonstrating that the strength and direction of the T-effect on the subjective ratings is related to that on the nonconscious zygomaticus activity. That is, on average, participants who were less happy under T with acceptance by attractive men, also showed this attenuated affective response in zygomaticus activity, and vice versa.
General assessment of EMG responses
in facial muscular activity from baseline. To this end, it was checked whether the activity in each condition significantly differed from zero. For the zygomaticus, this was only true for acceptance by attractive men under placebo (t(16) = 2.58, p = .020), and under T (t(16) = 2.26, p = .038), although it was previously demonstrated that the difference between activity under placebo and T was significant, reflecting substantial suppression by T. For the corrugator, activity likewise only differed from zero in the condition of acceptance by attractive men under placebo, t(19) = -2.53, p = .021. For all the other conditions, there was no significant rise or decline in muscle activity compared to baseline (ps > .05). It thus appears that the social feedback in the dating task only induced a substantial nonconscious affective response as reflected in facial muscles for positive (i.e., zygomaticus) and negative (i.e., corrugator) affect, in the case of acceptance by attractive men and this response is significantly suppressed (zygomaticus) or eliminated (corrugator) by T administration (see figure 3B and 3C, and figure 4 for visual illustration).
Mood measurements and self-confidence
To test for a possible effect of T on self-reported mood, a repeated-measures ANOVA with drug (T vs. placebo), time (pre-intake vs. post-intake) and valence (positive vs. negative affect) was performed. There were no significant main effects or interaction effects. Crucially, the interaction between drug x time was not significant,
F(1,19) = 0.36, p = .558, nor was the three-way interaction between drug x time x valence, F(1,19) = 0.53, p = .476, indicating that T administration did not significantly affect participants’ mood and can thus not explain
the effects above-mentioned effects.
Furthermore, in order to exclude the possibility of the confounding factor of baseline difference in participants’ rating of their own attractiveness (i.e., self-confidence) between the two sessions, differences in attractiveness ratings were compared between drug sessions7. A paired-sample t-test on the item measuring the
participants’ judgment of their own attractiveness between the placebo and T session was nonsignificant, t(19) = .57, p = .58, two-tailed, indicating that the current results were not due to a priori differences in self-confidence.
Manipulation checks
Although 70% of the participants correctly guessed when they received T and placebo, the subjective administration scores were still at chance level, binomial p = 0.12, two-tailed. Moreover, there was no relation between subjective guess of administration and dependent variables; the ANOVAs with T-effect for feedback and attractiveness as within-subjects factor and correctness of guess as between-subjects factor for ratings and zygomaticus activity showed no significant interactions (all ps > .05 and crucially, the three-way interaction with drug x feedback x attractiveness on ratings was not significant F(1,18) = 1.351, p = .26, the three-way interaction on zygomaticus activity disappeared, F(1,15) = 0.01, p = .97 and the three-way interaction on corrugator activity was also not significant, F(1,17) = 0.19, p = .67).
With regard to the believability of the task, as assessed with the open question, 60% of the participants indicated to have some doubts about the genuineness of some aspects of the dating task, mainly mentioning that the photos looked somewhat too professional to be taken by the men themselves. However, when asked to rate the believability of the task on a 1-10 scale (see Methods), the mean rating was 7.75 (SD = 1.99), with 5 being the lowest rating, indicating that participants generally took the task at face value. Moreover, this believability rating was unrelated to any of the dependent variables (all ps > .05) as indicated by correlations of the believability rating with zygomaticus and corrugator activity and subjective ratings in all conditions, as well with T-effect values (difference score between T and placebo) for zygomaticus and corrugator activity and subjective ratings in all conditions. The lack of significant correlations demonstrated that the extent to which participants believed the paradigm did not influence how they reacted on the task.
7 Note that judgment of participants’ own attractiveness was an item on the checklist, which was completed in the morning
Lastly, only one participant correctly mentioned the study purpose of investigating the effect of T on the experience of social evaluation. She believed that T makes one care less about the judgment of others. However, because her responses did not show this alleged pattern, it was decided to keep her data in the analyses.
