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Predictors and outcomes across the transition to fatherhood

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Predictors and outcomes across the transition to fatherhood

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Predictors and Outcomes Across the Transition to Fatherhood
Hannah Khoddam

A Dissertation Presented to the
FACULTY OF THE USC GRADUATE SCHOOL
UNIVERSITY OF SOUTHERN CALIFORNIA
In Partial Fulfillment of the
Requirements for the Degree
DOCTOR OF PHILOSOPHY
(PSYCHOLOGY)

August 2020

Copyright (2020) Hannah Khoddam

i
Acknowledgments
The research described in this dissertation was supported by the Eunice Kennedy Shriver
National Institute of Child Health and Human Development (Grant F31 HD093107-01A1; PI:
Hannah Lyden) and by the National Science Foundation (NSF) Career Award # 1552452 (PI:
Darby Saxbe, PhD). I am so grateful to have been awarded the time and money to be able to
answer the important questions addressed in this dissertation.
I am so grateful to my dissertation committee, Dr. Darby Saxbe, Dr. Gayla Margolin, Dr.
Jonas Kaplan, Dr. Richard John, and Dr. Lynn Miller who have been a part of this journey since
the conception of this project, and who have all been available at different times to help me,
challenge me, and aid in my growth as a clinical scientist. I am forever grateful to the
Neuroendocrinology of Social Ties (NEST) lab, for their support and friendship. Specifically, the
graduate students and lab managers of the Nest lab, Sarah Stoycos, Geoff Corner, Mona Khaled,
Alyssa Morris, Narcis Marshall, Sofi Cardenas, Katie Horton, Bryna Tsai and Nia Barbee; who
were not only integral to data collection and analysis of this project but also to helping me stay
sane and enjoy the ride. I would also like to extend my gratitude to Dr. Diane Goldenberg, a
postdoctoral fellow in NEST, and Dr. Xiaofei Yang, a research scientist in the Brain and
Creativity Institute, who helped me on a variety of data analytic techniques and data collection
for this project.
I am also so grateful for my cohort for being by my side every step of the way on this
crazy journey. You ladies have become some of my best friends and I am so grateful to have
lived the last 7 years of life with you all.
To my dissertation chair Dr. Darby Saxbe, who I literally wouldn’t be here without, I’m
so grateful to you for taking a chance on me 7 years ago and for allowing me to be the first

ii
graduate student of the Nest Lab. It has been an honor working with you and watching the lab
and the HATCH study grow from just an idea to the successful and blossoming lab that it is
today. I am so grateful to have had your support and friendship over the last 7 years and owe so
much of my success to you.
I, of course, would be remiss to not acknowledge my family; the one I was born into, the
one that married into my family, and the one that I married into, for all of your continuous
support and love. I’m so grateful to my Dad who has always acted as a mentor and has taught me
so much about what it means to be a scientist and a practitioner. To my mom and my sister for
their endless support and for pretending to read the papers I send them, and for always offering
to pay for me over the last 7 years of having a student stipend.
Lastly, to my husband Rubin Khoddam, whom I would not have met if not for this degree
and without whom I would not have been able to finish this dissertation. You are truly a role
model and an inspiration for what it means to be a clinical scientist and a human being. Your
guidance and mentorship in this program and in life has been invaluable. I would not have had
the successes both in this life and in this program if it weren’t for you. And to our beautiful
daughter Sahren Rose. Although this dissertation has often felt like my baby, nothing in this
world will come close to being your mom. You are truly the best thing that has ever happened in
my life. Having you changed this exploration of parenting from an intellectual exercise to a real,
complicated, and extraordinary experience.

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Table of Contents
Acknowledgments ............................................................................................................................ i

List of Tables ..................................................................................................................................v

List of Figures................................................................................................................................ vi

Abstract ........................................................................................................................................ vii

General Introduction ................................................................................................................... viii

Manuscript 1: Characterizing the Prenatal Response to Infant Cry ................................................8
Abstract .......................................................................................................................................8
Introduction .................................................................................................................................9
Methods......................................................................................................................................16
Results ...............................................................................................................................21
Discussion ..........................................................................................................................23
References .........................................................................................................................27
Table 1 ...............................................................................................................................42
Table 2 ...............................................................................................................................43
Table 3 ...............................................................................................................................44
Table 4 ...............................................................................................................................45
Table 5 ...............................................................................................................................46
Figure 1 ..............................................................................................................................47
Figure 2. ............................................................................................................................48

Manuscript 2: Longitudinal Examination of the Response to Infant Cry ....................................49
Abstract .....................................................................................................................................49
Introduction ................................................................................................................................50
Methods ....................................................................................................................................54
Results .......................................................................................................................................58
Discussion ..................................................................................................................................59
References .................................................................................................................................62
Table 1 ...................................................................................................................................76
Table 2 ...................................................................................................................................77
Table 3 ...................................................................................................................................78
Table 4 ...................................................................................................................................79

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Manuscript 3: Prenatal Factors Predicting Postpartum Relationship Decline ..............................80
Abstract .....................................................................................................................................80
Introduction ................................................................................................................................81
Methods .....................................................................................................................................86
Results .......................................................................................................................................90
Discussion .................................................................................................................................94
References .................................................................................................................................98
Table 1 .................................................................................................................................106
Table 2 .................................................................................................................................107
Table 3 .................................................................................................................................108
Table 4 .................................................................................................................................109
Figure 1 ................................................................................................................................110
Figure 2 ................................................................................................................................111
Figure 3 ................................................................................................................................112
Figure 4 ................................................................................................................................113
Figure 5 ................................................................................................................................114
Figure 6 ................................................................................................................................115

General Discussion .....................................................................................................................116

References for General Introduction and Discussion ................................................................120

v
Tables

1.1 Means and standard deviations for study variables…………………………………...…42

1.2 Bivariate correlations of main study variables……………………………………….…..43

1.3 Neural activation during infant cry………………………………………………............44

1.4 Summary of Multiple Regression Analysis for variables predicting right and left
amygdala activation in infant cry compared to white noise……………...…………...…45

1.5 Associations Between neural activation on task and prenatal testosterone…..................46

2.1 Means and standard deviations for study variables………………………..……………76

2.2 Bivariate correlations for all study variables………………………………..……...…....77

2.3 Summary of multiple regression analysis for psychological response to infant
cry predicting postpartum outcomes..................................................................................78

2.4 Summary of multiple regression analysis for neural response to infant cry predicting
postpartum bonding ……………........…………………………………...…...…...…….79

3.1. Means and standard deviations for study variables…………………………………….106

3.2 Bivariate correlations of main study variables……………………………………….…107

3.3 Unrestricted model with a between-dyads moderator……………………...……… ….108

3.4 Actor-partner interdependence moderation submodels………………...……….……...109

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Figures

1.1 Main effects for the contrast of interest infant cry > frequency matched white
noise……………..………………………………………………………………….…....47

1.2 Neural activation to infant cry associated with baseline testosterone level…………...…48

3.1 The actor-partner interdependence model with a between-dyads moderator…………..110

3.2 APIM examining couple prenatal sexual satisfaction on changes in relationship
satisfaction pre to postpartum……………………………….……………………........111

3.3 Path model examining the relationship between prenatal T and change in
relationship satisfaction pre to postpartum…………………………....……............….112

3.4 APIMOM examining fathers’ and mothers’ prenatal T as a moderator of prenatal sexual
satisfaction and relationship change………………………………………………........113

3.5 Actor and partner effects of prenatal sexual satisfaction ………………...................….114

3.6 Simple slopes analyses for APIMOM results……………………………….....……….115

vii
Abstract
The transition to fatherhood is a complex biological and psychological process with
myriad neuroendocrine changes which prepare the father for sensitive and effective parenting.
Biological mothers have nine months of physical and emotional changes to prepare for the
impending child, but when and how this process unfolds in biological fathers remains
understudied. Understanding how a father becomes a father, and which biological and
psychological factors contribute to this evolution, is an important first step in creating targeted
interventions to bolster father’s parenting efforts. Collectively, the included set of studies will
add substantially to our understanding of the prenatal psychological and physical factors
associated with the transition to fatherhood. Results from these studies may pinpoint important
prenatal variables in predicting difficulties in the co-parenting relationship and the father-child
relationship post birth and act as a first step in creating effective interventions for paternal
readiness.

1
General Introduction
The Transition to Fatherhood
The transition to fatherhood is a complex biological and psychological process with
myriad neuroendocrine changes which prepare the father for sensitive and effective parenting.
Biological mothers have nine months of physical and emotional changes to prepare for the
impending child, but when and how this process unfolds in biological fathers remains
understudied. Recent studies suggest that fathers are spending more time with their children than
any previous generation (Petts et al., 2018), and that sensitive and available fathers are important
for the child’s development and achievements (Wilson, Hansen, & Li, 2011). Therefore,
understanding how a father becomes a father, and which biological and psychological factors
contribute to this evolution, is an important first step in creating targeted interventions to bolster
father’s parenting efforts.
Hormonal and behavioral changes are hypothesized to ready the new father’s body and
mind to become a parent. Testosterone (T) is one such hormone that may change across the
transition to fatherhood and is implicated in paternal readiness and commitment to the parenting
relationship (Gettler et al., 2011). Another marker of parenting behavior that has been widely
studied (Bakermans-Kranenburg et al., 2012a; Compier-de Block et al., 2015; Crouch et al.,
2008; Joosen, et al., 2013; Riem et al., 2012; Rodriguez et al., 2015), reactivity to infant cry
sounds, has been linked with both sensitive and aggressive parenting. These responses to infant
cry include self-report of one’s interpretation of the infant (e.g. hostile, happy, sad etc.),
emotions felt while listening to the infant crying, as well as physiological responses such as skin
conductance, heart rate, neural response and behavioral responses. Studies of both perinatal
hormones and infant cry responses, however, rarely include men, and even fewer have included

2
expectant fathers. The prenatal period is a vitally important time wherein parent’s bodies, minds,
and relationships may be studied and potentially targeted for interventions designed to improve
parenting readiness. Additionally, parents’ reactivity to infant cry in this prenatal period might
predict a broad range of postpartum parenting outcomes, from sensitive to harsh or insensitive
parenting, as well as difficulties bonding with the infant (Oldbury & Adams, 2015). More
nuanced indicators of parenting risk, such as parenting stress or distressed bonding, have rarely
been investigated in regards to infant cry reactivity, and yet have important implications for child
development (Magill-Evans & Harrison, 2001; Molfese et al., 2010). The following studies are
an attempt to understand the prenatal variables that may be important in the eventual relationship
between the biological father and the child, as well as the biological father and mother, and act as
a first step in building interventions for paternal readiness.
Testosterone
Testosterone (T) is an androgenic steroid hormone that has been implicated in the
transition to fatherhood. Studies suggest that over time when men become partnered and when
they become fathers, T decreases, leading to more commitment to the co-parenting relationship
and more sensitive parenting (Gettler et al., 2011; Gray et al., 2007). However, higher T also
leads to more competitive mating advantages, creating a tradeoff between the benefits of high T
when finding a mate and low T when becoming a partnered father (Wingfield et al., 1990;
Wingfield, 2017). Additionally, although baseline T levels tend to decrease once men become
partnered fathers over time, T also increases acutely during parenting situations in which
children are crying and need protecting or nurturing (Van Anders et al., 2012). Therefore,
context may also be important in determining the significance of T in fathering. T reactivity

3
versus baseline T measurements may lead to different outcomes, and T in the context of an infant
crying may be important to explore.
Some studies have measured T in pregnant couples and found that fathers with steeper
declines in T during the prenatal period report higher postpartum paternal involvement and
investment in their relationship with their partner (Gray et al., 2007; Saxbe et al., 2017).
However, questions remain about the significance of T in the transition to parenthood. For
example, it remains unclear whether men’s T is associated with their relationship with the
biological mother of their child. It may be that men and women’s T coregulate during pregnancy
and that greater coregulation leads to better relationship outcomes postpartum (Saxbe et al.,
2017). However, the sexual relationship between expectant parents has never been investigated
as a potential and important indicator of relationship satisfaction that may be linked with T,
particularly during the transition to parenthood. This appears to be a notable gap given the
importance of T in sexuality, specifically in males, as well as the importance of T in the
transition to fatherhood. Although higher T is purported to increase successful mating challenges
(i.e., likelihood of fathering a child) in mammalian males, including humans (Wingfield et al.,
1990), research focusing on naturally occurring T in healthy men has often failed to find a
relationship between high T and more frequent sexual behaviors or increased sexual desire
(Brown et al., 1978; Persky et al., 1978; Raboch & Stárka, 1973; van Anders, 2013; Van Anders,
2012). Research on changes in sexual behavior or sexual desire across the transition to
parenthood is scarce, although studies suggest that changes in the sexual relationship in new
parents is common (Ahlborg et al., 2009; Condon et al., 2004; Pastore et al., 2007) and T levels
may interact with these changes (Gettler et al., 2013). Therefore, it is an intriguing but thus far

4
untested possibility that T levels in men before the birth of a child as well as sexual satisfaction
in their relationship may reflect relationship satisfaction across the transition to parenthood.
Infant Cry
Sensitive and prompt responding to an infant crying is important for the survival of the
infant. However, listening to an infant crying can cause stress and irritation and even aggression
toward the infant, especially in a new parent who may lack parenting experience. Therefore, how
a parent responds to a crying infant has vitally important implications for the child and for the
parent-child relationship. Studies have shown that physiological arousal to infant cry (i.e. greater
skin conductance, greater handgrip force), negative interpretations of the crying infant, and
activation in certain brain areas while listening to infant cry may be associated with risk along a
continuum with child abuse at the most extreme end, and less severe but widely experienced
outcomes, such as parenting stress and bonding difficulties, at the more normative end (Compier-
de Block et al., 2015; Crouch et al., 2008; Oldbury & Adams, 2015; Rodriguez & Green, 1997;
Witteman et al., 2019). These physiological, psychological, and neural studies of the response to
infant cry, however, seldom include fathers. Studies including new fathers have shown similar
neural responses to infant cry as new mothers, with activation in the bilateral auditory cortex,
dorsomedial prefrontal cortex, fronto-insular cortex, thalamocingulate circuits and midbrain
dopaminergic regions (Li et al., 2018). Few studies have investigated the implications of this
neural arousal, nor investigated multi-modal responses to infant cry (Bakermans-Kranenburg et
al., 2012b). However, one study found that greater activation in auditory cortices to infant cry in
new fathers was associated with rating the infant cry as more aversive (Li et al., 2018), whereas
another study of new mothers found no relationship between neural responses to infant cry and

5
mothers’ irritation with infant cry sounds. Therefore, questions remain about how the responses
to infant cry track together and influence the expectant father’s eventual parenting behavior.
Although a small, preliminary literature suggests that fathers may respond to infant cry
similarly to mothers, that these responses may have implications for future parenting, and
testosterone may be important for readying the man for fatherhood, many gaps in the literature
exist. Firstly, although physiological, behavioral, and psychological responses to infant cry are
assumed to correspond, few studies have measured each of these modalities in the same group of
expectant fathers to understand how they track together. Secondly, few longitudinal studies exist
investigating the real-world parenting implications of prenatal responses to infant cry. Thirdly, it
remains unclear how testosterone levels during the transition to fatherhood may interact with the
response to infant cry, specifically during the prenatal period during which testosterone may play
an important role in readying the man for fatherhood. Lastly, the role of prenatal factors such as
sexual satisfaction and testosterone on romantic relationship changes during the transition to
parenthood remains poorly understood.
Aims of Overall Project and Focus of Individual Manuscripts
The current set of studies aims to address these gaps. The Hormones Across the
Transition to Childrearing (HATCH) study follows pregnant couples from mid-to-late pregnancy
to six months postpartum to assess the biological and psychological processes involved in
becoming a parent. The three studies included within this dissertation focus on the prenatal
factors (such as testosterone level, sexual satisfaction ratings, and multimodal responses to infant
cry) that may predict postpartum family relationship functioning. To our knowledge, no studies
have prospectively investigated these questions in expectant fathers followed from the prenatal to
postpartum period. Three specific aims guided the inquiry for this project:

