Behavioral Neuroscience © 2017 American Psychological Association

2017, Vol. 131, No. 3, 262–273 0735-7044/17/$12.00 [Link]

Child Gender Influences Paternal Behavior, Language, and Brain Function

Jennifer S. Mascaro Kelly E. Rentscher

Emory University University of Arizona

Patrick D. Hackett Matthias R. Mehl

Emory University University of Arizona

James K. Rilling
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

Emory University
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Multiple lines of research indicate that fathers often treat boys and girls differently in ways that impact
child outcomes. The complex picture that has emerged, however, is obscured by methodological
challenges inherent to the study of parental caregiving, and no studies to date have examined the
possibility that gender differences in observed real-world paternal behavior are related to differential
paternal brain responses to male and female children. Here we compare fathers of daughters and fathers
of sons in terms of naturalistically observed everyday caregiving behavior and neural responses to child
picture stimuli. Compared with fathers of sons, fathers of daughters were more attentively engaged with
their daughters, sang more to their daughters, used more analytical language and language related to
sadness and the body with their daughters, and had a stronger neural response to their daughter’s happy
facial expressions in areas of the brain important for reward and emotion regulation (medial and lateral
orbitofrontal cortex [OFC]). In contrast, fathers of sons engaged in more rough and tumble play (RTP),
used more achievement language with their sons, and had a stronger neural response to their son’s neutral
facial expressions in the medial OFC (mOFC). Whereas the mOFC response to happy faces was
negatively related to RTP, the mOFC response to neutral faces was positively related to RTP, specifically
for fathers of boys. These results indicate that real-world paternal behavior and brain function differ as
a function of child gender.

Keywords: experience sampling, fathers, fMRI, gender socialization, play

Supplemental materials: [Link]

A number of studies argue that parents treat girls and boys 1977; Bussey & Bandura, 1999). For example, some research
differently, and learning theory proposes that parents model and indicates that mothers talk more (Leaper, Anderson, & Sanders,
reinforce gender stereotypes in their children (Bandura & Walters, 1998; MacDonald & Parke, 1984) and are more restrictive of
physical risk-taking with daughters (Morrongiello & Hogg, 2004)
than with sons. Other research indicates that western fathers and
mothers are more elaborative in autobiographical storytelling with
Jennifer S. Mascaro, Department of Family and Preventive Medicine, girls than boys (Fivush, 2011). Moreover, parents often encourage
Emory University School of Medicine, and Center for Translational Social gender-stereotyped play behavior and household chores and dis-
Neuroscience, Emory University; Kelly E. Rentscher, Department of Psy-
courage gender-atypical behavior (Lytton & Romney, 1991), and
chology, University of Arizona; Patrick D. Hackett, Department of An-
both mothers and fathers are more likely to engage in rough and
thropology, Emory University; Matthias R. Mehl, Department of Psychol-
ogy, University of Arizona; James K. Rilling, Center for Translational tumble (RTP) play with boys than with girls (McIntyre & Ed-
Social Neuroscience and Department of Anthropology, Emory University, wards, 2009; Paquette & Dumont, 2013; Pellegrini & Smith,
and Department of Psychiatry and Behavioral Sciences, Emory University 1998). Related lines of research also point to differences in paren-
School of Medicine. tal behavior related to child social and emotional development. For
This work was supported by a Positive Neuroscience Award from the example, fathers of young children attend more to daughters’
John Templeton Foundation and by NIH Grant R21HD078778. The study submissive emotional facial expressions (Chaplin, Cole, & Zahn-
was designed by Jennifer S. Mascaro, Matthias R. Mehl, and James K. Waxler, 2005), convey gender stereotypes while discussing emo-
Rillling Data collection, preparation, and analysis were conducted by
tions with their children (van der Pol et al., 2015), and are more
Jennifer S. Mascaro, Kelly E. Rentscher, and Patrick D. Hackett, and the
likely to express affection and respond to daughters’ pro-social
manuscript was written by Jennifer S. Mascaro, Matthias R. Mehl, and
James K. Rilling. behavior (Hastings, McShane, Parker, & Ladha, 2007).
Correspondence concerning this article should be addressed to James K. Despite the aforementioned research, there are considerable
Rilling, Department of Anthropology, Emory University, 207 Anthropology methodological challenges inherent to the study of gender social-
Building, 1557 Dickey Drive, Atlanta, GA 30322. E-mail: jrillin@[Link] ization that limit confidence in previous findings. First, the social-

262
 CHILD GENDER AND PATERNAL BEHAVIOR 263

ization of gender takes place in complex and multiply determined (Feldman, 2015; Rilling, 2013; Rilling & Mascaro, 2017). How-
environments that do not lend well to testing mechanistic models ever, to date no studies have examined the possibility that fathers’
(Eagly & Wood, 2013). Second, parents rarely self-report differ- neural responses differ depending on whether they have a son or
ential treatment of sons and daughters (e.g., (Lytton & Romney, daughter, nor have any studies examined whether these gender
1991), highlighting the importance of obtaining real-world obser- differences relate to differences in behavior outside of the labora-
vational measures (Fivush, 1998). Another challenge to the current tory context. The present study set out to examine whether fathers’
research on gender differences in caregiving is that a majority of brains respond differently to daughters versus sons and whether
this research has been conducted on mothers, and it may be these differences relate to gender differences in behavior.
particularly important to examine gender differences in paternal To investigate the impact of child gender on paternal behavior
behavior given research indicating that paternal gender socializa- and brain function in a sample of fathers with children aged 1–2
tion, compared with maternal socialization, may be disproportion- years, we first examined whether fathers of daughters were more
ately influential on child outcomes (Fitzgerald, 1977; Gottman, attentive to their toddler, expressed more empathy, and used more
Katz, & Hooven, 1997; Grossmann et al., 2002; Power, 1981) and affective expressions (laughing, singing, and complaining) and
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that fathers tend to more robustly differentiate between boys and emotion language (angry, happy, sad) than did fathers of sons. We
This document is copyrighted by the American Psychological Association or one of its allied publishers.