Effects of administration order
The order of drug administration was counterbalanced between participants to minimize the confounding effect of order. Indeed, no significant effects were found involving the between-subjects factor of order in which the drug was administered on any of the dependent variables (i.e., ratings, zygomaticus and corrugator activity) when added to the ANOVAs, except for a significant interaction between order x attractiveness on the ratings, F(1,18) = 6.98,
p = .017, ηp2 = .279. Further pairwise comparisons showed that participants who received T on the first session
and placebo on the second, were significantly less happy with feedback by unattractive men compared to attractive men (irrespective of the value of the feedback), F(1,18) = 18.44, p < .001, ηp2 = .506. For participants who first
received placebo and then T, there was no difference, F(1,18) = .31, p = .583. However, because there was no three-way interaction with drug x order x attractiveness, nor any other drug or order effects on the ratings, it was decided to omit this factor in further analyses.
Exploratory analyses
In addition to the primary confirmatory analyses, several additional statistical tests were performed to further explore the data. Notably, these analyses serve mainly to provide possible explanations for or links between the reported effects, and should only be interpreted as such.
Additional testosterone effects
In addition to the main analyses with the effect of T on the experience of social evaluation, some extra analyses were performed. Firstly, we analyzed whether T influenced attractiveness judgments of the male stimuli (conscious ratings and nonconscious responses). A paired-samples t-test was performed with regard to average attractiveness ratings, revealing no difference between the placebo and T condition on attractiveness judgments of the men in the task, t(19) = -1.41, p = .17, two-tailed. Additionally, two repeated-measures ANOVAs with drug and attractiveness were performed on zygomaticus and corrugator activity in response to the facial stimuli presentation. For zygomaticus activity, there was no main effect of drug, F(1,16) = 1.99, p = .177, nor an interaction between drug and attractiveness, F(1,16) = 0.12, p = .730. Similarly, for corrugator activity, there was no main effect of drug, F(1,19) = 1.79, p = .198, nor an interaction effect, F(1,19) = 1.47, p = .241.
In addition, due to the documented effects of T on approach motivation and punishment sensitivity, we examined the T-effect on the items assessing participants’ motivation to meet the men in the study and the extent to which they experienced feelings of rejection. Paired-sample t-tests revealed no effect of T on motivation for contact, t(19) = .64, p = .53, two-tailed, nor on rejection experience, t(19) = -1.14, p = .27, two-tailed. It should be noted though, that the last two concepts were measured with only one item on both sessions and might therefore not be sufficiently sensitive for subtle T effects.
Correlation subjective experience and zygomaticus activity
Lastly, in order to examine the relation between conscious affective ratings and nonconscious muscle responses, correlations were performed for each level of feedback and attractiveness of the stimuli. Due to large baseline individual differences in EMG activity, correlations between EMG measures and ratings did not yield any significant results (all ps > .05). Additionally, we calculated the T-effect for zygomaticus activity and the T-effect for the ratings in the acceptance by attractive men condition (as this was the only condition to display a significant T-effect on the EMG activity), by subtracting the value under placebo from that under T for each variable. This allowed the examination of a relation between the conscious ratings and nonconscious muscle activity specifically
for the effect of T (and eliminates distortion due to the large inter-individual variability in EMG signal). A significant correlation between the T-effect on zygomaticus activity and ratings was found, r(17) = .625, p = .007, indicating that participants who demonstrated the strongest attenuation of happiness by T in response to acceptance by attractive men, also showed the strongest attenuation of zygomaticus activity by T in this condition (see scatterplot figure 3D). Indeed, this correlation did not appear for any of the other conditions, which showed no substantial T-effect on facial measures at all (ps > .05).
Figure 4. Averaged EMG activity in mV of zygomaticus muscle under placebo (A) and under T (B), and corrugator muscle
under placebo (C) and under T (D) over time, from feedback presentation (time 0). Activity is displayed per feedback condition and attractiveness level. Error bars indicate standard errors.