6
Specific Aim 1: To characterize the neurobiological, psychological, and behavioral response to
infant cry in expectant fathers, and test the moderating role of testosterone in these responses.
Specific Aim 2: To test the relationship between reactivity to infant cry and parenting outcomes
in new fathers at three months postpartum.
Specific Aim 3: To investigate measurable prenatal factors (i.e. testosterone, sexual satisfaction)
that may predict postpartum relationship functioning in new mothers and fathers.
Manuscript 1: Characterizing the Prenatal Response to Infant Cry in Expectant Fathers
and the Role of Testosterone
The first study, “How Do Expectant Fathers Respond to Infant Cry? Examining Brain
and Behavioral Responses and the Moderating Role of Testosterone” (in press in Social
Cognitive and Affective Neuroscience) is the first study to characterize the neural, behavioral,
and psychological responses to infant cry in expectant fathers and examine the role of
testosterone in these responses. This study utilized novel physiological techniques (i.e. handgrip
dynamometer measurement), fMRI, and self-report measures of experiences of the crying infant,
to understand how expectant fathers respond to infant cry before the birth of their child and
establish how these responses may track together.
Manuscript 2: Longitudinal Examination of Prenatal Response to Infant Cry in Predicting
Postpartum Parenting Outcomes in Expectant Fathers.
The second study, “Prenatal Responses to Infant Cry and Postpartum Parenting Outcomes
in Expectant Fathers,” builds on Study 1’s goals by examining the predictive value of expectant
fathers’ responses to infant cry in postpartum parenting stress and bonding with the infant. This
is the first study that evaluates measurable prenatal factors (i.e. response to infant cry) in

7
predicting potential difficulties with parenting postpartum which may be important in crafting
future interventions to increase bonding between father and child.
Manuscript 3: Prenatal Factors Predicting Postpartum Relationship Outcomes Across the
Transition to Parenthood
The third and final study, “The Moderating Role of Testosterone on Sexual Satisfaction
and Marital Satisfaction Across the Transition to Parenthood,” (in revision at Hormones &
Behavior) examines how prenatal testosterone levels in expectant fathers and prenatal sexual
satisfaction in both expectant mothers and fathers may predict changes in relationship
satisfaction (in both mothers and fathers) across the transition to parenthood, as well as test the
moderating role of testosterone. Given the dyadic nature of parenting data, this study utilizes
path models, Actor-Partner Interdependence Models (APIM; Kenny, 1996; Kenny & Judd,
1986), and Actor-Partner Interdependence Moderation Models (APIMoM; Garcia et al., 2015).
APIM models account for the interdependence of dyadic data and allows for simultaneous,
independent estimation of the impact of a person’s characteristics on their own outcomes, actor
effects, and the impact of a person’s characteristics on their partner’s outcomes, partner effects.
To our knowledge this is the first study to employ dyadic data to examine how prenatal
testosterone and sexual satisfaction predict the potential decline in relationship satisfaction after
the birth of a child.
Collectively, these studies will add substantially to our understanding of the prenatal
psychological and physical factors associated with the transition to fatherhood. Results from
these studies may pinpoint important prenatal variables in predicting difficulties in the co-
parenting relationship and the father-child relationship post birth and act as a first step in creating
effective interventions for paternal readiness.

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How Do Expectant Fathers Respond to Infant Cry? Examining Brain and
Behavioral Responses and the Moderating Role of Testosterone
Hannah Khoddam, Diane Goldenberg, Sarah A. Stoycos, Katelyn Taline Horton, Narcis
Marshall, Sofia Cardenas, Jonas Kaplan and Darby Saxbe

In Press in Social Cognitive and Affective Neuroscience
Abstract

Expectant parents’ responses to infant cry may indicate future risk and resiliency in the
parent-child relationship. Most studies of parental reactivity to infant cry have focused on
mothers, and few studies have focused on expectant fathers, although fathers make important
contributions to parenting. Additionally, although different responses to infant cry (behavioral,
psychological, and neural) are hypothesized to track together, few studies have analyzed them
concurrently. The current investigation aimed to address these gaps by characterizing multimodal
responses to infant cry within expectant fathers and testing whether prenatal testosterone
moderates these responses. Expectant fathers responded to infant cry versus frequency-matched
white noise with increased activation in bilateral areas of the temporal lobe involved in
processing speech sounds and social and emotional stimuli. Handgrip force, which has been used
to measure parents’ reactivity to cry sounds in previous studies, did not differentiate cry from
white noise within this sample. Expectant fathers with higher prenatal testosterone showed
greater activation in the supramarginal gyrus, left occipital lobe and precuneus cortex to cry
sounds. Expectant fathers appear to interpret and process infant cry as a meaningful speech
sound and social cue, and testosterone may play a role in expectant fathers’ response to infant
cry.

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Introduction
An infant’s survival depends on the caregiving relationship. Caregivers who respond
sensitively to their infant’s needs can facilitate their child’s healthy development (Ainsworth,
1979; Malmberg et al., 2016). An infant’s primary form of communication, crying, may arouse
the parent to respond and attend to an infant’s distress (Brosch et al., 2007). Researchers have
studied endocrine, behavioral, and, more recently, neural responses to infant cry, finding that
hormone levels, handgrip strength, and patterns of brain activation may vary while listening to
infant cries (Crouch et al., 2008; Fleming et al., 2002; Kim et al., 2010). Previous studies have
suggested that calm parental responses to cry may be linked with approach-oriented, sensitive
responses to infants (e.g. Joosen et al., 2013), whereas irritated or hyperreactive responses may
be linked with risk for aggression or neglect (Reijman et al., 2014; Zeifman & St. James-Roberts,
2017).
Despite these intriguing findings, the literature on reactivity to infant cry has several
notable gaps. Surprisingly, given that fathers often participate in infant caregiving, few studies
have investigated responses to infant cry sounds in fathers and even fewer have included
expectant fathers (Alyousefi-Van Dijk et al., 2019; Thijssen et al., 2018). Interventions have
targeted parents’ possible responses to infant cry as a marker of aggressive responses to a crying
infant (Barr et al., 2009; Coster, 2017). However, it is imperative to thoroughly study expectant
fathers’ responses to infant cry in order to best tailor these interventions specifically before the
baby even arrives. Men’s hormones prior to the birth of their child (e.g. their testosterone levels
during their partner’s pregnancy) may also reflect their preparation for parenting (Saxbe et al.,
2017), and also warrant examination in conjunction with expectant fathers’ responses to infant
cry. Lastly, although responses across different measurement domains, such as brain and

10
behavioral responses, are assumed to relate to one another (Messina et al., 2016), few studies
have measured multiple responses to cry sounds within the same population and analyzed them
concurrently (e.g. Bakermans-Kranenburg et al., 2012). Understanding how behavior works in
tandem with the brain will help to clarify an overarching theory of the biological response to
infant cry and identify where this system may go awry in aggressive or abusive parenting
responses.
The current investigation aims to address these gaps by characterizing behavioral,
psychological, and neural responses to infant cry compared to frequency-matched white noise
within expectant fathers and by testing whether paternal prenatal testosterone, a hormone that
may reflect paternal investment, affects reactivity to infant cry.
Neural Responses to Infant Cry
Studies of parental brain responses to infant cry have focused primarily on mothers, with
few studies on fathers and even fewer investigating expectant fathers (for review see: Abraham
& Feldman, 2018; Feldman, 2015; Lynch, 2003; Rilling, & Young., 2014; Thijssen et al., 2018.;
Witteman et al., 2019). These studies have investigated how parents respond to own infant cry
versus unfamiliar infant cry, or an unfamiliar infant cry compared to a control sound. Infant cry
sounds, in comparison to video or picture stimuli of infants, have been linked with amygdala
activation in mothers, but not fathers or nonparents (Feldman, 2015). Recruitment of the
amygdala may underscore parental vigilance to infant distress cues (Abraham et al., 2014).
Studies have found that mothers listening to an unknown infant compared to a frequency-
matched white noise show activation in components of the midbrain dopamine system (i.e.
substantia nigra, and ventral tegmental area), anterior and posterior cingulate cortex, right fronto-
insular cortex, dorsomedial prefrontal cortex (dMPFC) (regions involved with emotional and

11
cognitive empathy), and right lateralized auditory cortices extending to the temporal pole
(Lorberbaum et al., 1998). A study of neural reactivity in first-time fathers listening to unknown
infant cry compared to a white noise control found bilateral activations in the medial prefrontal
cortex, bilateral anterior insula and inferior frontal gyrus (IFG), bilateral striatum, bilateral
thalamus, bilateral auditory cortex (including the planum temporale, Heschl’s gyrus, and
supramarginal gyrus) bilateral posterior cingulate, and bilateral midbrain structures (Li et al.,
2018). No differences were found between own infant cry and unfamiliar infant cry in these
first-time fathers. Similarly, a study of the effects of vasopressin on processing infant cry sounds
in expectant fathers found that infant crying (vs control sounds) was associated with increased
activation in the bilateral auditory cortex and posterior medial cortex (Thijssen et al., 2018). First
time fathers, similarly to first time mothers, appear to engage five neural systems while listening
to unknown infant cries: (1) auditory cortex (auditory perception), (2) dorsomedial prefrontal
cortex (perspective-taking, theory of mind); (3) fronto-insular cortex (emotional empathy), (4)
thalamocingulate circuits (parental caregiving), and (5) midbrain dopaminergic regions
(approach motivation). A recent meta-analysis similarly confirmed involvement of the cingulate,
the auditory system, the pre-supplementary motor area, the dorsal anterior insula, the
dorsomedial prefrontal cortex, and the inferior frontal gyrus in infant cry perception as well as
larger activations in the right IFG, temporal pole, and left angular gyrus in men in response to
infant cry compared to women (Witteman et al., 2019).
Handgrip Modulation Response to Infant Cry
Modulation of handgrip force is interpreted as a behavioral indicator of motivation to
respond to the distressed infant (Crouch et al., 2008; Riem et al., 2012; Zeifman & St James-
Roberts, 2017). The ability to modulate handgrip force is measured by a dynamometer, which

12
tracks grip strength while an individual listens to an infant crying. Studies have suggested that
excessive handgrip force, referred to as ‘poor modulation,’ may indicate risk for abuse or
aggressive responding toward an infant (Bakermans-Kranenburg et al., 2012b). Poorly
modulated handgrip response to infant cry has been linked with neglectful and physically
abusive mothers (Compier-de Block et al., 2015) and with parents at risk for child physical abuse
(Crouch et al., 2008). Other studies suggest that poor modulation of handgrip may be an
indicator of motivation to act or help while an infant is crying (Parsons et al., 2013). One study
of expectant fathers failed to find a difference in handgrip modulation between infant cry and
frequency-matched white noise control sounds (Alyousefi-Van Dijk et al., 2019). However, this
study involved vasopressin administration and viewing pictures of infant faces while listening to
infant cry. Earlier studies of infant cry and handgrip force also rarely utilized control sound
stimuli, making it difficult to assess whether poor modulation is tied specifically to infant cry or
more broadly to any distressing sound. The current study builds on previous studies in examining
the response to infant cry in expectant fathers, while also using a control condition of frequency
matched white noise.
Psychological Response to Infant Cry
In addition to neural and behavioral responses to infant cry, researchers have investigated
individual differences in parents’ self-reported interpretations of infant cry. Interpretations of a
crying infant as intentionally hostile or reporting increased frustration and negative emotion
while listening to infant cry have been associated with risk for aggressive or harsh parenting
behaviors (Crouch et al., 2008; Rodriguez et al., 2015). However, the question of whether these
interpretations are associated with neural or behavioral responses to cry sounds is underexplored.
Parents who rate infant cry sounds as more hostile may show more excessive handgrip force

13
while listening to infant cry (Crouch et al., 2008). Similarly, one study found that fathers who
rated infant cry as more aversive exhibited greater neural activation in auditory cortices (Li et al.,
2018). Li and authors interpreted this greater neural activation as reflecting a form of negative
emotional over-arousal in response to infant cry sounds. However, another study found no
relationship between mothers’ irritation with infant cry sounds and their neural responses to the
same sounds (Riem et al., 2012). In sum, this literature is small and inconclusive, and has
generally not included expectant fathers.
The Role of Testosterone
Many recent studies have investigated the neuroendocrine underpinnings of parenting
(Bos, 2017; Bos et al., 2010). Testosterone (T) appears to decline across the transition to
parenthood in men and may be associated with paternal sensitivity and involvement in childcare
(Gettler et al., 2011; Saxbe et al., 2017; Storey & Ziegler, 2016). Infant cry can elicit sensitive
caregiving (Murray, 1985) or frustration and annoyance (Barr et al., 2006; Del Vecchio et al.,
2009; Frodi, 1985). Some parents may be physiologically overly responsive to noxious child
stimuli, such as infant cry (Knutson, 1978). This hyperreactivity can lead to an increase in
“irritable aggression,” which may reflect heightened parenting stress and compromised parent-
child bonding.
Higher levels of T around the transition to parenthood may indicate the potential for
harsh or insensitive parenting, particularly when coupled with this hyperreactivity to infant cry.
The role of T in modulating paternal responses to infant cry, however, has received little
attention, with most studies focusing on T reactivity to infant stimuli rather than baseline levels
of T across the transition to parenthood. One of the only studies to investigate baseline levels of
T in first-time fathers (postpartum) failed to find a relationship between baseline T levels and

14
neural activation differences in response to infant cry versus frequency-matched white noise (Li
et al., 2018). In studies investigating T reactivity to infant cry, it has been found that men with
higher T reactivity (non-fathers and new fathers) in response to unfamiliar infant cries show less
sympathy for these infant cries (Fleming et al., 2002). Another study of reactivity to infant cry
video stimuli found greater activation in the left caudate in fathers whose T increased more after
interacting with their child (Kuo et al., 2012). These authors suggest that increased T and greater
neural activation may indicate the body readying itself to protect the baby as signaled by the
urgent cries (Kuo et al., 2012). Additionally, another study found that infant cries from a baby
doll decreased T levels when the father was allowed to care for the infant but increased T levels
when the father was blocked from nurturing the infant (Van Anders et al., 2012), while another
found lower T after interacting with an infant in fathers with low cortisol levels (Bos et al.,
2018). Therefore, context may be important for understanding the relationship between T and
infant cry reactivity in fathers and increased T levels (both baseline and reactivity) may indicate
a physiological hypereactivity to infant cry and be associated with other potential hyperactive
responses to infant cry such as neural activity and handgrip modulation. Notably, T level appears
to be positively associated with handgrip strength in men (Gallup et al., 2010), but the
relationship between T and handgrip modulation has not been thoroughly tested in the context of
infant cry. Few studies have investigated T levels in expectant fathers before the birth of their
child (prenatal T) and its potential role in reactivity to infant cry.
Current Study
Although neural, behavioral, and psychological responses to infant cry have been
examined separately in previous studies, multi-modal approaches are needed to characterize how
and whether these responses are correlated across domains. Moreover, understanding how

15
expectant fathers respond to infant cry, and the role of T in shaping these responses, might
elucidate how fathers transition to parenthood and prepare to care for their infants.
Within a sample of fathers expecting their first child, we tested four hypotheses:
1) In response to infant cry sounds (vs. white noise), we expected that expectant fathers
would show greater neural activation in regions that have been associated with infant cry
specifically (e.g. socio-cognitive areas such as the STG, insula, mPFC, dlPFC, auditory cortices
and IFG). We planned to use whole-brain analyses to test this hypothesis, and to supplement
these analyses with an a priori ROI focused on the amygdala.
2) We also expected fathers to show behavioral responses to infant cry sounds,
specifically poor handgrip modulation when listening to infant cry sounds compared with white
noise sounds.
3) We expected that fathers’ responses to infant cry would be consistent across neural,
behavioral, and self-report modalities. Specifically, given evidence that fathers who rated infant
cry more negatively also showed heightened neural activation to cry (Li et al., 2018), we
expected that fathers who showed greater neural activation to infant cry in hypothesized brain
areas would also show poorer handgrip modulation, greater interpretation of the infant as more
negative during infant cry, and greater self-reported negative emotions after infant cry compared
to white noise.
4) Given that prenatal T may reflect paternal investment in sensitive parenting, which
requires the ability to modulate negative responses to aversive stimuli such as infant cry, we
hypothesized that fathers with higher prenatal T levels would show more reactivity to infant cry,
including more negative ratings of the infant, more negative emotions after listening to infant
cry, poorer handgrip modulation, and more activation in hypothesized brain areas in response to