girls (Lytton & Romney, 1991). also investigated whether fathers of sons differed in their play
Systematic investigation of paternal caregiving may be partic- activities (nonphysical play, RTP, and reading) compared with
ularly imperative given the role that socialization plays in child fathers of daughters. Second, we examined whether fathers of
outcomes, particularly in the development of empathy and altru- daughters have a greater neural response to their daughters’ emo-
ism. Since the earliest meta-analyses over three decades ago tional facial expressions than fathers viewing their sons’ emotional
(Eisenberg & Lennon, 1983), a multitude of studies have reported facial expressions. In particular, we were interested in neural
on an apparent sex difference in empathy and its relevant compo- responses in areas of the brain important for reward and for
nents (Baron-Cohen et al., 2015; Christov-Moore et al., 2014; processing facial expressions of emotion, such as the ventral
Hoffman, 1977; Moriguchi, Touroutoglou, Dickerson, & Barrett, striatum, the OFC, the inferior frontal gyrus (IFG), and the anterior
2014). Women in industrialized Western cultures are often found insula (AI; Feldman, 2015; Rilling & Young, 2014; Swain, Day-
to have more interdependent concepts of self (Cross & Madson, ton, Kim, Tolman, & Volling, 2014). In an exploratory next step,
1997) and are more likely to have a nurturing response to psycho- we examined whether neural responses that differed by child
social stress (Taylor et al., 2000). Although these data are often gender were correlated with the behavioral variables that differed
interpreted as biologically determined sex differences, they may by child gender, and if so, whether brain activity mediated the
also emerge from gendered socialization of emotion. Parental impact of child gender on paternal behavior.
behavior is thought to influence the development of empathy and
social intelligence. For example, a retrospective study found that Method
adult empathy levels were related to paternal involvement in
caregiving and maternal inhibition of aggression (Koestner, Franz,
& Weinberger, 1990). Another found that maternal warmth and
Participants
discipline style predicted peer-rated empathy of her child in adult- As part of a larger study on paternal caregiving, we recruited 69
hood, controlling for childhood levels of empathy (Eisenberg, heterosexual, biological fathers of children age 1 or 2 who were
VanSchyndel, & Hofer, 2015). currently cohabiting with the child’s mother using flyers posted
Examining gender differences in paternal behavior will be cru- around the Emory campus, at local parks, daycare centers, and
cial for understanding gender socialization of empathy. Gender with an electronic advertisement on Facebook. The study was
socialization appears to moderate the relationship between care- approved by the Emory Institutional Review Board, and all par-
giving and child outcomes, and paternal impact may be particu- ticipants gave written informed consent. Participants had normal or
larly influential based in part on evidence that fathers may be more corrected-to-normal (with contact lenses) vision and were screened
likely to invoke gender stereotyped norms than mothers (Fabes & and excluded for self-reported history of head trauma, seizures or
Martin, 1991). Fathers of preschool children are more likely to other neurological disorder, psychiatric illness, alcoholism or any
engage in sex-typed socialization of empathy, with fathers of boys other substance abuse, serious medical illness, claustrophobia, and
reporting less inclination to reinforce pro-social behavior and less for ferrous metal in any part of body.
affection toward their sons (Hastings et al., 2007). Fathers of Fathers were between the age of 21 and 55 years (M ⫽ 33.0,
young children in the Netherlands also exhibit more explicit gen- SD ⫽ 5.67) and had between 1 and 4 children, with 2 as the modal
der stereotypes than mothers (Endendijk et al., 2013). number (M ⫽ 1.79, SD ⫽ 0.74; see Table 1 for descriptive
If parents differ in their behavioral responses toward sons and statistics). Thirty-four of the fathers had a daughter and 35 of the
daughters, brain function should mediate these gendered differ- fathers had a son. Fathers of daughters and sons did not differ in
ences. To date, several studies have examined the neurobiology age, child age, ethnic composition, yearly combined household
supporting paternal caregiving. These data converge to suggest income, or hours worked per week.
that paternal caregiving relies on brain systems similar to those
involved in maternal caregiving, including the midbrain dopamine
Procedure
system for approach motivation [the ventral tegmental area (VTA)
coupled with the substantia nigra (SN), the ventral striatum (VS), First, the mother completed self-report parenting questionnaires
and the medial orbitofrontal cortex (mOFC)], along with cortical (results published elsewhere: Mascaro, Hackett, Gouzoules, Lori,
regions involved in mirror simulation, empathy, and mentalizing & Rilling, 2014; Mascaro, Hackett, & Rilling, 2013, 2014) while
 264 MASCARO, RENTSCHER, HACKETT, MEHL, AND RILLING