Note: As the EMG was measured over the textual feedback presentation, rather than the facial stimuli, it represents an affective response, rather than a mimicry response. Therefore, these plots do not display the classical time course that is seen for facial mimicry (e.g., Hofman et al., 2012), However, they are informative for the sake of visualization.
Table 1
Acceptance Rejection
Attractive Unattractive Attractive Unattractive
Placebo T Placebo T Placebo T Placebo T
Subjective happiness ratings a 5.59(.69) 5.70(.53) 3.72(.76) 3.47(.76) 3.20(.76) 3.10(.65) 4.85(.93) 4.76(.89) Zygomaticus activity b .562(.90) .182(.33) -.068(.24) -.047(.29) .044(.44) .020(.45) .091(.66) -.198(.46) Corrugator activity c -.355(.61) -.237(.63) -.037(.75) .021(.20) .339(1.50) -.025(.58) -.044(.56) -.075(.32)
Note. Displays mean values, with standard deviations given in parentheses.
a Measured on a 7-point scale; n = 20 b n = 17 c n = 19
DISCUSSION
In the present study, we investigated the effects of T administration on affective responses to receiving social feedback of acceptance and rejection by a potential romantic partner in single women. Firstly, we validated the dating task by showing that, conform expectations, acceptance by attractive males elicited a substantial increase in happiness, and rejection by attractive men a decrease in happiness, whereas the pattern for unattractive men was reversed (i.e., the feedback x attractiveness interaction on the subjective ratings). This feedback x attractiveness interaction effect correspondingly appeared on physiological measures of facial muscle activity indicative of negative affect (corrugator supercilii), as well as partially on the muscle reflecting positive affect (zygomaticus major). Moreover, contrary to expectations, no T-effect was found on the subjective ratings or corrugator activity. However, T was found to indeed moderate the feedback x attractiveness interaction effect as exhibited in zygomaticus activity. Interestingly though, opposite predictions, T appeared to particularly suppress rather than amplify automatic positive affect in response to acceptance by a highly attractive male.
This seems puzzling, as bearing on reward theories, several studies demonstrated T to increase sensitivity towards positive outcomes, (Hermans et al., 2006a; Lombardo et al., 2012; Op de Macks et al., 2011; van Honk et al., 2004; Wu et al., 2016) and particularly so for outcomes with high mating-relevance, a finding often reported in female samples (Alexander & Sherwin, 1993; Tuiten et al., 2000; van Wingen et al., 2008a; Wang et al., 2014; Wingfield et al., 1990). The fact that we found T to attenuate positive affect (as reflected in zygomaticus activity) in response to acceptance feedback from attractive males, thus seems highly remarkable. Indeed, because it is exactly this condition which is most relevant with respect to mating behavior, as well as the most rewarding, a T-effect was expected in this condition. However, based on the abovementioned literature, an T-effect in precisely the
opposite direction, i.e., an increase in positive affect upon acceptance by attractive males, was predicted. Motivation vs appreciation
A few possible explanations for these results can be considered. First of all, however, we discard the probability of the absence (on ratings and corrugator activity) and inversion (on zygomaticus activity) of T-effects to be due to invalidity of the social feedback task itself. That is, based on the fact that 1) a strong feedback x attractiveness effect appeared on subjective ratings, which was (albeit to a lesser extent) mirrored in EMG measures, 2) the believability of the task was rather high, and 3) participants indicated to genuinely feel quite rejected by the males, it is deduced that the dating task did in fact target feelings of social acceptance and rejection. A more plausible explanation for the current results holds that T might in fact have very different – contrasting – effects on the passive emotional experience of social evaluation by a potential mate, than on the active motivation to approach a
potential mate. That is, T might particularly enhance motivation to acquire and defend one’s social status (e.g., through rewards) and find potential mates, and at the same time protect against potential social status attacks or rejection of courtship attempts, by providing a ‘shield’ against other people’s judgment. Crucially, the latter appears not to be restricted to negative social outcomes (e.g., rejection) but includes suppressing emotional responsiveness in general, promoting an “I don’t care” attitude. In other words, the reported emotional blunting resulting from T might not only occur for negative affective stimuli such as fearful (van Honk et al., 2005) and angry (Terburg et al., 2012) facial expressions, but also for positive outcomes.