16
infant cry.
Methods
Participants
Participants were drawn from the larger longitudinal Hormones and Attachment across
the Transition to Childrearing (HATCH) study. The study follows couples from mid-to-late
pregnancy across the first year postpartum. Recruitment occurred via flyers, social media
advertising, and word of mouth. The current study uses data from a prenatal laboratory visit,
conducted in mid-to-late pregnancy, and a separate MRI visit that occurred within two weeks of
the in-lab visit. Inclusion criteria included that couples were cohabiting, pregnant for the first-
time with a singleton fetus, and free of use of psychotropic medication. Exclusion criteria
included any contraindications for MR scanning, use of psychotropic medication, and left-
handedness.
Data for the current study was available for 41 expectant fathers who provided handgrip
and self-report data. Of these, 34 fathers also provided neuroimaging data and 32 of these fathers
provided testosterone data. Mean age was 31.7 years old (SD = 4.25 years). The sample was
highly educated with 80% of participants achieving a college degree or higher, and the
population was ethnically diverse (36% white, 7% black, 26% Hispanic or Latin, 24% Asian or
Pacific Islander, and 5% other).
Procedure
Expectant fathers participated in one in-lab visit scheduled mid-to-late pregnancy
(average weeks pregnant = 29 weeks, SD = 4.7 weeks, range = 18 to 38 weeks) and an MRI visit
an average of 1.05 (SD = 1.04, range 0-4 weeks) weeks later. The majority of fathers (31 out of
34 fathers) completed the scan visit within two weeks of the in-lab visit. During the prenatal in-

17
lab visit, each father provided three saliva samples for testosterone sampling over 90 minutes,
and completed the handgrip task after saliva collection, as described below. Additionally, after
completing the handgrip task, fathers were asked to listen to the infant cry noise and complete
the Emotional Reactions Questionnaire (ERQ) and Trait-Rating task. During the MRI visit,
fathers completed the same infant cry task as part of a larger MRI data collection protocol. All
procedures were approved by the University IRB and all participants signed informed consent
forms prior to participation.
Infant cry task. Using the same infant cry and control sounds as a previous study (Riem
et al., 2014), the cry task included six 30-second auditory clips of infant crying interspersed with
six 30-second clips of frequency-matched white noise counterbalanced across participants
leading to 12 trials. The stimuli were presented electronically using the E-Prime 3.0 software
(Psychology Software Tools, Pittsburgh, PA) in a block design. The task was six minutes long
and administered in one run.
Handgrip modulation. Established procedures for handgrip dynamometer data
collection (Bakermans-Kranenburg et al., 2012a; Crouch et al., 2008; Riem et al., 2012) were
followed. Prior to playing the infant cry task, a research assistant demonstrated correct hand
placement on the dynamometer and modeled the handgrip task (with their dominant hand).
Participants watched a line graph indicating grip-strength on the computer screen. The
participant was asked to perform a full-strength squeeze and a half-strength squeeze while
watching the line graph. The RA gave verbal feedback on each trial to demonstrate an accurate
full-strength and half-strength grip. Once the participant performed this task accurately for three
consecutive trials, the participant performed the infant cry task. During data collection,
participants were prompted to do a full-strength squeeze, followed two seconds later by a half-

18
strength squeeze one time per infant cry and white noise trial. Participants averaged 30 trials (of
one full-strength grip and one half-strength grip) during training, with a range of seven to 50
trials to master the task.
Testosterone. Saliva samples were collected in cryosafe collection tubes using passive
drool and then stored at -80c before shipment on dry ice to the Technical University of Dresden
(Kirschbaum, PI) to be assayed. Fathers were instructed not to eat, drink anything besides water,
and chew gum within an hour of before collection. Timing of collection was held constant across
participants to minimize variability, and T samples were taken during the first 90 minutes of the
prenatal laboratory visit and were not concurrent with the handgrip task described above which
occurred after all saliva samples were collected. Testosterone levels were averaged across the
three samples.
Neuroimaging protocol. Imaging was performed on a Siemens 3 Tesla MAGNETOM
Prisma scanner using a 20-channel matrix head coil. Functional images were collected using a
T2* weighted Echo Planar (EPI) sequence (32 transversal slices; TR = 2000ms; TE = 25 ms; flip
angle = 90
o
) with a voxel resolution of 3mm x 3mm x 2.5mm. Anatomical images were acquired
using a magnetization prepared rapid acquisition gradient (MPRAGE) sequence (TR = 2530 ms;
TE = 3.13ms; flip angle 10
o
; isotropic voxel resolution 1mm
3
). Task sounds were transmitted
using Siemens V14 sound headphone system.
Measures
Emotional reactions questionnaire. Following the in-lab cry task, participants
completed the Emotional Reactions Questionnaire (ERQ; Milner et al., 1995a) to indicate how
well each adjective describes their present mood (1, not at all, to 7, extremely well). The negative
emotions (bothered, irritated, annoyed and hostile) subscales were averaged to determine

19
negative emotions after listening to infant cry (Cronbach’s α = .89 in the current sample, Crouch
et al., 2008).
Trait rating task. Also following the in-lab infant cry task, each father was asked to rate
the infant on nine traits (i.e. hostile, negative, difficult, friendly, cooperative, sweet, content,
lively, attached). Following the procedures of previously conducted studies (Crouch et al., 2008;
Bakermans-Kranenburg et al., 2012a) the trait ratings were made on a 10-point scale (ranging
from 1 not at all to 10, extremely likely). Positive traits were also included to increase validity
and decrease bias toward negative traits. Trait ratings were averaged across the three negative
traits (hostile, negative, and difficult) to obtain a composite negative trait rating (Cronbach’s α =
.75 in the current sample).
Analyses
Hypothesis 1. Neural responses to infant cry were analyzed using FEAT (FMRI Expert
Analysis Tool) of FSL (FMRIB’s Software Library, www.FMRIb.ox.ac.uk/fsl; Smith et al.,
2004). First, motion correction using MCFLIRT, non-brain removal, spatial smoothing (5mm
FWHM Gaussian kernel), and registration to T1-weighted images using FSL FLIRT were done
for pre-processing. Then, functional activation was examined with general linear model analyses.
To identify regions involved in the perception of infant crying, contrasts of cry > white noise and
white noise > cry were assessed. Contrasts of parameter estimates (COPEs) for cry > white noise
and white noise > cry sound tested primary hypotheses regarding response to infant cry versus a
frequency matched white noise. First-level COPEs served as inputs to higher-level group
analyses conducted using FLAME to model random-effects components of mixed-effects
variance. Images were thresholded with clusters determined by Z > 2.3 and a cluster-corrected
significance threshold of p < .05 (Worsley et al., 2002) to identify regions that were activated

20
during cry versus white noise across the six blocks for each sound. Father’s age and weeks
pregnant were mean-centered and included as confound regressors in all models. Models were
run with and without covariates and yielded similar results. To visualize results, spherical ROI’s
(r = 5mm) centered on activation peaks were used to extract signal change for each condition.
Additionally, given our a priori hypotheses focusing on the amygdala, ROI analyses of
the bilateral amygdala were conducted. Parameter estimate values were converted to percentage
signal change values via scaling of the PE or COPE values by (100*) the peak-peak height of the
regressor (or effective regressor in the case of COPEs) and then by dividing by the mean over
time of the filtered functional data. A report was generated using Featquery with statistics
derived from each image’s values within the mask. Percent signal change was extracted from
bilateral amygdala using anatomically-defined masks created using the Harvard-Oxford
Subcortical Atlas.
Hypothesis 2. Consistent with previous studies (Bakermans-Kranenburg et al., 2012a;
Crouch et al., 2008; Riem et al., 2012), handgrip modulation was calculated by dividing the half-
squeeze intensity by the maximum squeeze intensity per block, and an average ratio of half
strength/full strength squeezes was calculated for infant cry and white noise per person.
Hypothesis 3. Demeaned self-report ratings of infant cry and negative emotions were
added (separately) as regressors into the general linear model described above. The first-level
contrast images of cry > white noise and white noise > cry were submitted to second-level whole
brain analysis to determine differences in activation depending on interpretations of the infant as
more negative, and self-reported negative emotions after infant cry. Both positive and negative
contrast weights were tested for each continuous predictor to determine whether it related to
increased or decreased neural response. Lastly, multivariate regression analyses were used to test

21
the relationship between interpretations of the infant as more negative, and negative emotions
during infant cry and signal change in the amygdala ROI.
Hypothesis 4. First, T was added as a regressor to the FSL models testing the
relationship between prenatal T and neural activation to infant cry as described above. Next,
multivariate regressions were run to determine the relationship between (a) testosterone and
negative infant interpretations, and (b) testosterone and self-reported negative emotions while
listening to infant cry. All analyses included gestational age of the infant and father’s age as
covariates.
We adjusted for multiple comparisons using the Holm-Bonferonni method (Holm, 1979),
in which the alpha value is adjusted such that the lowest p-value (I = 1) is expected to fall below
a/k, (where k is the number of analyses), and the higher values to progressively less restrictive
thresholds (a/(k-I + 1). Therefore, for six planned analyses, we would require at least one model
be significant at p = 0.008 (0.05/)6; one model significant at 0.01 (0.05/5); one at 0.013 (0.05/4);
one at 0.017 (0.05/3); and one at 0.025 (0.05/2).
Results
First, all data were explored for outliers and normality of distribution. All variables met
the criteria for normality. One subject was dropped due to neural activation in the contrast of
interest (cry > white noise) being three standard deviations above the mean. Means and standard
deviations of all study variables are presented in Table 1, and bivariate correlations of main study
variables are shown in Table 2.
Hypothesis 1. As depicted in Table 3 and Figure 1, greater activation to infant cry than
frequency-matched white noise emerged primarily in areas of the bilateral temporal lobes
consisting of the auditory cortices. These areas included bilateral planum temporale, left insular

22
cortex, R Heschl’s gyrus, R STG (posterior and anterior), R planum polare, and left
supramarginal gyrus, as well as the right IFG. No differences were found in amygdala activation
in response to infant cry versus white noise. No activation differences were found in response to
white noise > infant cry.
Hypothesis 2. A paired samples t-test was used to test for differences in handgrip
modulation during infant cry and white noise across fathers. This hypothesis was not supported.
No significant differences were found in the ratio of half-strength/full-strength during infant cry
versus white noise across fathers (t (40) = -1.21, p = 2.32). Given the lack of a main effect
finding of handgrip modulation during infant cry compared to white noise, no further analyses
were done testing the relationship between handgrip strength during infant cry and T or
psychological and neural responses to infant cry.
Hypothesis 3. This hypothesis was partially supported. Neural activation in whole-brain
analyses was not associated with fathers’ self-reported negative emotions or their negative trait
ratings of infant cry. Negative interpretations of infant cry (trait rating task) predicted right
amygdala percent signal change to infant cry (B = .36, p = .04; Table 4), but this effect was not
significant following correction for multiple comparisons. Handgrip force was not tested given
the null findings in the second hypothesis.
Hypothesis 4. As hypothesized, expectant fathers with higher prenatal T showed greater
activation to infant cry sounds relative to white noise sounds in the right supramarginal gyrus,
the left occipital cortex, and the precuneus cortex (Table 5, Figure 2). No activation was found to
be negatively associated with prenatal T level. Prenatal T level did not predict fathers’ negative
emotion ratings in response to infant cry (B = .12, p = .52), nor negative ratings of the infant
during infant cry (B = .22, p = .16).

23
Discussion
This study sought to characterize neural, behavioral, and psychological responses to
infant cry sounds among expectant fathers and to examine whether their prenatal T levels were
associated with these responses to cry sounds. Our hypotheses were partially supported. As
expected, infant cry (vs. white noise sounds) elicited activation in regions including the superior
temporal gyrus (STG), inferior frontal gyrus (IFG) and insula, although we did not find expected
differences in amygdala activation. Additionally, parts of the auditory cortices such as the
planum temporale, Heschl’s gyrus and supramarginal gyrus were additionally found to be more
active during infant cry than a frequency matched white noise, similarly to previous studies of
first-time fathers and expectant fathers (Li et al., 2018; Thijssen et al., 2018). The planum
temporale is situated posterior to the primary auditory cortex and has been found to be active
during the processing of speech related cues (Baars & Gage, 2010), locating sound in space
(Hickok, 2009), and stimulus selection and auditory attention (Hirnstein et al., 2013). The
supramarginal gyrus has been implicated in phonological processing (e.g. Church et al., 2011;
Saur et al., 2008) and in social cognition (Silani et al., 2013; Singer & Klimecki, 2014). Our
sample of expectant fathers appeared to recognize infant cry sounds as a meaningful speech
signal over and above a frequency-matched white noise.
Additionally, areas of the brain associated with social cognition (i.e. insula,
supramarginal gyrus) appear to be more active during cry than white noise. Previous research has
suggested that, in contrast to biological mothers, who form a physical bond with the infant prior
to birth, the paternal brain primarily comes “online” after the birth of a child and in interacting
with their newborn (Abraham et al., 2014). However, the current research suggests that infant
cry is nevertheless a salient stimulus for expectant fathers, even though they have not yet

24
participated in caregiving for their own infants. Fine-tuning of these processes may occur with
socialization with the infant as has been previously hypothesized (Abraham et al., 2014; Atzil et
al., 2012, Feldman, 2015). However, the current study suggests that expectant fathers show
processing of infant cry that is similar to mothers and to fathers whose child is already born.
Future studies would benefit from including expectant fathers at different stages of pregnancy, or
age-matched non-fathers to understand whether similar brain responses emerge in non-parents
and if not, when and how these brain responses come “online” during pregnancy.
Although we did not find a main effect for amygdala activation in response to infant cry
vs. white noise, we did find that fathers’ negative views of infant cry predicted greater right
amygdala activation during infant cry compared to white noise. This finding contradicts a
previous study that found no relationship between amygdala activation and mothers’ reported
irritation during infant cry (Riem et al., 2012b), suggesting that this finding may be specific to
expectant fathers. However, this result should be interpreted with caution given that it did not
reach the level of significance after controlling for multiple comparisons. Amygdala activation
has been found to be greater during infant cry compared to other forms of infant stimuli (such as
laughter), and activation in these areas has been purported to underlie parental vigilance to infant
distress cues (Abraham et al., 2014). Amygdala activation may also represent the hub of the
emotion-processing sub-network of the parenting brain (Abraham & Feldman, 2018; Feldman,
2015).
Counter to hypotheses, no difference in handgrip modulation was found during infant cry
compared to frequency-matched white noise. Several reasons may explain these null results.
Firstly, although handgrip has been used to reflect potential aggression in response to infant cry
in previous studies (Crouch et al., 2008), these studies did not include a control sound

25
comparison. Similar null results were found in another population of expectant fathers
(Alyousefi-Van Dijk et al., 2019) when comparing handgrip during infant cry and a frequency-
matched white noise. These results together suggest that handgrip modulation may not be a
specific response to infant cry per se but may reflect hyperarousal to aversive stimuli in general.
Our null results may also be due to limitations of the handgrip task itself, or an underpowered
sample size. It was challenging for participants to master the half-strength grip procedure and
many subjects became frustrated during the training. It may be that this task is not reliable
enough across participants to indicate a sensitive response to infant cry.
The relationship between psychological and neural response to infant cry was
additionally only partially supported in the current investigation, unlike previous studies which
found a relationship between neural activation in auditory cortices and greater irritation to infant
cry in new fathers (Li et al., 2018). However, these results are in line with a previous
investigation of mothers which failed to find a relationship between reported aversiveness of
infant cry and neural activation to the same sound (Riem et al., 2012b). These null results may be
due to a small sample size and an underpowered whole-brain analysis. However, the current
sample size is in line with previous investigations (Li et al., 2018; Riem et al., 2012b).
Additionally, this relationship may be stronger in fathers who could be at risk for aggressive
parenting or abuse. The current investigation sought to characterize the neural and psychological
response to infant cry in a community sample of expectant fathers and thus may not capture the
responses indicative of abusive parenting risk.
Higher prenatal T predicted greater activation in right supramarginal gyrus, left occipital
cortex and the precuneus cortex. As stated above, previous studies have found the supramarginal
gyrus to be implicated in social cognition (Silani et al., 2013; Singer & Klimecki, 2014), and the