Table 1 A group of eight trained research assistants coded the audio

Descriptive Statistics of Fathers of Daughters and Sons files. All audio files were independently coded by two research
assistants and, in all cases, one coder coded all of a participant’s
Girls’ Boys’ audio files. Coders were instructed to listen to each audio file at
Mean (SD) Mean (SD)
Statistic (n ⫽ 30) (n ⫽ 22) p least twice and to use context information (from previous and
consecutive sound files) to increase the accuracy of their codings.
Child age 1.98 (.65) 1.93 (.59) .74 Captured words by the father were transcribed verbatim (with the
Father age 33.0 (4.84) 33.0 (6.54) 1.00
Number of children 1.93 (.78) 1.64 (.66) .16 first coder transcribing the words and the second coder cleaning
Father ethnicity .33 the first coder’s verbatim transcript). Coders were blind to the
White 21 21 current research hypotheses. The data were collected in Atlanta,
Hispanic 4 1 Georgia and the audio files were coded in Tucson, Arizona,
Black 5 8
minimizing any chances that the research assistants would know or
Asian 3 1
Other or not reported 1 4 recognize any of the participants.
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Household income (year) $82,000 (53,000) $98,000 (53,000) .26 A new coding system for extracting a range of social and
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Father hours worked (week) 38.2 (15.3) 39.9 (13.4) .65 emotional aspects of father-child interactions from the recorded
ambient sounds was developed specifically for the purpose of this
study. A full description of the development process is beyond the
experimenters took naturalistic photographs of the child engaged scope of this paper but will be reported elsewhere. In short, the
in a play session. In a separate session, fathers were consented and new coding system was based on prior experiences with coding
given a mobile audio recording device, the Electronically Acti- EAR recordings (Alisic et al., 2015; Mehl et al., 2012), other
vated Recorder (EAR; Mehl, in press) that intermittently records existing lab-based observational coding systems, and important
ambient sounds around them (described below) and asked to wear information unique to this study (e.g., whether the participant was
the device on one weekend day (Sunday) and one weekday (Mon- talking to his child, spouse, or a coworker). The coding categories
day). In a third laboratory session, fathers completed self-report included father affective expression (e.g., laughing, singing, com-
parenting questionnaires and provided saliva and blood samples, plaining), toddler affective expression (e.g., laughing/giggling,
and were then positioned in the MRI scanner where they received fussing/crying), father-toddler activities (e.g., reading, nonphysical
structural and functional MRI scans of their brain. play, physical play), and paternal caregiving behaviors (e.g., en-
gaged attending to the child, showing empathy or affection, ex-
pressing praise and encouragement). All codings were binary,
Electronically Activated Recorder
reflecting the presence versus absence of a particular behavior in
Technically, the EAR is an audio recorder that intermittently any given 50-s audio file. For fathers with more than one child,
records snippets of ambient sounds while participants go about behaviors were coded as present only if they were directed toward
their lives. Conceptually, it is a naturalistic observation method the target child.
that yields an acoustic log of a person’s day as it unfolds. The For the purpose of this study, interrater reliability for the vari-
version of the EAR used in this study consisted of a Dell ables of interest, computed as one-way random intraclass correla-
Axim ⫻50 handheld computer with the EAR software pro- tions for the average measure (ICC[1,2]), was: Singing: .87 (M ⫽
grammed to record 50 seconds every 9 min. In reliability analyses, 1.8%, SD ⫽ 1.7%); Engaged Attending to the child: .86 (M ⫽
this sampling rate has yielded stable estimates of daily behaviors 14.0%, SD ⫽ 8.3%); Physical Play: .83 (M ⫽ 0.4%, SD ⫽ 0.7%);
and robust effect estimates with criterion variables (Mehl, Rob- Showing Empathy: .35 (M ⫽ 0.3%, SD ⫽ 0.4%). The empathy
bins, & Deters, 2012). variable was excluded from subsequent analyses due to its low
We collected naturalistic observation data on fathers’ real-world interrater reliability. Variables were calculated with reference to
behavior toward their children from 55 fathers (25 boys, 30 girls), the total number of (waking and compliant) audio files [M(t)], such
as 14 fathers experienced complete equipment failure (most fre- that a M(t) value of 1.0 indicates that 1% of all of the audio
quent reasons were they forgot to charge it or the device was set recordings contained that behavioral variable. We also report the
incorrectly). Participants were excluded if they had fewer than 50 percentage of the total number of audio files that included inter-
audio recordings during the 48-hour period (n ⫽ 3), so the final actions with the child [M(c)], such that a M(c) value of 1.0
EAR data set contained 52 participants (22 boys, 30 girls). Fathers indicates that 1% of the audio recordings where the father was
wore the EAR device in a protective case clipped to their waistline, interacting with the child contained that behavioral variable.
with an Olympus ME-15 external microphone clipped to their To analyze the language recorded by the EAR, we employed
lapels. The EAR was set to record between the hours of 8 a.m. Linguistic Inquiry and Word Count (LIWC) (Pennebaker, Boyd,
Sunday and 8 a.m. on Tuesday and participants were asked to store Jordan, & Blackburn, 2015), which is currently the most exten-
and charge the device in their child’s room at night. Participants sively validated word-count based text analysis tool in the social
had an average of 158.0 (SD ⫽ 30.6) valid (i.e., the father was sciences. To derive information about fathers’ language use when
wearing the device, and there were no technical problems), waking talking to their children, the overall verbatim transcripts were first
(i.e., the father was not sleeping) audio files that could be coded. separated by interaction partner. Information about whether the
For more information on the validity and ethical implications of participant was talking to his child (and specifically, the target
the EAR method for studying parent– child interactions see (Alisic child), spouse, a coworker, or someone else was derived from the
et al., 2015; Alisic, Barrett, Bowles, Conroy, & Mehl, 2016; Mehl behavioral codings. The fathers’ words spoken to their child,
et al., 2012; Slatcher & Robles, 2012). spouse, and coworker(s) were then submitted to three separate
 CHILD GENDER AND PATERNAL BEHAVIOR 265