However, before elaborating on the proposed account of attenuation of affective responses by T, it should be noted that it remains strange that we only found an effect on the zygomaticus, and only in the acceptance condition. That is, the account outlined above would also predict the attenuation of negative affect in response to rejection by attractive males, as this also reflects dampened emotional responsiveness. However, the absence of a T-effect for rejection on the corrugator might not denote the absence of a T-effect on negative affect per se (i.e., of a conceptual nature), but rather a failure of the EMG measures to pick it up (i.e., of a methodological nature). That is, the baseline analyses showed that under placebo, acceptance by attractive males was the only condition that elicited significant EMG activity from baseline on the zygomaticus and corrugator at all, which was significantly suppressed (zygomaticus) and eliminated (corrugator) under T. It could thus be argued that this way of feedback presentation was not sufficiently strong enough to generally induce substantial facial muscle activity, because the threshold for affective appraisals to be picked up with facial EMG measurements is rather high – in contrast with robust effects for facial mimicry responses (de Wied et al., 2012; Hermans et al., 2006b; Hofman et al., 2012; Sims et al., 2012). The zygomaticus response for acceptance by attractive males might have been the only exception that reached the threshold, and therefore constitute the only condition that could validly show attenuation by T to start with. A stronger manipulation with higher EMG sensitivity could elucidate on these suggestions. For instance, Wiggert et al. (2015) used more natural forms of acceptance and rejection feedback by means of video clips with individuals verbally expressing positive or negative evaluations. This form of presentation might be an interesting approach for future studies in a dating paradigm, as actually hearing and seeing a person expressing negative or positive feedback will be more intense than solely reading textual feedback. Obviously, the small sample size could also partially account for the failure to find statistically significant results, as the corrugator effects were in fact in the direction representing attenuation of negative affect in response to rejection by T.
“Wanting” vs “liking”
Nevertheless, regardless of the actual existence of a T-effect on negative affect in response to rejection, the demonstrated suppression of positive affect in response to acceptance deserves further scrutiny. Specifically, the dissociation between “wanting” (motivational) and “liking” (affective) in positive affect might provide an important distinction for interpreting the current findings, as well as for reconciling them with the well-founded reward hypothesis of T. That is, almost all existing studies on reward sensitivity exclusively focus on appetitive motivation and use a design in which rewards can be actively pursued, for instance in economic games (Aarts & van Honk, 2009; Coates & Herbert, 2008; Eisenegger et al., 2017; Johnson & Wood, 2001; Mehta & Josephs, 2006; van Honk et al., 2004; Wu et al., 2016), measure brain activity during those active behavioral tasks (Mehta & Beer, 2009; Op de Macks et al., 2011), or during reward anticipation (Hermans et al., 2010), rather than addressing the hedonic response to rewards (affective), as we did in our study. In other words, due to the adaptive and instrumental nature of T, rewards are indeed more pursued (and this pursuit is less inhibited by anxiety), yet perhaps not necessarily more, but even less enjoyed.
Several indirect lines of evidence could support the suggestion that T causes an attenuation of one’s passive emotional sensitivity for other people’s actions or emotions, in order to promote an active behavioral response. For instance, a prevailing explanation for the finding that T reduces empathy (Hermans et al., 2006b;
van Honk et al., 2011a), appertains to the rationale that closeness and affiliation with others should be suppressed in the face of competition because associated negative feelings of guilt diminish one’s motivation to act and one’s competitive drive (Eisenberg, 2000; Van Kleef et al., 2010). Crucially, the finding by Hermans et al. (2006b) that T administration decreased empathy as reflected in facial mimicry, is evidently in line with the postulation that T dampens positive emotions elicited by other individuals’ actions or emotions. Moreover, T might motivate sexual and romantic interest (i.e., "wanting"; Lopez et al., 2009; Roney, 2009; Slatcher et al., 2011; Tuiten et al., 2000; van Anders et al., 2007; van der Meij et al., 2008; Zilioli et al., 2014), but not necessarily satisfaction with and emotional appreciation of sexual behavior and romantic interactions (i.e., “liking”). Indeed, Edelstein et al. (2014) reported a negative correlation between T levels and emotional connectedness and relationship satisfaction.