26
precuneus cortex to be associated with arousal and reward learning (Swain, 2011). Previous
studies of first-time fathers and baseline T found no neural activation differences in infant cry vs.
white noise in conjunction with T level (Li et al., 2018). However, T reactivity to infant stimuli
has been found to be associated with greater neural activation in the left caudate to infant cry
while listening to infant cry and watching infant video stimuli (Kuo et al., 2012). Given that this
is the first investigation to look at neural activation to infant cry in expectant fathers and baseline
prenatal T levels this finding may be specific to the prenatal period. Additionally, greater
activation to infant cry in association with higher T level may indicate a hyperreactivity to infant
cry in line with previous investigations that have found a positive relationship between neural
activation and greater reports of irritation to infant cry in new fathers (Li et al., 2008). However,
the current investigation did not measure parenting behaviors postpartum in these fathers with
higher T. Future studies can investigate whether fathers’ prenatal responses to cry sounds predict
their actual parenting behaviors following the birth of their child, and how this relates to T level
across pregnancy.
The current study had a number of limitations. Our sample size was small (34 fathers),
albeit larger than the samples used in most published studies of the parenting brain. Moreover,
we used a multi-modal approach, incorporating behavioral (handgrip), hormonal (testosterone),
and self-report data as well as MRI data. Another limitation is that because our sample consisted
of expectant fathers, we were not able to present own-infant sounds, and instead played
unfamiliar cry sounds. However, this limitation is balanced by the advantage that the stimuli was
standardized across participants and has been used in previous studies even with participants
who were already parents. Additionally, T levels and neuroimaging responses were collected on
different days, and therefore may not fully map onto each other. Finally, these data are cross-

27
sectional, focusing only on expectant fathers. Future studies can extend this work by focusing on
prenatal responses to cry as a predictor of later parenting behavior. Given the exploratory nature
of this study and the novel population included, future research should endeavor for an increased
sample size to replicate and clarify these results.
Despite its important limitations, this study is one of the first to investigate responses to
infant cry in expectant fathers across multiple modalities. Our sample was ethnically diverse and
contributed hormonal, behavioral, neural, and self-report data. Our findings suggest that
expectant fathers process infant cry as a salient and meaningful speech sound that may require
empathic responding, even before the child is born. Additionally, it appears that T may moderate
this effect, with expectant fathers who were higher in T also showing stronger neural responses
to infant cry. Given the importance of fathers to healthy child development, this work contributes
to our understanding of the fathering brain and can ultimately improve the detection of fathers at
risk and inform the development of interventions that target expectant fathers.
References
Abraham, E., & Feldman, R. (2018). The neurobiology of human allomaternal care; implications
for fathering, coparenting, and children’s social development. Physiology and Behavior,
193(May), 25–34. https://doi.org/10.1016/j.physbeh.2017.12.034
Abraham, E., Hendler, T., Shapira-Lichter, I., Kanat-Maymon, Y., Zagoory-Sharon, O., &
Feldman, R. (2014). Father’s brain is sensitive to childcare experiences. Proceedings of the
National Academy of Sciences. https://doi.org/10.1073/pnas.1402569111
Ahlborg, T., Misvaer, N., & Möller, A. (2009). Perception of marital quality by parents with
small children: A follow-up study when the firstborn is 4 years old. Journal of Family
Nursing, 15(2), 237–263. https://doi.org/10.1177/1074840709334925

28
Ainsworth, M. S. (1979). Infant-mother attachment. American Psychologist.
https://doi.org/10.1037/0003-066X.34.10.932
Alyousefi-Van Dijk, K., van ‘t Veer, A. E., Meijer, W. M., Lotz, A. M., Rijlaarsdam, J.,
Witteman, J., & Bakermans-Kranenburg, M. J. (2019). Vasopressin differentially affects
handgrip force of expectant fathers in reaction to own and unknown infant faces. Frontiers
in Behavioral Neuroscience, 13. https://doi.org/10.3389/fnbeh.2019.00105
Atzil, S., Hendler, T., Zagoory-Sharon, O., Winetraub, Y., & Feldman, R. (2012). Synchrony and
specificity in the maternal and the paternal brain: Relations to oxytocin and vasopressin.
Journal of the American Academy of Child and Adolescent Psychiatry.
https://doi.org/10.1016/j.jaac.2012.06.008
Baars, B. J., & Gage, N. M. (2010). Cognition, Brain and Consciousness. In Cognition, Brain
and Consciousness. https://doi.org/10.1016/C2009-0-01556-6
Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Riem, M. M. E., Tops, M., & Alink, L. R.
A. (2012a). Oxytocin decreases handgrip force in reaction to infant crying in females
without harsh parenting experiences. Social Cognitive and Affective Neuroscience, 7(8),
951–957. https://doi.org/10.1093/scan/nsr067
Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Riem, M. M. E., Tops, M., & Alink, L. R.
A. (2012b). Oxytocin decreases handgrip force in reaction to infant crying in females
without harsh parenting experiences. Social Cognitive and Affective Neuroscience, 7(8),
951–957. https://doi.org/10.1093/scan/nsr067
Barr, R. G., Rivara, F. P., Barr, M., Cummings, P., Taylor, J., Lengua, L. J., & Meredith-Benitz,
E. (2009). Effectiveness of Educational Materials Designed to Change Knowledge and
Behaviors Regarding Crying and Shaken-Baby Syndrome in Mothers of Newborns: A

29
Randomized, Controlled Trial. Pediatrics, 123(3), 972–980.
https://doi.org/10.1542/peds.2008-0908
Barr, Ronald G., Fairbrother, N., Pauwels, J., Green, J., Chen, M., & Brant, R. (2014). Maternal
frustration, emotional and behavioural responses to prolonged infant crying. Infant Behavior
and Development, 37(4), 652–664. https://doi.org/10.1016/j.infbeh.2014.08.012
Barr, Ronald G., Trent, R. B., & Cross, J. (2006). Age-related incidence curve of hospitalized
Shaken Baby Syndrome cases: Convergent evidence for crying as a trigger to shaking.
Child Abuse and Neglect, 30(1), 7–16. https://doi.org/10.1016/j.chiabu.2005.06.009
Beebe, S. A., Casey, R., & Pinto-Martin, J. (1993). Association of Reported Infant Crying and
Maternal Parenting Stress. Clinical Pediatrics, 32(1), 15–19.
https://doi.org/10.1177/000992289303200103
Berry, J. O., & Jones, W. H. (1995). The Parental Stress Scale: Initial Psychometric Evidence.
Journal of Social and Personal Relationships, 12(3), 463–472.
https://doi.org/10.1177/0265407595123009
Blunt Bugental, D., Ellerson, P. C., Lin, E. K., Rainey, B., Kokotovic, A., & O’hara, N. (2002).
A Cognitive Approach to Child Abuse Prevention. https://doi.org/10.1037/0893-
3200.16.3.243
BORNSTEIN, M. (2008). Handbook of Parenting. Pratical Issues in Parenting.
Bos, P. A. (2017). The endocrinology of human caregiving and its intergenerational
transmission. Development and Psychopathology, 29(3), 971–999.
https://doi.org/10.1017/S0954579416000973
Bos, P. A., Hechler, C., Beijers, R., Shinohara, K., Esposito, G., & de Weerth, C. (2018).
Prenatal and postnatal cortisol and testosterone are related to parental caregiving quality in

30
fathers, but not in mothers. Psychoneuroendocrinology, 97, 94–103.
https://doi.org/10.1016/j.psyneuen.2018.07.013
Bos, P. A., Hermans, E. J., Montoya, E. R., Ramsey, N. F., & van Honk, J. (2010). Testosterone
administration modulates neural responses to crying infants in young females.
Psychoneuroendocrinology, 35(1), 114–121.
https://doi.org/10.1016/j.psyneuen.2009.09.013
Brockington, I. F., Fraser, C., & Wilson, D. (2006). The Postpartum Bonding Questionnaire: A
validation. Archives of Women’s Mental Health, 9(5), 233–242.
https://doi.org/10.1007/s00737-006-0132-1
Brosch, T., Sander, D., & Scherer, K. R. (2007). That baby caught my eye... attention capture by
infant faces. Emotion (Washington, D.C.), 7(3), 685–689. https://doi.org/10.1037/1528-
3542.7.3.685
Brown, W. A., Monti, P. M., & Corriveau, D. P. (1978). Serum testosterone and sexual activity
and interest in men. Archives of Sexual Behavior, 7(2), 97–103.
Caselles, C. E., & Milner, J. S. (2000). Evaluations of child transgressions, disciplinary choices,
and expected child compliance in a no-cry and a crying infant condition in physically
abusive and comparison mothers. Child Abuse & Neglect, 24(4), 477–491.
http://www.ncbi.nlm.nih.gov/pubmed/10798838
Church, J. A., Balota, D. A., Petersen, S. E., & Schlaggar, B. L. (2011). Manipulation of length
and lexicality localizes the functional neuroanatomy of phonological processing in adult
readers. Journal of Cognitive Neuroscience. https://doi.org/10.1162/jocn.2010.21515
Compier-de Block, L. H. C. G., Alink, L. R. A., Reijman, S., Werner, C. D., Maras, A., Rijnberk,
C., van IJzendoorn, M. H., & Bakermans-Kranenburg, M. J. (2015). Handgrip force of

31
maltreating mothers in reaction to infant signals. Child Abuse and Neglect, 40, 124–131.
https://doi.org/10.1016/j.chiabu.2014.03.006
Condon, J. T., Boyce, P., & Corkindale, C. J. (2004). The first-time fathers study: A prospective
study of the mental health and wellbeing of men during the transition to parenthood.
Australian and New Zealand Journal of Psychiatry, 38(1–2), 56–64.
https://doi.org/10.1111/j.1440-1614.2004.01298.x
Coster, D. (2017). Changing Parents’ Behaviour Using a Psycho-Educational Film as a
Preventative Measure to Reduce the Risk of Non-accidental Head Injury. Child Abuse
Review, 26(6), 465–476. https://doi.org/10.1002/car.2469
Crouch, J. L., & Behl, L. E. (2001). Relationships among parental beliefs in corporal
punishment, reported stress, and physical child abuse potential. Child Abuse and Neglect,
25(3), 413–419. https://doi.org/10.1016/S0145-2134(00)00256-8
Crouch, J. L., Skowronski, J. J., Milner, J. S., & Harris, B. (2008). Parental responses to infant
crying: The influence of child physical abuse risk and hostile priming. Child Abuse and
Neglect, 32(7), 702–710. https://doi.org/10.1016/j.chiabu.2007.11.002
Del Vecchio, T., Walter, A., & O’Leary, S. G. (2009). Affective and physiological factors
predicting maternal response to infant crying. Infant Behavior and Development, 32(1),
117–122. https://doi.org/10.1016/j.infbeh.2008.10.005
Ellett, M. L. C., & Swenson, M. (2005). Living With a Colicky Infant. Gastroenterology
Nursing, 28(1), 19–25. https://doi.org/10.1097/00001610-200501000-00005
Fairbrother, N., Barr, R. G., Pauwels, J., Brant, R., & Green, J. (2015). Maternal thoughts of
harm in response to infant crying: an experimental analysis. Archives of Women’s Mental
Health, 18(3), 447–455. https://doi.org/10.1007/s00737-014-0471-2

32
Feldman, R. (2015). Sensitive periods in human social development: New insights from research
on oxytocin, synchrony, and high-risk parenting. Development and Psychopathology, 27(2),
369–395. https://doi.org/10.1017/s0954579415000048
Fleming, A. S., Corter, C., Stallings, J., & Steiner, M. (2002). Testosterone and prolactin are
associated with emotional responses to infant cries in new fathers. Hormones and Behavior,
42(4), 399–413. http://www.ncbi.nlm.nih.gov/pubmed/12488107
Frodi, A. (1985). When Empathy Fails. In Infant Crying (pp. 263–277). Springer US.
https://doi.org/10.1007/978-1-4613-2381-5_12
Frodi, A. M., & Lamb, M. E. (1980). Child abusers’ responses to infant smiles and cries. Child
Development, 51(1), 238–241. http://www.ncbi.nlm.nih.gov/pubmed/7363736
Gallup, A. C., O’Brien, D. T., White, D. D., & Wilson, D. S. (2010). Handgrip strength and
socially dominant behavior in male adolescents. Evolutionary Psychology, 8(2), 229–243.
Garcia, R. L., Kenny, D. A., & Ledermann, T. (2015). Moderation in the actor-partner
interdependence model. Personal Relationships, 22(1), 8–29.
https://doi.org/10.1111/pere.12060
Gettler, L. T., McDade, T. W., Feranil, A. B., & Kuzawa, C. W. (2011). Longitudinal evidence
that fatherhood decreases testosterone in human males. Proceedings of the National
Academy of Sciences, 108(39), 16194–16199. https://doi.org/10.1073/pnas.1105403108
Gettler, Lee T., McDade, T. W., Agustin, S. S., Feranil, A. B., & Kuzawa, C. W. (2013). Do
testosterone declines during the transition to marriage and fatherhood relate to men’s sexual
behavior? Evidence from the Philippines. Hormones and Behavior, 64(5), 755–763.
https://doi.org/10.1016/j.yhbeh.2013.08.019
Gray, P. B., Parkin, J. C., & Samms-Vaughan, M. E. (2007). Hormonal correlates of human

33
paternal interactions: a hospital-based investigation in urban Jamaica. Hormones and
Behavior, 52(4), 499–507. https://doi.org/10.1016/j.yhbeh.2007.07.005
Hickok, G. (2009). The functional neuroanatomy of language. Physics of Life Reviews, 6(3),
121–143. https://doi.org/10.1016/j.plrev.2009.06.001
Hillson, J., Review, N. K.-C. P., & 1994, undefined. (n.d.). A stress and coping model of child
maltreatment. Elsevier. Retrieved March 5, 2020, from
https://www.sciencedirect.com/science/article/pii/0272735894900256
Hirnstein, M., Westerhausen, R., Korsnes, M. S., & Hugdahl, K. (n.d.). Sex differences in
language asymmetry are age-dependent and small: a large-scale, consonant-vowel dichotic
listening study with behavioral and fMRI data. Cortex; a Journal Devoted to the Study of
the Nervous System and Behavior, 49(7), 1910–1921.
https://doi.org/10.1016/j.cortex.2012.08.002
Holm, S. (1979). Board of the Foundation of the Scandinavian Journal of Statistics A Simple
Sequentially Rejective Multiple Test Procedure. In Source: Scandinavian Journal of
Statistics (Vol. 6, Issue 2).
Joosen, K. J., Mesman, J., Bakermans-Kranenburg, M. J., Pieper, S., Zeskind, P. S., & Van
Ijzendoorn, M. H. (2013). Physiological Reactivity to Infant Crying and Observed Maternal
Sensitivity. Infancy, 18(3), 414–431. https://doi.org/10.1111/j.1532-7078.2012.00122.x
Joosen, K. J., Mesman, J., Bakermans-Kranenburg, M. J., & van IJzendoorn, M. H. (2013).
Maternal Overreactive Sympathetic Nervous System Responses to Repeated Infant Crying
Predicts Risk for Impulsive Harsh Discipline of Infants. Child Maltreatment, 18(4), 252–
263. https://doi.org/10.1177/1077559513494762
Kenny, D. A. (1996). Models of non-independence in dyadic research. Journal of Social and