LIWC analyses (in addition to a LIWC analysis of the fathers’ mm, 34 axial slices. TE was minimally decreased from the typical
overall word use). LIWC counts words by matching them to value (32 ms) to reduce magnetic susceptibility artifact in the
categories of dictionary words that represent specific domains, orbitofrontal region. Subjects were imaged while viewing pictures
such as affect words (e.g., sadness), social words (e.g., family), of happy (H), sad (S), and neutral (N) facial expressions in three
core drives and needs (e.g., achievement), and biological processes different people: (a) Their own child (O), (b) An unknown child
(e.g., body). The variables based on these analyses are expressed (U), and (c) An unknown adult (A). Participants were instructed to
as the percentages of all words spoken to the child, spouse, and “please observe each picture and try to share the emotions of the
coworker, respectively, in which a given LIWC category applied person in the picture.” For each expression, fathers viewed 8
(e.g., percentage of all words the father spoke to the child that were different pictures of the person making that expression over the
sadness or anger words). course of 4 blocks, and each picture was viewed twice (see Figure
To address our first aim, we conducted independent sample t S2). During a single block, 4 photographs of the same type were
tests and calculated Cohen’s d effect size values to examine shown, each for 3 seconds. There was a 0.5 second fixation
whether behavior and language variables of interest differed be- between each photograph. Thus the duration of each block was 14
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tween fathers of daughters and fathers of sons. For variables that seconds. After every 6 blocks, subjects viewed a fixation block of
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violated normality assumptions, we used the standard SPSS ad- equal duration. The total duration of the task was 9 min 48 seconds
justment of degrees of freedom using the Welch-Satterthwaite (36 face blocks ⫹ 6 fixation blocks ⫻ 14 s per block). Photographs
method. Behavioral variables of interest included engaged attend- were presented in pseudorandom order, and fathers always viewed
ing to the child (e.g., child is fussing and father responds), RTP own children at the end so that their responses to unknown chil-
(e.g., running, chasing, tickling, or swinging), and affectively dren could be compared to the responses from nonfathers in a
laden interactions such as laughing, singing, and complaining. For related study.
the LIWC analysis, we were particularly interested in emotion
language, such as that related to anger, sadness, and happiness.
Functional Image Analysis
Given the exploratory nature of our analyses, we first used an
alpha level of .05. Tests of the hypotheses were also conducted Image preprocessing was conducted with Brain Voyager QX
using Bonferroni adjusted alpha levels of .004 (.05/14). Statistics (version 2.0.8) software (Brain Innovation, Maastricht, The Neth-
were performed on M(t), but did not differ substantively when erlands). The first 8 volumes of each run were discarded to allow
performed on M(c). the tissue magnetization to equilibrate. Preprocessing involved
image realignment by six-parameter 3-D motion correction, slice
scan time correction using linear interpolation, spatial smoothing
Photograph Stimuli
with a 8-mm full width at half maximum (FWHM) Gaussian
As part of the larger study, fathers viewed photographs of an kernel, and temporal smoothing with voxel-wise linear detrending
unknown adult, an unknown child, and their own child. A descrip- and high-pass filtering of frequencies below three cycles per run
tion of the methodology related to the unknown adult and child are length. Six subjects were excluded because of movement greater
found elsewhere (Mascaro, Hackett, Gouzoules, et al., 2014; Mas- than 2 mm in the x, y, or z direction, so the final data set contained
caro, Hackett, & Rilling, 2013; Mascaro et al., 2014). For the 59 participants. Images were subsequently normalized into Ta-
present study, we were interested in fathers’ responses to their own lairach space (Talairach & Tournoux, 1988).
child’s face. We captured eight pictures of the father’s child A separate general linear model (GLM) was defined for each
making each facial expression during a play session. If the child subject that examined the neural response to the following nine
did not make one of the facial expressions naturally, sad faces were regressors: own child’s happy face (OH), own child’s sad face
elicited by the mother leaving the room or taking a favorite toy or (OS), own child’s neutral face (ON), unknown child’s happy face
cell phone from the child, and happy faces were elicited with (UH), unknown child’s sad face (US), unknown child’s neutral
singing, dancing, or tickling. See supplementary Figure S1 for face (UN), unknown adult’s happy face (AH), unknown adult’s
examples of each condition. sad face (AS), and unknown adult’s neutral face (AN). Because
there was substantial attenuation of the BOLD response upon
viewing each block repetition, only the first block of each condi-
Anatomical Image Acquisition
tion was included in the GLM. Each regressor was convolved with
Subjects were positioned in the Siemens Trio 3T MRI scanner. a standardized model of the hemodynamic response. The resulting
Subjects lay motionless in a supine position in the scanner with GLM was corrected for temporal autocorrelation using a first-
padded head restraint to minimize head movement during scan- order autoregressive model. For each subject, contrasts of param-
ning. Each scanning session began with a 15-s scout, followed by eter estimates for various predictors were computed at every voxel
a 5-min T1-weighted MPRAGE scan (TR ⫽ 2600 ms, TE ⫽ 3.02 of the brain.
ms, matrix ⫽ 256 ⫻ 256, FOV ⫽ 256 mm, slice thickness ⫽ 1.00 To address our second aim, we examined whether fathers of
mm, gap ⫽ 0 mm). daughters responded differently to their child’s happy, sad, or
neutral facial expressions than fathers of sons. To this end, we
conducted second level analyses using a standard analysis of
fMRI Image Acquisition
variance (ANCOVA) in which sex of the child was entered as a
Functional scans used an EPI sequence with the following between groups factor, which computes this effect separately for
parameters: TR ⫽ 2000 ms, TE ⫽ 28 ms, matrix ⫽ 64 ⫻ 64, each voxel in a whole brain exploratory analysis. Results were
FOV ⫽ 224 mm, slice thickness ⫽ 2.5 mm, gap thickness ⫽ 1.05 thresholded at p ⬍ .001, and were corrected for multiple compar-
 266 MASCARO, RENTSCHER, HACKETT, MEHL, AND RILLING

isons at the cluster level using Monte Carlo simulations as imple- t(25) ⫽ ⫺2.24, p ⫽ .03, d ⫽ 0.67. Levene’s test indicated unequal
mented by the Brain Voyager QX Cluster Threshold Estimator variances (F ⫽ 13.68, p ⫽ .001), so degrees of freedom were
plugin. For the exploratory aim, we extracted beta contrast values adjusted from 49 to 25. Fathers of daughters used more analytical
from the regions identified in the ANCOVA and entered them in language (M ⫽ 36.9%, SD ⫽ 19.3) than did fathers of sons (M ⫽
bivariate correlation analyses with the behavior and language 27.2%, SD ⫽ 13.5), t(49) ⫽ 2.015, p ⫽ .04, d ⫽ 0.58. To examine
variables identified as differing between fathers of sons and daugh- whether gender of child impacted the language fathers used in
ters. other contexts, the same analyses were performed for fathers’ use
of sadness, body, achievement and analytical language in the
Results context of coworkers and friends and there were no significant
differences between fathers of daughters and fathers of sons.
Behavior Results are summarized in Table 2.

The results of independent samples t tests comparing gender

Brain Activity
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means in behavior and language as a percentage of all audio