In addition, a parallel can be drawn in an enhanced behavioral tendency – related to “wanting” and acting – combined with a reduced emotional responsiveness – related to “liking” and passively feeling – by T between both mating behavior and parenting behavior. For instance, while T has been related to reduced nurturance behavior (including emotional responsiveness to infants) based on the mating/parenting tradeoff (Archer, 2006; Kaplan & Gangestad, 2005; Wingfield et al., 1990), exogenous T administration has in fact also induced enhanced neural responsiveness to infant crying (Bos et al., 2010). Notably, while perception of infant crying is associated with enhanced neural activation in areas related to parental care under T, it is at the same time subjectively experienced as less aversive (i.e., dampened emotional response) This combination of enhanced motivation for active caring and reduced emotional responsiveness to infants by T (e.g., Fleming et al., 2002; Weisman et al., 2014) can be understood to adaptively enable parents to more readily act on crying. Likewise, high endogenous cyclic T levels are related to high reward motivation, yet unaffected appreciation ratings regarding baby faces (Hahn et al., 2015).
A possible psychological mechanism: self-confidence
The finding that T did indeed dampen rather than enhance responsiveness to positive feedback by attractive males in our dating task, reflecting an evolutionarily adaptive response, raises the question of how this mechanism actually works. In other words, what psychological process can underlie the effects? As T is instrumental in approach behavior, by means of anxiety-reducing mechanisms (Boissy & Bouissou, 1994; Hermans et al., 2006a; van Honk et al., 2005), it appears reasonable that these approach tendencies are mediated by a rise in self-confidence induced by T. Although there is little direct evidence, various studies do indeed suggest that a boost in confidence might underlie considerable behavioral effects of T. For instance, Eisenegger et al. (2017) recently demonstrated that basal T levels are related to motivation to compete, as well as to confidence in one’s cognitive performance, and Coates and Herbert (2008) discussed confidence to explain T effects in financial decisions. Additionally, prenatal T exposure has also been related to confidence, with higher T exposure linked to more (cognitive) confidence (Dalton & Ghosal, 2014; Neyse et al., 2016). Moreover, it has been argued that (over)confidence can be understood as a psychological mechanism aimed at achieving higher social status (Anderson et al., 2012), which can in turn be attributed to T mechanisms. More specifically related to social behavior, T has been shown to enhance confidence in patient populations with social anxiety disorder (Enter et al., 2016) and low T levels were found to be a marker for social insecurity in anorexia nervosa patients (Miller et al., 2007). A rise in confidence has furthermore been used to account for the positive relation between T levels and safe sex, a finding that at first glance contradicts the risk-taking feature of T, yet can logically be explained through the confidence link; high confidence is required for actually choosing safe sex, which is understood as the ‘bolder choice’ (van Anders et al., 2012). Lastly, Bedgood et al. (2014) argued that T might be mobilized to deal with stress associated with social evaluations specifically, emphasizing the adaptive role of T due to its stress-reducing and confidence-boosting properties. Unfortunately, in our study self-confidence was only measured during baseline, in order to exclude it as a potentially confounding factor. Therefore, no T-effects on self-confidence could be assessed in our data. Based on the suggestion of confidence being an interesting mediating factor in
T-effects on the experience of social feedback, we recommend future studies to include this concept as a dependent (mediating) variable in a sensitive manner8.