34
Personal Relationships, 13(2), 279–294. https://doi.org/10.1177/0265407596132007
Kenny, D. A., & Judd, C. M. (1986). Consequences of Violating the Independence Assumption
in Analysis of Variance. Psychological Bulletin, 99(3), 422–431.
https://doi.org/10.1037/0033-2909.99.3.422
Kim, P., Capistrano, C., & Congleton, C. (2016). Socioeconomic disadvantages and neural
sensitivity to infant cry: Role of maternal distress. Social Cognitive and Affective
Neuroscience, 11(10), 1597–1607. https://doi.org/10.1093/scan/nsw063
Kim, P., Leckman, J. F., Mayes, L. C., Newman, M. A., Feldman, R., & Swain, J. E. (2010).
Perceived quality of maternal care in childhood and structure and function of mothers’
brain. Developmental Science, 13(4), 662–673. https://doi.org/10.1111/j.1467-
7687.2009.00923.x
Knutson, J. F. (1978). Child abuse as an area of aggression research. Journal of Pediatric
Psychology, 3(1), 20–27. https://doi.org/10.1093/jpepsy/3.1.20
Kuo, P. X., Carp, J., Light, K. C., & Grewen, K. M. (2012). Neural responses to infants linked
with behavioral interactions and testosterone in fathers. Biological Psychology, 91(2), 302–
306. https://doi.org/10.1016/j.biopsycho.2012.08.002
Kurth, E., Kennedy, H. P., Spichiger, E., Hösli, I., & Zemp Stutz, E. (2011). Crying babies, tired
mothers: What do we know? A systematic review. Midwifery, 27(2), 187–194.
https://doi.org/10.1016/j.midw.2009.05.012
Ladd, A. (2010). Infant cries as predictors of child abuse potential: Associations of working
models of attachment, cognitive appraisals and emotional reactions.
http://search.proquest.com/openview/41ebd00d886190638de53e05e95a06cc/1?pq-
origsite=gscholar&cbl=18750&diss=y

35
Li, T., Horta, M., Mascaro, J. S., Bijanki, K., Arnal, L. H., Adams, M., Barr, R. G., & Rilling, J.
K. (2018). Explaining individual variation in paternal brain responses to infant cries.
Physiology and Behavior, 193(December 2017), 43–54.
https://doi.org/10.1016/j.physbeh.2017.12.033
Lorberbaum, J. P., Newman, J. D., Dubno, J. R., Horwitz, A. R., Nahas, Z., Teneback, C.,
Johnson, M. R., Lydiard, R. B., Ballenger, J. C., & George, M. S. (1998). Feasibility Of
Using fMRI To Study Mothers Responding To Infant Cries. NeuroImage, 7(4), S907.
https://doi.org/10.1016/S1053-8119(18)31740-3
Lynch, M. (2003). Consequences of Children ’ s Exposure to Community Violence. 6(4).
Magill-Evans, J., & Harrison, M. J. (2001). Parent-child interactions, parenting stress, and
developmental outcomes at 4 years. Children’s Health Care, 30(2), 135–150.
https://doi.org/10.1207/S15326888CHC3002_4
Malmberg, L.-E., Lewis, S., West, A., Murray, E., Sylva, K., & Stein, A. (2016). The influence
of mothers’ and fathers’ sensitivity in the first year of life on children’s cognitive outcomes
at 18 and 36 months. Child: Care, Health and Development, 42(1), 1–7.
https://doi.org/10.1111/cch.12294
Megel, M. E., Wilson, M. E., Bravo, K., McMahon, N., & Towne, A. (2011). Baby Lost and
Found: Mothers’ Experiences of Infants Who Cry Persistently. Journal of Pediatric Health
Care, 25(3), 144–152. https://doi.org/10.1016/j.pedhc.2009.10.005
Messina, I., Cattaneo, L., Venuti, P., de Pisapia, N., Serra, M., Esposito, G., Rigo, P., Farneti, A.,
& Bornstein, M. H. (2016). Sex-specific automatic responses to infant cries: TMS reveals
greater excitability in females than males in motor evoked potentials. Frontiers in
Psychology, 6(JAN). https://doi.org/10.3389/fpsyg.2015.01909

36
Milner, J. S., Halsey, L. B., & Fultz, J. (1995a). Empathic responsiveness and affective reactivity
to infant stimuli in high- and low-risk for physical child abuse mothers. Child Abuse and
Neglect, 19(6), 767–780. https://doi.org/10.1016/0145-2134(95)00035-7
Milner, J. S., Halsey, L. B., & Fultz, J. (1995b). Empathic responsiveness and affective reactivity
to infant stimuli in high- and low-risk for physical child abuse mothers. Child Abuse and
Neglect. https://doi.org/10.1016/0145-2134(95)00035-7
Molfese, V. J., Rudasill, K. M., Beswick, J. L., Jacobi-Vessels, J. L., Ferguson, M. C., & White,
J. M. (2010). Infant Temperament, Maternal Personality, and Parenting Stress as
Contributors to Infant. In Source: Merrill-Palmer Quarterly (Vol. 56, Issue 1).
Murray, A. D. (1985). Aversiveness is in the Mind of the Beholder. In Infant Crying (pp. 217–
239). Springer US. https://doi.org/10.1007/978-1-4613-2381-5_10
Musser, E. D., Kaiser-Laurent, H., & Ablow, J. C. (2012). The neural correlates of maternal
sensitivity: An fMRI study. Developmental Cognitive Neuroscience, 2(4), 428–436.
https://doi.org/10.1016/j.dcn.2012.04.003
Neece, C. L., Green, S. A., & Baker, B. L. (2012). Parenting stress and child behavior problems:
A transactional relationship across time. American Journal on Intellectual and
Developmental Disabilities, 117(1), 48–66. https://doi.org/10.1352/1944-7558-117.1.48
Oldbury, S., & Adams, K. (2015). The impact of infant crying on the parent-infant relationship.
Community Practitioner, 88(3), 29–34.
Out, D., Bakermans-Kranenburg, M. J., van Pelt, J., & van IJzendoorn, M. H. (2012). Salivary α-
Amylase and Intended Harsh Caregiving in Response to Infant Crying: Evidence for
Physiological Hyperreactivity. Child Maltreatment, 17(4), 295–305.
https://doi.org/10.1177/1077559512464427

37
Parsons, C. E., Stark, E. A., Young, K. S., Stein, A., & Kringelbach, M. L. (2013).
Understanding the human parental brain: A critical role of the orbitofrontal cortex. Social
Neuroscience, 8(6), 525–543. https://doi.org/10.1080/17470919.2013.842610
Pastore, L., Owens, A., & Raymond, C. (2007). Postpartum sexuality concerns among first-time
parents from one U.S. Academic Hospital. Journal of Sexual Medicine, 4(1), 115–123.
https://doi.org/10.1111/j.1743-6109.2006.00379.x
Persky, H., Lief, H. I., Strauss, D., Miller, W. R., & O’Brien, C. P. (1978). Plasma testosterone
level and sexual behavior of couples. Archives of Sexual Behavior, 7(3), 157–173.
https://doi.org/10.1007/BF01542376
Petts, R. J., Shafer, K. M., & Essig, L. (2018). Does Adherence to Masculine Norms Shape
Fathering Behavior? Journal of Marriage and Family, 80(3), 704–720.
https://doi.org/10.1111/jomf.12476
Raboch, J., & Stárka, L. (1973). Reported coital activity of men and levels of plasma
testosterone. Archives of Sexual Behavior, 2(4), 309–315.
https://doi.org/10.1007/BF01541004
Reijman, S., Alink, L. R. A., Block, L. H. C. G. C. de, Werner, C. D., Maras, A., Rijnberk, C.,
IJzendoorn, M. H. va., & Bakermans-Kranenburg, M. J. (2014). Autonomic Reactivity to
Infant Crying in Maltreating Mothers. Child Maltreatment, 19(2), 101–112.
https://doi.org/10.1177/1077559514538115
Riem, M. M. E., Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Out, D., & Rombouts, S.
A. R. B. (2012). Attachment in the brain: Adult attachment representations predict
amygdala and behavioral responses to infant crying. Attachment and Human Development,
14(6), 533–551. https://doi.org/10.1080/14616734.2012.727252

38
Riem, M. M. E., Voorthuis, A., Bakermans-Kranenburg, M. J., & van Ijzendoorn, M. H. (2014).
Pity or peanuts? Oxytocin induces different neural responses to the same infant crying
labeled as sick or bored. Developmental Science, 17(2), 248–256.
https://doi.org/10.1111/desc.12103
Rilling, J., K., & Young, L., J. (2014). The biology of mammalian parenting and its effect on
offspring social development. Science, 345(6198), 771–776.
https://doi.org/10.1126/science.1252723
Rodriguez, C. M., & Green, A. J. (1997). Parenting stress and anger expression as predictors of
child abuse potential. Child Abuse and Neglect, 21(4), 367–377.
https://doi.org/10.1016/S0145-2134(96)00177-9
Rodriguez, C. M., Russa, M. B., & Kircher, J. C. (2015). Analog assessment of frustration
tolerance: Association with self-reported child abuse risk and physiological reactivity. Child
Abuse and Neglect, 46, 121–131. https://doi.org/10.1016/j.chiabu.2015.02.017
Saur, D., Kreher, B. W., Schnell, S., Kummerer, D., Kellmeyer, P., Vry, M.-S., Umarova, R.,
Musso, M., Glauche, V., Abel, S., Huber, W., Rijntjes, M., Hennig, J., & Weiller, C. (2008).
Ventral and dorsal pathways for language. Proceedings of the National Academy of
Sciences. https://doi.org/10.1073/pnas.0805234105
Saxbe, D.E., Edelstein, R. S., Lyden, H. M., Wardecker, B. M., Chopik, W. J., & Moors, A. C.
(2017). Fathers’ decline in testosterone and synchrony with partner testosterone during
pregnancy predicts greater postpartum relationship investment. Hormones and Behavior,
90. https://doi.org/10.1016/j.yhbeh.2016.07.005
Saxbe, Darby E., Schetter, C. D., Simon, C. D., Adam, E. K., & Shalowitz, M. U. (2017). High
paternal testosterone may protect against postpartum depressive symptoms in fathers, but

39
confer risk to mothers and children. Hormones and Behavior, 95, 103–112.
https://doi.org/10.1016/j.yhbeh.2017.07.014
Silani, G., Lamm, C., Ruff, C. C., & Singer, T. (2013). Right supramarginal gyrus is crucial to
overcome emotional egocentricity bias in social judgments. The Journal of Neuroscience :
The Official Journal of the Society for Neuroscience, 33(39), 15466–15476.
https://doi.org/10.1523/JNEUROSCI.1488-13.2013
Singer, T., & Klimecki, O. M. (2014). Empathy and compassion. In Current Biology.
https://doi.org/10.1016/j.cub.2014.06.054
Storey, A. E., & Ziegler, T. E. (2016). Primate paternal care: Interactions between biology and
social experience. Hormones and Behavior, 77, 260–271.
https://doi.org/10.1016/j.yhbeh.2015.07.024
Swain, J. E. (2011). The human parental brain: In vivo neuroimaging. Progress in Neuro-
Psychopharmacology and Biological Psychiatry, 35(5), 1242–1254.
https://doi.org/10.1016/j.pnpbp.2010.10.017
Tabuchi, N., & Shimada, K. (2008). Maternal characteristics and feelings of distress resulting
from the crying of high-risk infants. Japan Journal of Nursing Science, 5(2), 99–108.
https://doi.org/10.1111/j.1742-7924.2008.00108.x
Thijssen, S., Van ’t Veer, A. E., Witteman, J., Meijer, W. M., Van Ijzendoorn, M. H.,
Bakermans-Kranenburg, M. J., & Nl, B. L. (n.d.). Effects of Vasopressin on Neural
Processing of Infant Crying in Expectant Fathers.
https://doi.org/10.1016/j.yhbeh.2018.05.014
van Anders, S. M. (2013). Beyond masculinity: Testosterone, gender/sex, and human social
behavior in a comparative context. In Frontiers in Neuroendocrinology.

40
https://doi.org/10.1016/j.yfrne.2013.07.001
Van Anders, S. M. (2012). Testosterone and sexual desire in healthy women and men. Archives
of Sexual Behavior, 41(6), 1471–1484. https://doi.org/10.1007/s10508-012-9946-2
Van Anders, S. M., Tolman, R. M., & Volling, B. L. (2012). Baby cries and nurturance affect
testosterone in men. Hormones and Behavior, 61(1), 31–36.
https://doi.org/10.1016/j.yhbeh.2011.09.012
Wilson, Katherine R., & Prior, M. R. (2011). Father involvement and child well-being. Journal
of Paediatrics and Child Health, 47(7), 405–407. https://doi.org/10.1111/j.1440-
1754.2010.01770.x
Wilson, Kathryn R., Hansen, D. J., & Li, M. (2011). The traumatic stress response in child
maltreatment and resultant neuropsychological effects. Aggression and Violent Behavior,
16(2), 87–97. https://doi.org/10.1016/j.avb.2010.12.007
Wingfield, J. C., Hegner, R. E., Dufty, A. M., & Ball, G. F. (1990). The “challenge hypothesis’’:
theoretical implications for patterns of testosterone secretion, mating systems, and breeding
strategies.” American Naturalist, 136(6), 829–846. https://doi.org/10.1086/285134
Wingfield, John C. (2017). The challenge hypothesis: Where it began and relevance to humans.
In Hormones and Behavior (Vol. 92, pp. 9–12). Academic Press Inc.
https://doi.org/10.1016/j.yhbeh.2016.11.008
Witteman, J., Van IJzendoorn, M. H., Rilling, J. K., Bos, P. A., Schiller, N. O., & Bakermans-
Kranenburg, M. J. (2019). Towards a neural model of infant cry perception. In
Neuroscience and Biobehavioral Reviews (Vol. 99, pp. 23–32). Elsevier Ltd.
https://doi.org/10.1016/j.neubiorev.2019.01.026
Worsley, K. J., Liao, C. H., Aston, J., Petre, V., Duncan, G. H., Morales, F., & Evans, A. C.

41
(2002). A general statistical analysis for fMRI data. NeuroImage, 15(1), 1–15.
https://doi.org/10.1006/nimg.2001.0933
Zeifman, D. M., & St James-Roberts, I. (2017). Parenting the crying infant. Current Opinion in
Psychology, 15, 149–154. https://doi.org/10.1016/j.copsyc.2017.02.009

Central Opercular Cortex

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Table 1
Means and standard deviations for study variables
N Mean (SD)
Average Handgrip Cry 39 .584 (.124)
Average Handgrip Control 40 .597 (.126)
Emotional Reactions Questionnaire 39 11.28 (5.77)
Trait Rating Task 39 4.93 (2.17)
Aggregate Testosterone 33 56.61 (20.65)
Age of Expectant Father 41 31.70 (4.25)
Days Pregnant at Prenatal Visit 41 205.80 (33.01)

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44

Table 3

Neural activation during infant cry (N = 34)
Trial Type Region x y z Z-max Voxels
Main effect

Cry > Control

L Supramarginal
Gyrus
-46 -10 2 5.48 4673
L Planum
Temporale
-50 -36 14 5.12
L Insula -46 4 -6 4.86
L Temporal Pole -46 14 -8 4.68
R Planum
Temporale
62 -10 4 5.93 4430
R Superior
Temporal Gyrus,
posterior
66 -14 -4 5.20
R Heschl’s Gyrus 50 -18 4 4.93
R Planum Porale 48 2 -8 4.81
R Superior
Temporal Gyrus,
anterior
66 0 -6 4.70
R Inferior
Frontal Gyrus
53 29 12 2.58
Note: x, y, and z refer to MNI coordinates; Z-max refers to the peak level of
activation intensity; Voxels refers to the number of voxels in each significant
cluster; L and R refer to left and right hemispheres.