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recordings are summarized in Table 2. Fathers of daughters en- A whole brain exploratory analysis revealed that fathers of
gaged in significantly more singing or whistling with their child daughters respond more to their daughters’ happy facial expression
(M(t) ⫽ 1.40%, SD ⫽ 1.40; M(c) ⫽ 4.75%, SD ⫽ 5.08) than did in areas of the brain important for visual processing (BA 17 and
fathers of sons (M(t) ⫽ 0.30%, SD ⫽ 0.50; M(c) ⫽ 0.96%, SD ⫽ 18) and for reward, emotion regulation, and face processing,
1.36), t(38) ⫽ 3.69, p ⫽ .001, d ⫽ 1.05 (Figure 1). Levene’s test including medial OFC (mOFC) [BA 10, 11] and lateral OFC
indicated unequal variances (F ⫽ 7.80, p ⫽ .01), so degrees of (lOFC) [BA 47] and left middle frontal gyrus [BA 10] compared
freedom were adjusted from 50 to 38. Differences in singing by with fathers of sons (Figure 2a).
fathers of daughters and son were significant at the Bonferroni Fathers of boys responded more robustly to their sons’ neutral
adjusted alpha levels of .004. Fathers of sons engaged in signifi- facial expressions in the thalamus, mOFC, and inferior temporal
cantly more RTP play (M(t) ⫽ 0.70%, SD ⫽ 0.90; M(c) ⫽ 3.20%, sulcus (Figure 3a). There were no significant differences between
SD ⫽ 4.21) than did fathers of daughters (M(t) ⫽ 0.20%, SD ⫽ 0.40; fathers of daughters and fathers of sons in the neural response to
M(c) ⫽ 0.90%, SD ⫽ 1.38), t(26) ⫽ ⫺2.48, p ⫽ .02, d ⫽ 0.72. sad facial expressions. Results are summarized in Table 3.
Levene’s test indicated unequal variances (F ⫽ 9.581, p ⫽ .003), so
degrees of freedom were adjusted from 50 to 26. Fathers of daughters
Behavior–Brain Activity Associations
attentively responded to their child (M(t) ⫽ 16.1%, SD ⫽ 8.20;
M(c) ⫽ 54.5%, SD ⫽ 18.1) significantly more than fathers of sons Regarding the extent to which behavior was related to neural
(M(t) ⫽ 10.8%, SD ⫽ 7.40; M(c) ⫽ 42.6%, SD ⫽ 20.6), t(50) ⫽ 2.40, responses, bivariate correlation analyses revealed that singing was
p ⫽ .02, d ⫽ 0.68. positively correlated with activity in the right lOFC (r(40) ⫽ 0.31,
When talking to their children, fathers of daughters also used p ⫽ .05; Fathers of boys (B): r(16) ⫽ 0.34, p ⫽ .18; Fathers of girls
more language related to sadness (M ⫽ 0.45%, SD ⫽ 0.72) than (G): r(22) ⫽ 0.22, p ⫽ .30; Figure 2b) and left frontal pole (r(40) ⫽
fathers of sons (M ⫽ 0.15%, SD ⫽ 0.238), t(36) ⫽ 2.14, p ⫽ .04, 0.32, p ⫽ .04; B: r(16) ⫽ 0.49, p ⫽ .04; G: r(22) ⫽ 0.15, p ⫽ .48).
d ⫽ 0.57. Levene’s test indicated unequal variances (F ⫽ 4.085, RTP was inversely correlated with neural responses to happy faces in
p ⫽ .049), so degrees of freedom were adjusted from 49 to 36. the right lOFC (r(40) ⫽ ⫺0.40, p ⫽ .01; B: r(16) ⫽ ⫺0.32, p ⫽ .19;
Fathers of daughters also used more language about the body (M ⫽ G: r(22) ⫽ ⫺0.53, p ⫽ .01) and mOFC (r(40) ⫽ ⫺0.32, p ⫽ .04; B:
1.49%, SD ⫽ 1.165) than did fathers of sons (M ⫽ 0.82%, SD ⫽ r(16) ⫽ ⫺0.34, p ⫽ .17; G: r(22) ⫽ ⫺0.09, p ⫽ .68; Figure 2c) and
0.731), t(49) ⫽ 2.36, p ⫽ .02, d ⫽ 0.69. Fathers of sons used left lOFC (r(40) ⫽ ⫺0.32, p ⫽ .04; B: r(16) ⫽ ⫺0.11, p ⫽ .66; G:
significantly more achievement language (M ⫽ 0.91%, SD ⫽ 1.17) r(22) ⫽ ⫺0.47, p ⫽ .02). Amount of analytical language was posi-
than did fathers of daughters (M ⫽ 0.33%, SD ⫽ 0.41), tively correlated with activity in the right lOFC (r(39) ⫽ 0.32, p ⫽
.04; B: r(16) ⫽ 0.09, p ⫽ .74; G: r(21) ⫽ 0.35, p ⫽ .10). Activity in
the mOFC in response to neutral facial expressions was positively
Table 2 correlated with RTP (r(40) ⫽ 0.36, p ⫽ .02; B: r(16) ⫽ 0.49, p ⫽ .04;
Summary of Independent Samples t Tests Interrogating G: r(22) ⫽ ⫺0.21, p ⫽ .27; Figure 3b); correlations are summarized
Differences in Behavior and Language Between Fathers of in Table 4). In a final exploratory step, we examined whether brain
Daughters and Fathers of Sons activity meditated the impact of child gender on paternal behavior and
did not find any significant mediation effects.
Girls’ Boys’
Mean (SD) Mean (SD)
Behavior/Language (n ⫽ 30) (n ⫽ 22) t p Discussion
Rough and tumble play .20% (.40) .70% (.90) ⫺2.48 .02 Previous studies indicate that the paternal brain responds to
Singing or whistling 1.40% (1.40) .30% (.50) 3.69 .001ⴱ
Engaged responding 16.1% (8.20) 10.8% (7.40) 2.40 .02
pictures of children’s faces in much the same way as the maternal
Achievement .32% (.41) .91% (1.17) ⫺2.24 .03 brain, specifically in neural systems important for reward and
Analytical thinking 36.9% (19.3) 27.2% (13.5) 2.02 .05 motivation (mOFC, VTA, and ventral striatum), emotion regula-
Body 1.49% (1.17) .82% (.73) 2.36 .02 tion (lOFC and dorsolateral PFC), and empathy (anterior insula
Sadness .45% (.72) .14% (.24) 2.14 .04 and IFG; Feldman, 2015; Rilling, 2013; Rilling & Mascaro, 2017;
ⴱ
Significant at the Bonferroni adjusted alpha level of .004 (.05/14). Rilling & Young, 2014). The current project extends this work by
 CHILD GENDER AND PATERNAL BEHAVIOR 267
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Figure 1. (a) Fathers of daughters sang more, used more language related to the body and to sadness, were
more attentively engaged, and used more analytical language. (b) Fathers of sons engaged in more “rough and
tumble play” and used more achievement language. All variables are represented as a percentage with reference
to the total number of waking and compliant audio files. ⴱ indicates findings significant at the Bonferroni
adjusted alpha level of .004. All other findings are significant at the alpha level of .05.