A possible neurobiological mechanism: OFC-limbic crosstalk
An underlying neurobiological mechanism that could potentially account for the attenuation of emotional responsiveness following social evaluation under T, involves the orbitofrontal cortex (OFC) and more specifically the OFC – limbic crosstalk. The OFC is typically related to the regulation of emotional responses (Ochsner & Gross, 2005) and the OFC – limbic crosstalk is argued to be particularly important for affective appraisals and emotional modulation of social information (Bachevalier & Loveland, 2006). T has indeed been shown to reduce OFC reactivity to social cues (Mehta & Beer, 2009; van Wingen et al., 2008a; van Wingen et al., 2009), as well as to reduce cortical-subcortical crosstalk (Schutter & van Honk, 2004), specifically in the connectivity between the OFC and amygdala (van Wingen et al., 2010). This neurobiological mechanisms also nicely accounts for the finding that T administration increased amygdala reactivity to social stimuli, reflected in facial anger, but reduced appraisal of these stimuli, reflected in poorer conscious recognition of facial anger (van Honk & Schutter, 2007). Moreover, the OFC - amygdala decoupling has been associated with the upregulation of dopamine in the OFC by T (van Honk et al., 2011b) and T has also been related to dopaminergic pathways in other brain areas, including the ventral striatum (Bell & Sisk, 2013; Campbell et al., 2010; de Souza Silva et al., 2009; Thiblin et al., 1999). Notably, dopamine activity in the latter in particular has been associated with motivational “wanting” – in contrast with hedonic “liking” (Berridge et al., 2009), in line with the suggested dissociation of T being more causally involved in “wanting” than “liking”. This, in turn, is corroborated by Hermans et al. (2010) who suggested that the increased blood oxygenation level-dependent (BOLD) response in the ventral striatum in anticipation of rewards observed after T administration, primarily reflects enhancement of appetitive reward pursuit (“wanting”) rather than hedonic appreciation (“liking”). However, the OFC – limbic decoupling and the role of dopamine remain highly speculative with respect to emotional appraisal regarding social evaluation, and should be investigated in future research before any conclusions can be drawn.
Open questions and limitations
The lack of a T-effect on the subjective ratings might be due to strength of feedback x attractiveness interaction effect, which overshadows the subtle T-effect and leaves it redundant. However, as previously mentioned, several T administration studies have demonstrated a dissociation between consciously reported and nonconsciously expressed affect or sensations (e.g., Schutter & van Honk, 2004; Tuiten et al., 2002; van Honk et al., 2005; van Honk et al., 2004). The absence of T-effects on the ratings could thus also reflect the genuine absence of T-effects on the overt affective experience of social acceptance and rejection by a potential mate. Future studies could provide further insight into the relation between conscious and nonconscious measure of emotional appraisal with regard to social evaluation particularly.
Notably, with regard to ecological validity, a considerable potential limitation of the present experiment refers to the fact that in spite of the relatively high believability ratings, the dating task might still not have been realistic enough, possibly also due to the lab setting. Crucially, the males might not have been seen as actual potential partners, because the women knew that they were foreign and could not be contacted. Finally, although it was explicitly not the focus of this study, it would definitely be interesting to investigate the influence of T on real-life dating behavior, in which, contrary to our study, motivational processes and self-agency play an essential role. A dissociation between the dampening of passive emotional reactivity observed in the current study, and
8 The sensitivity relates to the factor that in our study, self-confidence was not only assessed before drug administration, but
also with solely one item. As mentioned for the motivation item, which was also assessed with one question in our study, this one-item construct lacks sensitivity. Bearing on the significance of the outlined dissociation between the motivational “wanting” and hedonic “liking”, it is recommended that future studies, especially those with more ecological validity, should also target the motivation factor more sensitively.
potential enhancement of behavioral reactivity in a face-to-face dating setting would be of great interest for a more elaborate and nuanced interpretation of the effect of T on social evaluative experiences in a mating context.
Conclusion
In summary, the challenge hypothesis and reward theories might predict that T enhances positive affect for positive outcomes in a mating context. However, the present findings show that as opposed to increasing emotional appraisal in response to acceptance by potential mates, T rather suppresses passive positive affect. We argue this can be understood in light of the dissociation between motivational (“wanting”) and hedonic (“liking”) effects of T with respect to rewards, status or mates. Bearing on its adaptive nature, T appears to enhance mating motivation, but also dampen emotional reactivity to the judgment or actions of potential mates. These diverging effects both contribute to promoting future mating attempts and subsequent reproductive success.
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