45

46

Table 5
Associations between neural activation on task and prenatal testosterone (N =
32)
Trial Type Region x y z Z-max Voxels
Positive Associations with Testosterone

Cry > Control

R Supramarginal
Gyrus
37 -44 60 4.24 184
L Occipital
Cortex
-34 -68 52 4.18 262
Precuneus
Cortex
-10 -68 62 3.85
Note: x, y, and z refer to MNI coordinates; Z-max refers to the peak level of
activation intensity; Voxels refers to the number of voxels in each significant cluster;
L and R refer to left and right hemispheres.

47

48

R L
L Occipital Cortex
X = -10 Y= -68= Z = 62
Precuneus
X = -34 Y = -68, Z = 52

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Prenatal Response to Infant Cry and Postpartum Parenting Outcomes in Expectant
Fathers

Hannah Khoddam, Diane Goldenberg, Alyssa Morris, & Darby Saxbe

Abstract

The current study tested whether psychological and neural responses to infant cry sounds
predicted self-reported postpartum parenting stress and bonding in a sample of expectant fathers
followed across the transition to parenthood. Expectant fathers who interpreted infant cry more
negatively reported greater parenting stress and weaker father-child bonding at three months
postpartum. However, self-reported negative emotions while listening to cry sounds, as well as
neural reactivity associated with processing infant cry sounds, did not predict postpartum stress
or bonding in this sample of expectant fathers. This study is one of the first longitudinal
investigations of responses to infant cry in expectant fathers across multiple modalities and
postpartum parenting outcomes. Expectant fathers’ negative interpretations of infant cry may
reflect risk for parenting stress and impaired bonding. Prenatal parenting interventions can target
expectant fathers who may interpret infant cry as more hostile or negative

L
L
R L

50
Introduction

Prenatal Response to Infant Cry as a Predictor of Postpartum Parenting

An infant crying can make a parent feel helpless, angry, or frustrated, which may lead to
difficulties bonding with the infant (Oldbury & Adams, 2015). Parents’ responses to infant cry
sounds may indicate risk in the parenting relationship, with the harshest outcome being child
abuse or neglect (Crouch et al., 2008; Frodi & Lamb, 1980). Parenting stress, defined as negative
psychological responses to the obligations of being a parent (Bornstein, 2008), has been found to
predict the potential for child abuse (Crouch & Behl, 2001; Hillson & Kuipur, 1994; Rodriguez
& Green, 1997), and excessive crying has been linked to increased parenting stress (Beebe et al.,
1993). Therefore, responses to a crying infant might be linked to risk along a continuum, with
abuse at the most extreme end and less severe but widely experienced outcomes such as
parenting stress and bonding difficulties at the more normative end.
Recent studies have focused on physiological or biological responses to infant cry in
investigating risk in the parent-child relationship given that “under the skin” measurements may
allow researchers and clinicians to characterize risk before aversive behaviors actually occur
(Compier-de Block et al., 2015; Joosen et al.,, 2013; Out et al., 2012; Riem et al., 2012).
However, these “under the skin” measurements have been less frequently employed to predict
less severe but more prevalent difficulties in adjustment to parenthood. Studies to date have
focused on physiological and behavioral responses (i.e. skin conductance, handgrip modulation,
and neural reactivity) to infant cry, interpretations of the infant cry as negative or hostile, and the
relationship between these responses and eventual parenting behavior. Most studies have focused
on mothers, although fathers also contribute to parenting ( Wilson & Prior, 2011). Few studies
have investigated the prenatal period as an important window in which to capture risk in the

51
parenting relationship once the child is born. The prenatal period, a potentially important
window in which to understand how parenting behaviors may come “online,” is characterized by
hormonal and behavioral changes that may prepare expectant mothers and fathers for parenthood
( Gettler et al., 2011; Saxbe et al., 2017). A paucity of research exists on the potential responses
to infant cry in expectant fathers and how these responses may predict subsequent parenting.
Neural Response to Infant Cry
Previously, we reported that expectant fathers show neural responses to infant cry sounds
in areas associated with speech processing and social cognition (Khoddam et al., in press).
Specifically, we found that when hearing cry sounds, contrasted with frequency-matched white
noise, fathers activated regions in the bilateral temporal lobes including the auditory cortex,
supramarginal gyrus, and insula. However, that study did not include data collected in the
postpartum period. In other work, responses to infant cry have been associated with parenting
behaviors. For example, activation to infant cry in the medial, middle and lateral PFC as well as
sensory and auditory cortices have been positively associated with maternal sensitivity during an
observed interaction with an infant (Atzil et al., 2012; Kim et al., 2010; Musser et al., 2012).
Consistent with this, reduced activation in the medial prefrontal gyrus (mPFG) (involved in
evaluating emotional values of stimuli) and the superior temporal gyrus (STG) (involved in
sensory information processing) to infant cry have been associated with socioeconomic
disadvantage and greater parenting stress in first time mothers, as well as more negative views of
parenting (Kim et al., 2016). Conversely, one study of new fathers found that greater ratings of
aversiveness to cry was associated with greater neural activation in auditory cortices (Li et al.,
2018). Another study found that mothers’ amygdala activation to infant cry was not associated
with their reported irritation during infant cry, but was linked with their insecure attachment

52
(Riem et al., 2012). Taken together, these studies suggest that parenting-related challenges have
been associated with differences in activation in regions including the auditory cortices, mPFC,
STG, IFG, insula and bilateral amygdala, although the direction of these relationships remains
unclear. Of note, no studies have followed expectant fathers into the postpartum period to
examine whether prenatal infant cry responses predict subsequent parenting.
Psychological Response to Infant Cry
Infant crying can challenge the parent-child bond, particularly when the parent feels
rejected by the crying infant (Ellett & Swenson, 2005) or reports feeling ambivalent or less
empathetic toward their baby (Kurth et al., 2011; Megel et al., 2011; Tabuchi & Shimada, 2008).
Additionally, an individual’s interpretations and emotional reactions to infant cry might reflect
risk for harsh or insensitive parenting behaviors (Crouch et al., 2008; Fairbrother, Barr, Pauwels,
Brant, & Green, 2015; Rodriguez, Russa, & Kircher, 2015; Zeskind & Shingler, 1991).
Importantly, how a parent interprets the intentions or personality of a crying infant can be
distinct from the emotions the parent feels while listening to the crying infant. For example, a
parent may feel distressed and bothered that their infant is crying but attribute the crying to
communication rather than being intentionally hostile (Bugental et al., 2002). Additionally, an
individual’s interpretation or emotional reaction to infant cry appears to only occur while parents
and non-parents are listening to infant cry versus a control sound (Bakermans-Kranenburg et al.,
2012b; Caselles & Milner, 2000; Compier-de Block et al., 2015) and thus these responses are
specific to the context of an infant crying. Most of the studies focusing on how an individual
interprets infant cry have focused on individuals who are already parents, or individuals who are
not parents but are shown to be at risk for harsh parenting on a measure of child abuse potential
before they actually have children (Barr et al., 2014; Fairbrother et al., 2015; Ladd, 2010; Milner

53
et al., 1995b). No studies have investigated whether expectant fathers’ interpretation of infant cry
predict the quality of the postpartum parenting relationship or examined more normative
outcomes such as parenting stress or bonding difficulties. Indicators of a difficult parenting
relationship such as parenting stress or bonding difficulties between parent and child may lend
more information to the spectrum of insensitive and aggressive parenting that has a higher base
rate in the general public (Neece et al., 2012).
Current Study
Our previous work (Khoddam et al., in press) examined cross-sectional associations
between expectant fathers’ neural, psychological, and behavioral responses to infant cry. The
current investigation builds on this work and includes a longitudinal design to understand how
fathers’ neural and psychological responses to infant cry predict subsequent parenting risk
(parenting stress and parent-infant bonding) at three months postpartum. We tested three
hypotheses:
1) We expect that expectant fathers who interpret the personality traits of the crying
infant more negatively will report (1) more parenting stress and (2) more distressed
bonding at three months postpartum.
2) We expect that expectant fathers who report that they experience more negative
emotions during infant cry will report (1) more parental stress and (2) more distressed
bonding in early infancy, and
3) We expect that fathers who show greater neural activation to infant cry sounds in brain
regions previously associated with infant cry responding (i.e. STG, bilateral amygdala,
insula and IFG) will report (1) more parental stress and (2) more distressed bonding in
early infancy. We will test these hypotheses using both whole brain and ROI analyses.

54
Methods
Participants
Participants were drawn from the larger longitudinal Hormones Across the Transition to
Childrearing (HATCH) study. The study follows couples from mid-to-late pregnancy across the
first year postpartum. Recruitment occurred via flyers, social media advertising, and word of
mouth. The current study uses data from a prenatal laboratory visit, conducted in mid-to-late
pregnancy; a separate MRI visit that occurred within two weeks of the in-lab visit; and online
questionnaires that were done at approximately three months postpartum. Inclusion criteria
included that couples were cohabiting and pregnant for the first time with a singleton fetus.
Exclusion criteria included any contraindications for MR scanning, daily use of psychotropic
medication, and left-handedness.
Data was available for 41 expectant fathers. Of these 37 expectant fathers participated in
the MRI visit and 34 completed the Parenting Stress Scale (PingSS) and the Parental Bonding
Questionnaire (PBQ). The mean age was 31.7 years old (SD = 4.13 years), the sample was well-
educated with 80% of participants achieving a college degree or higher, and the population was
ethnically diverse, reflective of the population from which it was collected (36% white, 7.3%
black, 26% Hispanic or Latinx, 24% Asian or Pacific Islander, and 5% other).
Procedure
Expectant fathers participated in an in-lab visit scheduled between 25-30 weeks of their
partners’ pregnancy (average weeks pregnant = 29 weeks) and an MRI visit an average of 1.05
(SD = 1.04) weeks later. During the prenatal in-lab visit fathers were asked to listen to the infant
cry noise and complete the Emotional Reactions Questionnaire and Trait-Rating task. During the
MRI visit, fathers completed the same infant cry task in the MRI scanner. Postpartum parenting

55
outcomes were complete via online surveys at approximately three months postpartum (average
of 14.12 weeks, SD = 2.05 weeks).
Protocol and Measures
Neuroimaging protocol. Imaging was performed on a Siemens 3 Tesla MAGNETOM
Prisma scanner using a 20-channel matrix head coil. Functional images were collected using a
T2* weighted Echo Planar (EPI) sequence (32 transversal slices; TR = 2000ms; TE = 25 ms; flip
angle = 90
o
) with a voxel resolution of 3mm x 3mm x 2.5mm. Anatomical images were acquired
using a magnetization prepared rapid acquisition gradient (MPRAGE) sequence (TR = 2530 ms;
TE = 3.13ms; flip angle 10
o
; isotropic voxel resolution 1mm
3
). Task sounds were transmitted
using Siemens V14 sound headphone system.
Infant cry task. Using the same infant cry sounds as a previous study (Riem et al., 2014)
the cry task included six 30-second auditory clips of infant crying interspersed with six 30-
second clips of frequency-matched white noise counterbalanced across participants leading to 12
trials. The stimuli were presented electronically using the E-Prime 3.0 software (Psychology
Software Tools, Pittsburgh, PA) in a block design. The task was six minutes long and
administered in one run.
Emotional reactions questionnaire. Fathers’ negative emotions experienced while
listening to the infant cry was measured using the Emotional Reactions Questionnaire (ERQ;
Milner et al., 1995b). The ERQ asks respondents to indicate how well each adjective describes
their present mood (1, not at all, to 7, extremely well). The negative emotions (bothered, irritated,
annoyed and hostile) subscales were averaged to determine negative emotions after listening to
infant cry (Cronbach’s α = .89 in the current sample, Crouch et al., 2008).

56
Trait rating task. Similarly, after listening to the infant cry each father completed the
Trait Rating Task (Crouch et al., 2008; Riem et al., 2012) to determine how negatively the father
rated the infant who was crying. For the trait-rating task, fathers were asked to rate the infant on
nine traits (i.e. hostile, negative, difficult, friendly, cooperative, sweet, content, lively, attached).
Following the procedures of previously conducted studies the trait ratings were made on a 10-
point scale (ranging from 1 not at all to 10, extremely likely). Positive traits were also included
to increase validity and decrease bias toward negative traits. Trait ratings were averaged across
the three negative traits to obtain a composite negative trait rating (Cronbach’s α = .75 in the
current sample).
Parenting stress. At three months postpartum, parenting stress was measured with the
Parenting Stress Scale (PingSS). The PingSS is an 18-item questionnaire representing positive
themes and negative components of parenthood. The scale is intended to be used for the
assessment of parental stress for both mothers and fathers and for parents of children with and
without clinical problems. It has demonstrated satisfactory level of internal reliability
(Cronbach’s α = .83) and test-retest reliability (Cronbach’s α = .81; Berry & Jones, 1995).
Postpartum bonding questionnaire. Bonding between the father and the child was
measured with the Postpartum Bonding Questionnaire (PBQ), a 25-item questionnaire that
assesses parent-child bonding. Respondents rate how strongly true statements are regarding their
relationship with their infant. Four subscales are identified: Impaired Bonding, Rejection and
Anger, Anxiety about Care and Risk for Abuse (Brockington, et al., 2006; Cronbach’s α = .67-
.97). A total score was used in the current analysis.
Analyses

57
First, all data was explored for outliers and normality of distribution. Means and standard
deviations of all study variables can be found in Table 1. All analyses were run with baby age
and paternal age as covariates.
Neuroimaging analysis. Neural responses to infant cry were analyzed using FEAT
(FMRI Expert Analysis Tool) of FSL (FMRIB’s Software Library, www.FMRIb.ox.ac.uk/fsl;
Smith et al., 2004). First, motion correction using MCFLIRT, non-brain removal, spatial
smoothing (5mm FWHM Gaussian kernel), and registration to T1-weighted images using FSL
FLIRT were done for pre-processing. Next, functional activation was examined with general
linear model analyses. To identify regions involved in the perception of infant crying, contrasts
of cry > white noise and white noise > cry were assessed. Contrasts of parameter estimates
(COPEs) for cry > white noise and white noise > cry sound tested primary hypotheses regarding
response to infant cry versus a frequency matched white noise. First-level COPEs served as
inputs to higher-level group analyses conducted using FLAME to model random-effects
components of mixed-effects variance. Images were thresholded with clusters determined by Z >
2.3 and a cluster-corrected significance threshold of p < .05 (Worsley et al., 2002) to identify
regions that were activated during cry versus white noise across the six blocks for each sound.
Demeaned postpartum bonding (PBQ) and parenting stress (PingSS) were added
(separately) as regressors into the general linear model described above. The first-level contrast
images of cry > white noise and white noise > cry were submitted to second-level whole brain
analysis to determine differences in activation depending on interpretations of the infant as more
negative, and self-reported negative emotions after infant cry. Both positive and negative
contrast weights were tested for each continuous predictor to determine whether it related to
increased or decreased neural response.

58
Additionally, given our a priori hypotheses based on previous published results
(Khoddam et al., in press) and previous literature, ROI analyses of the insula, IFG and STG
(anterior and posterior) and bilateral amygdala were conducted. Parameter estimate values were
converted to percentage signal change values via scaling of the PE or COPE values by (100*) the
peak-peak height of the regressor (or effective regressor in the case of COPEs) and then by
dividing by the mean over time of the filtered functional data. A report was generated using
Featquery with statistics derived from each image’s values within the mask. Percent signal
change for each participant was used in the following analyses. Percent signal change was
extracted from each of the aforementioned areas using anatomically-defined masks created using
the Harvard-Oxford Subcortical Atlas.
Multivariate regression analyses were used to test the relationship between (1) negative
trait ratings of the infant and (2) self-reported emotions after infant cry predicting parental stress
and bonding at three months postpartum, and (3) neural reactivity during infant cry predicting
postpartum parental stress and bonding at three months.
Results
First, all data were explored for outliers and normality of distribution. Two outliers that
were two standard deviations above the mean were identified in the Postpartum Bonding
Questionnaire (PBQ). Analyses remained unchanged with and without including the outliers,
thus all data was included in the final analyses to increase the final sample size. Means and
standard deviations of all study variables are presented in Table 1, and bivariate correlations of
main study variables are shown in Table 2.