demonstrating that fathers’ brains respond differently to the emo- are fathers of sons. This difference was only significant for the
tional facial expressions of daughters compared with sons. Specif- response to happy faces, which is consistent with previous research
ically, fathers of daughters had a more robust response to happy showing that fathers attend more to daughters’ happy facial ex-
facial expressions in visual processing areas, likely reflecting pressions (Chaplin et al., 2005) and are more likely to associate
increased attention of fathers to their daughters’ happy faces happiness with girls (van der Pol et al., 2015). We were surprised
(Lane, Chua, & Dolan, 1999). Moreover, fathers of daughters that fathers of daughters and sons did not differ in their response
exhibited a greater response in a region of the lateral OFC (BA 47) to sad facial expressions, but it is possible that the sad picture
involved in emotion regulation, especially implicated in augment- stimuli, many of which were of children actively crying, were so
ing positive emotion (S. H. Kim & Hamann, 2007). Greater salient and evocative that they elicited a robust response in all
activity here may also be interpreted as an effect of increased fathers.
attention. A recent meta-analysis indicated that lateral OFC (BA Fathers of daughters also had a more robust response to happy
10) and the inferior frontal gyrus orbitalis (BA 47) are often faces in the mOFC, a component of the mesolimbic dopamine
coactive during face processing tasks (Zald et al., 2014), providing system identified in diverse lines of research as important for
support for the interpretation that fathers of daughters are attending parental responding (Feldman, 2015; Morten L. Kringelbach et al.,
to and processing happy facial expressions to a greater degree than 2008; Rilling, 2013). For example, our previous work comparing

Figure 2. (a) Results of whole brain exploratory second level analysis indicating where fathers of daughters
have more activity in response to their own child’s happy faces. All results are thresholded at p ⬍ .001, corrected
for multiple comparisons using the Brain Voyager QX Cluster Threshold Estimator plugin. (b) Scatter plot of
paternal behaviors and beta contrast values derived from the R lOFC and medial OFC ROIs, plotted as a function
of group status. See the online article for the color version of this figure.
 268 MASCARO, RENTSCHER, HACKETT, MEHL, AND RILLING
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Figure 3. (a) Results of whole brain exploratory second level analysis indicating where fathers of sons have
more activity in response to their own child’s neutral faces. All results are thresholded at p ⬍ .001, corrected
for multiple comparisons using the Brain Voyager QX Cluster Threshold Estimator plugin. (b) Scatter plot of
“Rough and tumble” play and beta contrast values derived from the med OFC ROI, plotted as a function of group
status. See the online article for the color version of this figure.

fathers with age-matched nonfathers identified the mOFC as a positively correlated with RTP. In fact, the correlation between
region that was significantly more active in fathers in response to mOFC activity and RTP was only significant for fathers of boys.
unknown child faces (Mascaro et al., 2014). This region may be Neutral faces should not to be misinterpreted as blank faces,
integral for coding the hedonic pleasure associated with a stimulus devoid of meaning (Carvajal et al., 2013), and other lines of
(Kringelbach & Berridge, 2009), and intriguingly, it is this region research indicate that neutral facial expressions may be more
that appears to respond early and selectively in response to infant negatively or positively biased depending on context and charac-
faces (Kringelbach et al., 2008). Although speculative, this finding teristics of the perceiver (Demenescu, Kortekaas, den Boer, &
indicates that fathers of daughters may experience more pleasure Aleman, 2010; Leppänen, Milders, Bell, Terriere, & Hietanen,
when viewing their child’s happy faces than fathers of sons and 2004). The fact that fathers of sons respond more robustly to
future research can more definitively test this hypothesis by in- neutral expressions renders it possible that these fathers are more
cluding pleasure ratings to query fathers’ reactions to their chil- strongly negatively or positively biasing neutral facial expressions
dren’s happy faces. of their sons. Future research may employ an affective bias task to
It is interesting, then, that fathers of sons had a greater mOFC examine the possibility that fathers of sons are more likely to
response to neutral facial expressions, and that this activity was attribute emotional valence to boys’ neutral facial expressions, and

Table 3
Results of Whole Brain Exploratory Second Level Analyses Using a Standard Analysis of Variance (ANCOVA) in Which Sex of the
Child was Entered as a Between-Groups Factor for Own Happy [OH] and Own Neutral [ON]. Voxels are 1MM Isotropic.

Variable Brodmann’s area x y z Peak t Voxels

OH乆–OH么 (n ⫽ 59)
R Inferior frontal gyrus (orbitalis), extending into middle frontal gyrus 47,46 51 31 0 5.12 2133
R Cerebellum 17 ⫺74 ⫺30 3.89 468
R Striate cortex 17,18 14 ⫺89 ⫺3 3.78 740
Rostral sulcus, extending into inferior rostral gyrus 11,10 0 37 ⫺12 4.47 732
L Cerebellum ⫺31 ⫺77 ⫺30 4.35 840
L Middle frontal gyrus, extending into inferior rostral gyrus 10 ⫺34 53 ⫺3 5.04 1523
ON么–ON乆 (n ⫽ 59)
R Cerebellum 26 ⫺80 ⫺33 4.72 1038
R Thalamus 11 ⫺14 12 4.49 4296
L Medial orbital gyrus 11 ⫺13 22 ⫺13 5.18 593
L Inferior temporal sulcus 20 ⫺58 ⫺29 ⫺19 4.26 546
 CHILD GENDER AND PATERNAL BEHAVIOR 269

Table 4
Correlation Table of All Behavioral and Neural Variables

Right lOFC MedOFC Left frontal Thalamus MedOFC Engaged Analytical

Variable [OH] [OH] pole [OH] [ON] [ON] RTP Singing responding Achievement thinking Body Sadness

Right lOFC [OH] .43ⴱⴱ .79ⴱⴱ ⫺.22 ⫺.11 ⫺.40ⴱⴱ .31ⴱ .06 ⫺.22 .32ⴱ .05 ⫺.19
.00 .00 .17 .48 .01 .05 .69 .17 .04 .74 .22
Med OFC [OH] .43ⴱⴱ .47ⴱⴱ ⫺.18 ⫺.10 ⫺.32ⴱ .23 .01 ⫺.08 .16 .04 .21
.00 .00 .25 .51 .04 .14 .94 .60 .31 .81 .18
Left frontal pole [OH] .79 .47ⴱⴱ ⫺.27 ⫺.12 ⫺.32ⴱ .32ⴱ .18 ⫺.31ⴱ .23 .11 ⫺.04
.00 .00 .08 .43 .04 .04 .25 .05 .15 .51 .82
Thalamus [ON] ⫺.22 ⫺.18 ⫺.27 .11 .06 ⫺.13 ⫺.19 .08 ⫺.22 ⫺.09 .08
.17 .25 .08 .47 .71 .42 .24 .62 .17 .59 .64
MedOFC [ON] ⫺.11 ⫺.10 ⫺.12 .11 .36ⴱ ⫺.29 ⫺.12 .30 ⫺.30 .00 .03
.48 .51 .43 .47 .02 .06 .43 .06 .06 .98 .84
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RTP ⫺.40ⴱⴱ ⫺.32ⴱ ⫺.32ⴱ .06 .36ⴱ ⫺.14 .05 ⫺.03 ⫺.20 .01 .10
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.01 .04 .04 .71 .02 .33 .71 .84 .15 .97 .46
Singing .31ⴱ .23 .32ⴱ ⫺.13 ⫺.29 ⫺.14 .46ⴱⴱ ⫺.27 .18 .45ⴱⴱ .08
.05 .14 .04 .42 .06 .33 .00 .05 .20 .00 .57
Engaged responding .06 .01 .18 ⫺.19 ⫺.13 .05 .46ⴱⴱ ⫺.22 .14 .25 .04
.69 .94 .25 .24 .43 .71 .00 .12 .33 .08 .75
Achievement ⫺.22 ⫺.09 ⫺.31ⴱ .08 .30 ⫺.03 ⫺.27 ⫺.22 ⫺.33ⴱ ⫺.11 ⫺.09
.17 .60 .05 .62 .06 .84 .05 .12 .02 .44 .52
Analytical thinking .32ⴱ .16 .23 ⫺.22 ⫺.30 ⫺.20 .18 .14 ⫺.33ⴱ .32ⴱ ⫺.24
.05 .31 .15 .17 .06 .15 .20 .33 .02 .02 .09
Body .05 .04 .11 ⫺.09 .00 .01 .45ⴱⴱ .25 ⫺.11 .32ⴱ .01
.74 .81 .51 .60 .98 .97 .00 .08 .44 .02 .94
Sadness ⫺.19 .22 ⫺.04 .08 .03 .10 .08 .04 ⫺.09 ⫺.24 .01
.23 .18 .82 .64 .84 .46 .57 .75 .52 .09 .94
ⴱ ⴱⴱ
Correlation is significant at the .05 level (2-tailed). Correlation is significant at the .01 level (2-tailed).

whether this has any bearing on their emotional interactions. A icantly more engaged and responsive to their daughters. The latter
second possible interpretation of these data is that fathers of sons finding is germane to the research on gender socializations of
may find more reward in interactions that include ambiguous empathy, suggesting that fathers of daughters are more present and
emotions, a characteristic of RTP. A longitudinal study found that attentive, as opposed to distracted, during interpersonal dyadic
new fathers experienced a decrease in gray matter in the mOFC interactions. This may facilitate the development of increased
that correlated with paternal intrusiveness during a play session (P. empathy in girls. Unfortunately, while our more specific empathy
Kim et al., 2014). Although the decrease is surprising in light of variable also differed by gender, it proved difficult to code reliably
research reviewed above highlighting the importance of the mOFC from the recorded ambient sounds and future research may exam-
for parental responding, it is consistent with the idea that one role ine more precise real-world indicators of empathic parental behav-
of the mOFC is to support the social ambiguity of RTP. It may be ior. Examining the behavioral findings as a whole, an intriguing
that fathers of sons employ more reward-related activity in re- hypothesis is that there are multiple routes for augmenting the
sponse to neutral facial expressions, which then supports their development of empathy and emotion regulation, with fathers of
engagement in RTP outside of the laboratory. daughters using more emotion language and greater attentional
In addition to differences in neural responding, fathers of daugh- engagement to emotions, and fathers of sons engaging in more
ters and sons differed in behavioral and linguistic variables. The RTP. Future research can examine the possibility that fathers
finding that fathers engage in more RTP with sons is consistent engage in differing approaches to foster emotion regulation, and
with a large body or prior research (Fletcher, StGeorge, & Free- can evaluate how children’s emotional and social outcomes fare as
man, 2013; Lewis, 1997; McIntyre & Edwards, 2009; Paquette & a result.
Dumont, 2013; Pellegrini & Smith, 1998). RTP involves dynamic Perhaps the most novel observation in these data were gender
and forceful behaviors, such as tickling, poking, and tumbling, differences in linguistic patterns that were not predicted a priori.
which would be hostile in many circumstances and which can only While fathers of sons use more language related to achievement
be interpreted as play given the particular social context. For this (e.g., “top,” “win,” “proud”), fathers of daughters use more ana-
reason, it is thought to both require and entrain emotion regulation lytical language, which a previous study linked with future aca-
and empathy. Research across many mammalian species attests to demic success (Pennebaker, Chung, Frazee, Lavergne, & Beaver,
the importance of RTP for social competence and brain develop- 2014). Previous research highlights the importance of paternal use
ment (Bell, McCaffrey, Forgie, Kolb, & Pellis, 2009; MacDonald of language with young children for shaping later verbal skills,
& Parke, 1984). In humans, paternal rough and tumble play is even after controlling for other aspects of quality child care (Panc-
positively correlated with children’s popularity with their peers sofar & Vernon-Feagans, 2006). Taken together then, if the results
(MacDonald & Parke, 1984). hold in a future replication, the current findings suggest that gender
Whereas fathers engaged in more RTP with sons, fathers of differences in fathers’ language may relate to future cognitive
daughters engaged in more whistling and singing and were signif- outcomes in meaningful ways and may help explain the consistent
 270 MASCARO, RENTSCHER, HACKETT, MEHL, AND RILLING