59
Hypothesis 1. As seen in Table 3, fathers who made more negative interpretations of the
crying infant (Trait Rating Task) subsequently reported greater postpartum parenting stress (B =
.47, p = .006), and more distressed bonding between father and child (B = .49, p = .004).
Hypothesis 2. Fathers who reported more negative emotions after listening to infant cry
(Emotional Reactions Questionnaire) did not report greater postpartum parenting stress (B = .19,
p = .29) nor distressed bonding (B = .04, p = .81)
Hypothesis 3. Fathers’ prenatal neural activation to infant cry, whether measured via
whole brain or ROI analyses, did not predict their parenting stress or distressed bonding at three
months postpartum (Table 4).
Discussion
The current study tested whether expectant fathers’ psychological and neural responses to
infant cry predicted their subsequent reports of postpartum parenting stress and bonding. Our
hypotheses were partially supported. We found that expectant fathers who rated a crying infant
as more hostile, difficult, and negative showed greater parenting stress and distressed father-child
bonding approximately six months later, at three months postpartum. However, expectant
fathers’ self-reported negative emotions while listening to cry sounds and their neural responses
to infant cry sounds did not predict their postpartum stress or bonding.
Negative trait interpretations of a crying infant have been associated with risk for child
physical abuse (Crouch et al., 2008; Rodriguez, et al., 2015) or have been found in parents who
have previously abused their children (Zeskind & Shingler, 1991), as well as in parents with
difficulty bonding or being empathetic to their children (Oldbury & Adams, 2015). The current
results indicate that expectant fathers who listened to an unknown infant crying and rated that
infant as hostile, negative or difficult reported more difficulty adjusting to the parenting

60
relationship with their own infant. The relationship between more negative interpretations of a
crying infant and difficulties with bonding with the child has been found in groups of parents
who already have children (Oldbury & Adams, 2015). To our knoweldge, this is the first study of
expectant fathers to have found this relationship between interpretations of an unknown infant
before the birth of their child and more difficulties adjusting to parenthood and bonding with the
child postpartum. This finding is in line with previous studies that have consistently found that
parents who view their children as more intentionally hostile or negative, not just while they are
crying, are more likely to act aggressively or negatively toward their children (Bugental et al.,
2002). The current finding adds to this literature and suggests that negative interpretations of an
unfamiliar crying infant may indicate a potential difficulty or likelihood of reacting negatively
toward their own child once the child is born.
Contrary to our hypotheses, the current investigation failed to find a relationship between
expectant fathers’ neural reactivity to infant cry and their subsequent postpartum parenting stress
or distressed bonding. The current study is the first longitudinal study, to our knowledge, to test
the relationship between prenatal neural activation during infant cry and self-reported postpartum
parenting outcomes. Although in a previous study we found that fathers who rated the crying
infant more negatively showed differences in their neural responses to infant cry, these neural
responses did not predict our parenting outcomes. These null results may be due to our small
sample, although 33 subjects is a larger than samples used in most published studies of the
parenting brain. Additionally, it may be that prenatal negative interpretations of the crying infant
is a more robust indicator of potential difficulties in the parent-child relationship than neural
reactivity to infant cry. However, actual observed parenting behaviors, rather than self-report
measures, may have a more direct relationship with neural activity. Therefore, future studies

61
should include observed parenting interactions, as well as include self-report measurements of
negative interpretations of infant cry to validate brain imaging findings.
The current study had a number of limitations. The sample size was small (33 fathers),
albeit larger than the samples used in most published studies of the parenting brain. Another
limitation is that because the sample consisted of expectant fathers, only unfamiliar cry sounds
were played instead of familiar own infant cry sounds. However, this limitation is balanced by
the advantage that the stimuli was standardized across participants and has been used in previous
studies even with participants who were already parents. Additionally, parenting was measured
with self-report questionnaires. Measures of observed parenting, such as either at home or in-lab
recorded parenting interactions that may be coded may have more ecological validity. Given the
exploratory nature of this study and the novel population included, future research should
endeavor for an increased sample size to replicate and clarify these results.
Despite its important limitations, this study is the first to measure both neural and
psychological responses to infant cry in expectant fathers and follow them into the postpartum
period to examine parenting outcomes. The sample was ethnically diverse. The present findings
suggest that expectant fathers who made more negative attributions about a crying infant
subsequently reported greater postpartum parenting stress and difficulties bonding with their
child. Importantly, recent investigations seek to measure “under the skin” risk factors for
distressed parenting or abuse with the assumption that self-report measures are subject to social
desirability and thus will not pick up on risk of insensitive or harsh parenting. However, the
current results suggest that inexpensive and simple prenatal measures, such as the trait rating
task, may indicate risk for postpartum parenting challenges. Expectant fathers who may interpret

62
an unknown crying infant as more hostile or negative may benefit from interventions before the
birth of their child to facilitate their adjustment to fatherhood.
References
Abraham, E., & Feldman, R. (2018). The neurobiology of human allomaternal care; implications
for fathering, coparenting, and children’s social development. Physiology and Behavior,
193(May), 25–34. https://doi.org/10.1016/j.physbeh.2017.12.034
Abraham, E., Hendler, T., Shapira-Lichter, I., Kanat-Maymon, Y., Zagoory-Sharon, O., &
Feldman, R. (2014). Father’s brain is sensitive to childcare experiences. Proceedings of the
National Academy of Sciences. https://doi.org/10.1073/pnas.1402569111
Ahlborg, T., Misvaer, N., & Möller, A. (2009). Perception of marital quality by parents with
small children: A follow-up study when the firstborn is 4 years old. Journal of Family
Nursing, 15(2), 237–263. https://doi.org/10.1177/1074840709334925
Ainsworth, M. S. (1979). Infant-mother attachment. American Psychologist.
https://doi.org/10.1037/0003-066X.34.10.932
Alyousefi-Van Dijk, K., van ‘t Veer, A. E., Meijer, W. M., Lotz, A. M., Rijlaarsdam, J.,
Witteman, J., & Bakermans-Kranenburg, M. J. (2019). Vasopressin differentially affects
handgrip force of expectant fathers in reaction to own and unknown infant faces. Frontiers
in Behavioral Neuroscience, 13. https://doi.org/10.3389/fnbeh.2019.00105
Atzil, S., Hendler, T., Zagoory-Sharon, O., Winetraub, Y., & Feldman, R. (2012). Synchrony and
specificity in the maternal and the paternal brain: Relations to oxytocin and vasopressin.
Journal of the American Academy of Child and Adolescent Psychiatry.
https://doi.org/10.1016/j.jaac.2012.06.008
Baars, B. J., & Gage, N. M. (2010). Cognition, Brain and Consciousness. In Cognition, Brain

63
and Consciousness. https://doi.org/10.1016/C2009-0-01556-6
Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Riem, M. M. E., Tops, M., & Alink, L. R.
A. (2012a). Oxytocin decreases handgrip force in reaction to infant crying in females
without harsh parenting experiences. Social Cognitive and Affective Neuroscience, 7(8),
951–957. https://doi.org/10.1093/scan/nsr067
Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Riem, M. M. E., Tops, M., & Alink, L. R.
A. (2012b). Oxytocin decreases handgrip force in reaction to infant crying in females
without harsh parenting experiences. Social Cognitive and Affective Neuroscience, 7(8),
951–957. https://doi.org/10.1093/scan/nsr067
Barr, R. G., Rivara, F. P., Barr, M., Cummings, P., Taylor, J., Lengua, L. J., & Meredith-Benitz,
E. (2009). Effectiveness of Educational Materials Designed to Change Knowledge and
Behaviors Regarding Crying and Shaken-Baby Syndrome in Mothers of Newborns: A
Randomized, Controlled Trial. Pediatrics, 123(3), 972–980.
https://doi.org/10.1542/peds.2008-0908
Barr, Ronald G., Fairbrother, N., Pauwels, J., Green, J., Chen, M., & Brant, R. (2014). Maternal
frustration, emotional and behavioural responses to prolonged infant crying. Infant Behavior
and Development, 37(4), 652–664. https://doi.org/10.1016/j.infbeh.2014.08.012
Barr, Ronald G., Trent, R. B., & Cross, J. (2006). Age-related incidence curve of hospitalized
Shaken Baby Syndrome cases: Convergent evidence for crying as a trigger to shaking.
Child Abuse and Neglect, 30(1), 7–16. https://doi.org/10.1016/j.chiabu.2005.06.009
Beebe, S. A., Casey, R., & Pinto-Martin, J. (1993). Association of Reported Infant Crying and
Maternal Parenting Stress. Clinical Pediatrics, 32(1), 15–19.
https://doi.org/10.1177/000992289303200103

64
Berry, J. O., & Jones, W. H. (1995). The Parental Stress Scale: Initial Psychometric Evidence.
Journal of Social and Personal Relationships, 12(3), 463–472.
https://doi.org/10.1177/0265407595123009
Blunt Bugental, D., Ellerson, P. C., Lin, E. K., Rainey, B., Kokotovic, A., & O’hara, N. (2002).
A Cognitive Approach to Child Abuse Prevention. https://doi.org/10.1037/0893-
3200.16.3.243
BORNSTEIN, M. (2008). Handbook of Parenting. Pratical Issues in Parenting.
Bos, P. A. (2017). The endocrinology of human caregiving and its intergenerational
transmission. Development and Psychopathology, 29(3), 971–999.
https://doi.org/10.1017/S0954579416000973
Bos, P. A., Hechler, C., Beijers, R., Shinohara, K., Esposito, G., & de Weerth, C. (2018).
Prenatal and postnatal cortisol and testosterone are related to parental caregiving quality in
fathers, but not in mothers. Psychoneuroendocrinology, 97, 94–103.
https://doi.org/10.1016/j.psyneuen.2018.07.013
Bos, P. A., Hermans, E. J., Montoya, E. R., Ramsey, N. F., & van Honk, J. (2010). Testosterone
administration modulates neural responses to crying infants in young females.
Psychoneuroendocrinology, 35(1), 114–121.
https://doi.org/10.1016/j.psyneuen.2009.09.013
Brockington, I. F., Fraser, C., & Wilson, D. (2006). The Postpartum Bonding Questionnaire: A
validation. Archives of Women’s Mental Health, 9(5), 233–242.
https://doi.org/10.1007/s00737-006-0132-1
Brosch, T., Sander, D., & Scherer, K. R. (2007). That baby caught my eye... attention capture by
infant faces. Emotion (Washington, D.C.), 7(3), 685–689. https://doi.org/10.1037/1528-

65
3542.7.3.685
Brown, W. A., Monti, P. M., & Corriveau, D. P. (1978). Serum testosterone and sexual activity
and interest in men. Archives of Sexual Behavior, 7(2), 97–103.
Caselles, C. E., & Milner, J. S. (2000). Evaluations of child transgressions, disciplinary choices,
and expected child compliance in a no-cry and a crying infant condition in physically
abusive and comparison mothers. Child Abuse & Neglect, 24(4), 477–491.
http://www.ncbi.nlm.nih.gov/pubmed/10798838
Church, J. A., Balota, D. A., Petersen, S. E., & Schlaggar, B. L. (2011). Manipulation of length
and lexicality localizes the functional neuroanatomy of phonological processing in adult
readers. Journal of Cognitive Neuroscience. https://doi.org/10.1162/jocn.2010.21515
Compier-de Block, L. H. C. G., Alink, L. R. A., Reijman, S., Werner, C. D., Maras, A., Rijnberk,
C., van IJzendoorn, M. H., & Bakermans-Kranenburg, M. J. (2015). Handgrip force of
maltreating mothers in reaction to infant signals. Child Abuse and Neglect, 40, 124–131.
https://doi.org/10.1016/j.chiabu.2014.03.006
Condon, J. T., Boyce, P., & Corkindale, C. J. (2004). The first-time fathers study: A prospective
study of the mental health and wellbeing of men during the transition to parenthood.
Australian and New Zealand Journal of Psychiatry, 38(1–2), 56–64.
https://doi.org/10.1111/j.1440-1614.2004.01298.x
Coster, D. (2017). Changing Parents’ Behaviour Using a Psycho-Educational Film as a
Preventative Measure to Reduce the Risk of Non-accidental Head Injury. Child Abuse
Review, 26(6), 465–476. https://doi.org/10.1002/car.2469
Crouch, J. L., & Behl, L. E. (2001). Relationships among parental beliefs in corporal
punishment, reported stress, and physical child abuse potential. Child Abuse and Neglect,

66
25(3), 413–419. https://doi.org/10.1016/S0145-2134(00)00256-8
Crouch, J. L., Skowronski, J. J., Milner, J. S., & Harris, B. (2008). Parental responses to infant
crying: The influence of child physical abuse risk and hostile priming. Child Abuse and
Neglect, 32(7), 702–710. https://doi.org/10.1016/j.chiabu.2007.11.002
Del Vecchio, T., Walter, A., & O’Leary, S. G. (2009). Affective and physiological factors
predicting maternal response to infant crying. Infant Behavior and Development, 32(1),
117–122. https://doi.org/10.1016/j.infbeh.2008.10.005
Ellett, M. L. C., & Swenson, M. (2005). Living With a Colicky Infant. Gastroenterology
Nursing, 28(1), 19–25. https://doi.org/10.1097/00001610-200501000-00005
Fairbrother, N., Barr, R. G., Pauwels, J., Brant, R., & Green, J. (2015). Maternal thoughts of
harm in response to infant crying: an experimental analysis. Archives of Women’s Mental
Health, 18(3), 447–455. https://doi.org/10.1007/s00737-014-0471-2
Feldman, R. (2015). Sensitive periods in human social development: New insights from research
on oxytocin, synchrony, and high-risk parenting. Development and Psychopathology, 27(2),
369–395. https://doi.org/10.1017/s0954579415000048
Fleming, A. S., Corter, C., Stallings, J., & Steiner, M. (2002). Testosterone and prolactin are
associated with emotional responses to infant cries in new fathers. Hormones and Behavior,
42(4), 399–413. http://www.ncbi.nlm.nih.gov/pubmed/12488107
Frodi, A. (1985). When Empathy Fails. In Infant Crying (pp. 263–277). Springer US.
https://doi.org/10.1007/978-1-4613-2381-5_12
Frodi, A. M., & Lamb, M. E. (1980). Child abusers’ responses to infant smiles and cries. Child
Development, 51(1), 238–241. http://www.ncbi.nlm.nih.gov/pubmed/7363736
Gallup, A. C., O’Brien, D. T., White, D. D., & Wilson, D. S. (2010). Handgrip strength and

67
socially dominant behavior in male adolescents. Evolutionary Psychology, 8(2), 229–243.
Garcia, R. L., Kenny, D. A., & Ledermann, T. (2015). Moderation in the actor-partner
interdependence model. Personal Relationships, 22(1), 8–29.
https://doi.org/10.1111/pere.12060
Gettler, L. T., McDade, T. W., Feranil, A. B., & Kuzawa, C. W. (2011). Longitudinal evidence
that fatherhood decreases testosterone in human males. Proceedings of the National
Academy of Sciences, 108(39), 16194–16199. https://doi.org/10.1073/pnas.1105403108
Gettler, Lee T., McDade, T. W., Agustin, S. S., Feranil, A. B., & Kuzawa, C. W. (2013). Do
testosterone declines during the transition to marriage and fatherhood relate to men’s sexual
behavior? Evidence from the Philippines. Hormones and Behavior, 64(5), 755–763.
https://doi.org/10.1016/j.yhbeh.2013.08.019
Gray, P. B., Parkin, J. C., & Samms-Vaughan, M. E. (2007). Hormonal correlates of human
paternal interactions: a hospital-based investigation in urban Jamaica. Hormones and
Behavior, 52(4), 499–507. https://doi.org/10.1016/j.yhbeh.2007.07.005
Hickok, G. (2009). The functional neuroanatomy of language. Physics of Life Reviews, 6(3),
121–143. https://doi.org/10.1016/j.plrev.2009.06.001
Hillson, J., Review, N. K.-C. P., & 1994, undefined. (n.d.). A stress and coping model of child
maltreatment. Elsevier. Retrieved March 5, 2020, from
https://www.sciencedirect.com/science/article/pii/0272735894900256
Hirnstein, M., Westerhausen, R., Korsnes, M. S., & Hugdahl, K. (n.d.). Sex differences in
language asymmetry are age-dependent and small: a large-scale, consonant-vowel dichotic
listening study with behavioral and fMRI data. Cortex; a Journal Devoted to the Study of
the Nervous System and Behavior, 49(7), 1910–1921.