finding that girls outperform boys in school achievement outcomes gendered behavior, often obfuscated by social desirability (Mehl,
in countries committed to equal education opportunities (Epstein, in press).
1998; Spinath, Eckert, & Steinmayr, 2014). Although we have primarily interpreted these findings in the
Additionally, fathers of daughters used more language referenc- context of gender socialization, it is crucial to consider the possi-
ing the body (e.g., “belly,” “foot,” “tummy”) relative to fathers of bility that gender differences in paternal behavior may be the result
sons. This finding is relevant to research indicating that preado- of fathers responding to differential cues from the children, some
lescent girls are more likely to report body dissatisfaction (Wood, of which may be highly influenced by biological sex differences
Becker, & Thompson, 1996) as well as more negative effects of (e.g., hormone or neurobiological differences between male and
body dissatisfaction on self-esteem (Furnham, Badmin, & Sneade, female children). Males are exposed to higher levels of fetal
2002). Other lines of research shows that body stigmatism begins testosterone, which plays an important role in brain organization,
in early childhood (e.g., (Cramer & Steinwert, 1998) and that and there is literature causally linking fetal androgen exposure to
parents exert an important influence on young children’s body rough and tumble play (Collaer & Hines, 1995; Pasterski et al.,
image (Tatangelo, McCabe, Mellor, & Mealey, 2016). The EAR 2011). Not only does fetal androgen exposure influence play style,
This article is intended solely for the personal use of the individual user and is not to be disseminated broadly.

methodology may be useful for better understanding how parental but there is some evidence that it influences playmate selection
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language about the body influences child body image and whether such that both boys and girls exposed to higher levels of fetal T are
the gender of the child moderates this link. more likely to choose a playmate with a stereotypically masculine
The finding that fathers of girls used more sadness language play style (Pasterski et al., 2011). Further support for the notion that
when talking to their child is consistent with previous studies in fathers may alter their behavior in response to their toddler’s cues
which parents report more acceptance of girls’ sadness (Birnbaum comes from a study indicating that fathers decrease their levels of
& Croll, 1984) and attend more to girls’ sadness and anxiety physical play if their child is high on internalizing behavior (Jia,
(Chaplin et al., 2005). This gender difference has real health Kotila, & Schoppe-Sullivan, 2012). The authors interpreted their
consequences. Across diverse lines of research, restrictive emo- findings as reflecting fathers’ adaptation to children that respond
tionality in adult men is linked to marital dissatisfaction and negatively to play. In combination these findings lend support for the
depression (Shepard, 2002) and interferes with social intimacy possibility that the male children of the fathers in our sample may
“pull” RTP from fathers and highlights the important point that
(Sharpe & Heppner, 1991) and mental health help-seeking behav-
father-child interactions take place in a complex micro- and macroen-
ior (Robertson & Fitzgerald, 1992). Our findings using unobtru-
vironment (Cabrera, Fitzgerald, Bradley, & Roggman, 2014).
sive real-world observational assessments of language reveal that
Other limitations warrant mention. First, this was an exploratory
gender differences in emotion language begin early, in particular in
study with a relatively short EAR monitoring period. Although
the extent to which fathers use sadness language. We cannot
these data are far more comprehensive than a laboratory assess-
determine the functional implications of this language difference,
ment, they were acquired over a period of two days (one weekend
but future research can test the hypothesis that fathers’ reduced use
day and one weekday) and it is possible that for some fathers the
of sadness language with sons impedes the socioemotional learn-
study period was not accurately representative of their typical
ing so integral for positive outcomes. Nor can we rule out the
behaviors and interactions. Related, the EAR data are acoustic
possibility that daughters somehow solicited sadness language
observations only, and as such visual cues were not available for
more than sons, a real possibility given previous findings that very behavioral coding. This may be particularly important with respect
young girls use more emotion language than boys (e.g., (Cervantes to our assessments of paternal affective behavior, and our assess-
& Callanan, 1998). An intriguing possibility is that there exists a ment of empathy may have suffered from this limitation. In addi-
positive feedback loop in which caregivers use more emotion tion, this was a wide-ranging exploratory analysis and only the
language with girls, which then leads girls to use and solicit gender difference related to singing survived Bonferroni correction
emotion language with their intimate caregivers. for multiple comparisons. Several of our findings were not predicted
The present study establishes a novel method for naturalistically a priori and should be replicated in future research. Moreover, we
observing and assessing meaningful parenting behaviors, and can were not able to detect any neural responses that significantly medi-
be used in the future to more thoroughly investigate the biological ated the link between child gender and paternal behavior. Although
bases of paternal and maternal caregiving. An important strength paternal brain responses to the photograph stimuli were relatively
of the EAR methodology is that we can examine whether language robust, it may be that the neural response to the child’s dynamic
variables directed at romantic partners or coworkers also differs as behaviors in everyday life, more so than the neural response to still
a function of the sex of the child. None of the language variables photographs of the child in the laboratory environment, mediates the
identified here as differing between fathers of daughters and fa- relationship between child gender and paternal behavior. Finally,
thers of sons differed in the coworker or friend context, allowing although the sample size provided sufficient statistical power to detect
us to determine with confidence that gender differences in paternal the gender differences in brain function, it is relatively small for
language were specific to the domain of direct interactions with the studies examining parental behavior and was likely underpowered to
child. Fathers had an average of 150 sound files, which equates to detect mediation effects of brain function on behavior.
more than two hours of data spread over 2 days. Compared with a Despite inherent limitations, the present findings advance our
15-min play session in a laboratory setting, the EAR allows for a understanding of the neural bases of paternal caregiving by un-
more thorough evaluation of patterns of paternal involvement and covering neural systems that differentially respond to daughters
the brain activity that support them. Moreover, as an unobtrusive versus sons and how those neural responses, in turn, may contrib-
observation method, the EAR is not beset with self-report biases, ute to gendered paternal caregiving behaviors. Additionally, the
and may be particularly important for examining questions, such as use of the EAR revealed differences in paternal behavior and
 CHILD GENDER AND PATERNAL BEHAVIOR 271

language use that may relate to child social, emotional, and cog- major depression. PLoS ONE, 5, e15058. [Link]
nitive outcomes. Future research can employ this methodology for [Link].0015058
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