68
https://doi.org/10.1016/j.cortex.2012.08.002
Holm, S. (1979). Board of the Foundation of the Scandinavian Journal of Statistics A Simple
Sequentially Rejective Multiple Test Procedure. In Source: Scandinavian Journal of
Statistics (Vol. 6, Issue 2).
Joosen, K. J., Mesman, J., Bakermans-Kranenburg, M. J., Pieper, S., Zeskind, P. S., & Van
Ijzendoorn, M. H. (2013). Physiological Reactivity to Infant Crying and Observed Maternal
Sensitivity. Infancy, 18(3), 414–431. https://doi.org/10.1111/j.1532-7078.2012.00122.x
Joosen, K. J., Mesman, J., Bakermans-Kranenburg, M. J., & van IJzendoorn, M. H. (2013).
Maternal Overreactive Sympathetic Nervous System Responses to Repeated Infant Crying
Predicts Risk for Impulsive Harsh Discipline of Infants. Child Maltreatment, 18(4), 252–
263. https://doi.org/10.1177/1077559513494762
Kenny, D. A. (1996). Models of non-independence in dyadic research. Journal of Social and
Personal Relationships, 13(2), 279–294. https://doi.org/10.1177/0265407596132007
Kenny, D. A., & Judd, C. M. (1986). Consequences of Violating the Independence Assumption
in Analysis of Variance. Psychological Bulletin, 99(3), 422–431.
https://doi.org/10.1037/0033-2909.99.3.422
Khoddam, H., Goldenberg, D., Stoycos, S.A., Horton, K.T., Marshall, N., Cardenas, S.I, Kaplan,
J & Saxbe, D. (in press). How do expectant fathers respond to infant cry? Examining brain
and behavioral responses and the moderating role of testosterone. Social Cognitive and
Affective Neuroscience. https://doi.org/10.1093/scan/nsaa051
Kim, P., Capistrano, C., & Congleton, C. (2016). Socioeconomic disadvantages and neural
sensitivity to infant cry: Role of maternal distress. Social Cognitive and Affective
Neuroscience, 11(10), 1597–1607. https://doi.org/10.1093/scan/nsw063

69
Kim, P., Leckman, J. F., Mayes, L. C., Newman, M. A., Feldman, R., & Swain, J. E. (2010).
Perceived quality of maternal care in childhood and structure and function of mothers’
brain. Developmental Science, 13(4), 662–673. https://doi.org/10.1111/j.1467-
7687.2009.00923.x
Knutson, J. F. (1978). Child abuse as an area of aggression research. Journal of Pediatric
Psychology, 3(1), 20–27. https://doi.org/10.1093/jpepsy/3.1.20
Kuo, P. X., Carp, J., Light, K. C., & Grewen, K. M. (2012). Neural responses to infants linked
with behavioral interactions and testosterone in fathers. Biological Psychology, 91(2), 302–
306. https://doi.org/10.1016/j.biopsycho.2012.08.002
Kurth, E., Kennedy, H. P., Spichiger, E., Hösli, I., & Zemp Stutz, E. (2011). Crying babies, tired
mothers: What do we know? A systematic review. Midwifery, 27(2), 187–194.
https://doi.org/10.1016/j.midw.2009.05.012
Ladd, A. (2010). Infant cries as predictors of child abuse potential: Associations of working
models of attachment, cognitive appraisals and emotional reactions.
http://search.proquest.com/openview/41ebd00d886190638de53e05e95a06cc/1?pq-
origsite=gscholar&cbl=18750&diss=y
Li, T., Horta, M., Mascaro, J. S., Bijanki, K., Arnal, L. H., Adams, M., Barr, R. G., & Rilling, J.
K. (2018). Explaining individual variation in paternal brain responses to infant cries.
Physiology and Behavior, 193(December 2017), 43–54.
https://doi.org/10.1016/j.physbeh.2017.12.033
Lorberbaum, J. P., Newman, J. D., Dubno, J. R., Horwitz, A. R., Nahas, Z., Teneback, C.,
Johnson, M. R., Lydiard, R. B., Ballenger, J. C., & George, M. S. (1998). Feasibility Of
Using fMRI To Study Mothers Responding To Infant Cries. NeuroImage, 7(4), S907.

70
https://doi.org/10.1016/S1053-8119(18)31740-3
Lynch, M. (2003). Consequences of Children ’ s Exposure to Community Violence. 6(4).
Magill-Evans, J., & Harrison, M. J. (2001). Parent-child interactions, parenting stress, and
developmental outcomes at 4 years. Children’s Health Care, 30(2), 135–150.
https://doi.org/10.1207/S15326888CHC3002_4
Malmberg, L.-E., Lewis, S., West, A., Murray, E., Sylva, K., & Stein, A. (2016). The influence
of mothers’ and fathers’ sensitivity in the first year of life on children’s cognitive outcomes
at 18 and 36 months. Child: Care, Health and Development, 42(1), 1–7.
https://doi.org/10.1111/cch.12294
Megel, M. E., Wilson, M. E., Bravo, K., McMahon, N., & Towne, A. (2011). Baby Lost and
Found: Mothers’ Experiences of Infants Who Cry Persistently. Journal of Pediatric Health
Care, 25(3), 144–152. https://doi.org/10.1016/j.pedhc.2009.10.005
Messina, I., Cattaneo, L., Venuti, P., de Pisapia, N., Serra, M., Esposito, G., Rigo, P., Farneti, A.,
& Bornstein, M. H. (2016). Sex-specific automatic responses to infant cries: TMS reveals
greater excitability in females than males in motor evoked potentials. Frontiers in
Psychology, 6(JAN). https://doi.org/10.3389/fpsyg.2015.01909
Milner, J. S., Halsey, L. B., & Fultz, J. (1995a). Empathic responsiveness and affective reactivity
to infant stimuli in high- and low-risk for physical child abuse mothers. Child Abuse and
Neglect, 19(6), 767–780. https://doi.org/10.1016/0145-2134(95)00035-7
Milner, J. S., Halsey, L. B., & Fultz, J. (1995b). Empathic responsiveness and affective reactivity
to infant stimuli in high- and low-risk for physical child abuse mothers. Child Abuse and
Neglect. https://doi.org/10.1016/0145-2134(95)00035-7
Molfese, V. J., Rudasill, K. M., Beswick, J. L., Jacobi-Vessels, J. L., Ferguson, M. C., & White,

71
J. M. (2010). Infant Temperament, Maternal Personality, and Parenting Stress as
Contributors to Infant. In Source: Merrill-Palmer Quarterly (Vol. 56, Issue 1).
Murray, A. D. (1985). Aversiveness is in the Mind of the Beholder. In Infant Crying (pp. 217–
239). Springer US. https://doi.org/10.1007/978-1-4613-2381-5_10
Musser, E. D., Kaiser-Laurent, H., & Ablow, J. C. (2012). The neural correlates of maternal
sensitivity: An fMRI study. Developmental Cognitive Neuroscience, 2(4), 428–436.
https://doi.org/10.1016/j.dcn.2012.04.003
Neece, C. L., Green, S. A., & Baker, B. L. (2012). Parenting stress and child behavior problems:
A transactional relationship across time. American Journal on Intellectual and
Developmental Disabilities, 117(1), 48–66. https://doi.org/10.1352/1944-7558-117.1.48
Oldbury, S., & Adams, K. (2015). The impact of infant crying on the parent-infant relationship.
Community Practitioner, 88(3), 29–34.
Out, D., Bakermans-Kranenburg, M. J., van Pelt, J., & van IJzendoorn, M. H. (2012). Salivary α-
Amylase and Intended Harsh Caregiving in Response to Infant Crying: Evidence for
Physiological Hyperreactivity. Child Maltreatment, 17(4), 295–305.
https://doi.org/10.1177/1077559512464427
Parsons, C. E., Stark, E. A., Young, K. S., Stein, A., & Kringelbach, M. L. (2013).
Understanding the human parental brain: A critical role of the orbitofrontal cortex. Social
Neuroscience, 8(6), 525–543. https://doi.org/10.1080/17470919.2013.842610
Pastore, L., Owens, A., & Raymond, C. (2007). Postpartum sexuality concerns among first-time
parents from one U.S. Academic Hospital. Journal of Sexual Medicine, 4(1), 115–123.
https://doi.org/10.1111/j.1743-6109.2006.00379.x
Persky, H., Lief, H. I., Strauss, D., Miller, W. R., & O’Brien, C. P. (1978). Plasma testosterone

72
level and sexual behavior of couples. Archives of Sexual Behavior, 7(3), 157–173.
https://doi.org/10.1007/BF01542376
Petts, R. J., Shafer, K. M., & Essig, L. (2018). Does Adherence to Masculine Norms Shape
Fathering Behavior? Journal of Marriage and Family, 80(3), 704–720.
https://doi.org/10.1111/jomf.12476
Raboch, J., & Stárka, L. (1973). Reported coital activity of men and levels of plasma
testosterone. Archives of Sexual Behavior, 2(4), 309–315.
https://doi.org/10.1007/BF01541004
Reijman, S., Alink, L. R. A., Block, L. H. C. G. C. de, Werner, C. D., Maras, A., Rijnberk, C.,
IJzendoorn, M. H. va., & Bakermans-Kranenburg, M. J. (2014). Autonomic Reactivity to
Infant Crying in Maltreating Mothers. Child Maltreatment, 19(2), 101–112.
https://doi.org/10.1177/1077559514538115
Riem, M. M. E., Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Out, D., & Rombouts, S.
A. R. B. (2012). Attachment in the brain: Adult attachment representations predict
amygdala and behavioral responses to infant crying. Attachment and Human Development,
14(6), 533–551. https://doi.org/10.1080/14616734.2012.727252
Riem, M. M. E., Voorthuis, A., Bakermans-Kranenburg, M. J., & van Ijzendoorn, M. H. (2014).
Pity or peanuts? Oxytocin induces different neural responses to the same infant crying
labeled as sick or bored. Developmental Science, 17(2), 248–256.
https://doi.org/10.1111/desc.12103
Rilling, J., K., & Young, L., J. (2014). The biology of mammalian parenting and its effect on
offspring social development. Science, 345(6198), 771–776.
https://doi.org/10.1126/science.1252723

73
Rodriguez, C. M., & Green, A. J. (1997). Parenting stress and anger expression as predictors of
child abuse potential. Child Abuse and Neglect, 21(4), 367–377.
https://doi.org/10.1016/S0145-2134(96)00177-9
Rodriguez, C. M., Russa, M. B., & Kircher, J. C. (2015). Analog assessment of frustration
tolerance: Association with self-reported child abuse risk and physiological reactivity. Child
Abuse and Neglect, 46, 121–131. https://doi.org/10.1016/j.chiabu.2015.02.017
Saur, D., Kreher, B. W., Schnell, S., Kummerer, D., Kellmeyer, P., Vry, M.-S., Umarova, R.,
Musso, M., Glauche, V., Abel, S., Huber, W., Rijntjes, M., Hennig, J., & Weiller, C. (2008).
Ventral and dorsal pathways for language. Proceedings of the National Academy of
Sciences. https://doi.org/10.1073/pnas.0805234105
Saxbe, D.E., Edelstein, R. S., Lyden, H. M., Wardecker, B. M., Chopik, W. J., & Moors, A. C.
(2017). Fathers’ decline in testosterone and synchrony with partner testosterone during
pregnancy predicts greater postpartum relationship investment. Hormones and Behavior,
90. https://doi.org/10.1016/j.yhbeh.2016.07.005
Saxbe, Darby E., Schetter, C. D., Simon, C. D., Adam, E. K., & Shalowitz, M. U. (2017). High
paternal testosterone may protect against postpartum depressive symptoms in fathers, but
confer risk to mothers and children. Hormones and Behavior, 95, 103–112.
https://doi.org/10.1016/j.yhbeh.2017.07.014
Silani, G., Lamm, C., Ruff, C. C., & Singer, T. (2013). Right supramarginal gyrus is crucial to
overcome emotional egocentricity bias in social judgments. The Journal of Neuroscience :
The Official Journal of the Society for Neuroscience, 33(39), 15466–15476.
https://doi.org/10.1523/JNEUROSCI.1488-13.2013
Singer, T., & Klimecki, O. M. (2014). Empathy and compassion. In Current Biology.

74
https://doi.org/10.1016/j.cub.2014.06.054
Storey, A. E., & Ziegler, T. E. (2016). Primate paternal care: Interactions between biology and
social experience. Hormones and Behavior, 77, 260–271.
https://doi.org/10.1016/j.yhbeh.2015.07.024
Swain, J. E. (2011). The human parental brain: In vivo neuroimaging. Progress in Neuro-
Psychopharmacology and Biological Psychiatry, 35(5), 1242–1254.
https://doi.org/10.1016/j.pnpbp.2010.10.017
Tabuchi, N., & Shimada, K. (2008). Maternal characteristics and feelings of distress resulting
from the crying of high-risk infants. Japan Journal of Nursing Science, 5(2), 99–108.
https://doi.org/10.1111/j.1742-7924.2008.00108.x
Thijssen, S., Van ’t Veer, A. E., Witteman, J., Meijer, W. M., Van Ijzendoorn, M. H.,
Bakermans-Kranenburg, M. J., & Nl, B. L. (n.d.). Effects of Vasopressin on Neural
Processing of Infant Crying in Expectant Fathers.
https://doi.org/10.1016/j.yhbeh.2018.05.014
van Anders, S. M. (2013). Beyond masculinity: Testosterone, gender/sex, and human social
behavior in a comparative context. In Frontiers in Neuroendocrinology.
https://doi.org/10.1016/j.yfrne.2013.07.001
Van Anders, S. M. (2012). Testosterone and sexual desire in healthy women and men. Archives
of Sexual Behavior, 41(6), 1471–1484. https://doi.org/10.1007/s10508-012-9946-2
Van Anders, S. M., Tolman, R. M., & Volling, B. L. (2012). Baby cries and nurturance affect
testosterone in men. Hormones and Behavior, 61(1), 31–36.
https://doi.org/10.1016/j.yhbeh.2011.09.012
Wilson, Katherine R., & Prior, M. R. (2011). Father involvement and child well-being. Journal

75
of Paediatrics and Child Health, 47(7), 405–407. https://doi.org/10.1111/j.1440-
1754.2010.01770.x
Wilson, Kathryn R., Hansen, D. J., & Li, M. (2011). The traumatic stress response in child
maltreatment and resultant neuropsychological effects. Aggression and Violent Behavior,
16(2), 87–97. https://doi.org/10.1016/j.avb.2010.12.007
Wingfield, J. C., Hegner, R. E., Dufty, A. M., & Ball, G. F. (1990). The “challenge hypothesis’’:
theoretical implications for patterns of testosterone secretion, mating systems, and breeding
strategies.” American Naturalist, 136(6), 829–846. https://doi.org/10.1086/285134
Wingfield, John C. (2017). The challenge hypothesis: Where it began and relevance to humans.
In Hormones and Behavior (Vol. 92, pp. 9–12). Academic Press Inc.
https://doi.org/10.1016/j.yhbeh.2016.11.008
Witteman, J., Van IJzendoorn, M. H., Rilling, J. K., Bos, P. A., Schiller, N. O., & Bakermans-
Kranenburg, M. J. (2019). Towards a neural model of infant cry perception. In
Neuroscience and Biobehavioral Reviews (Vol. 99, pp. 23–32). Elsevier Ltd.
https://doi.org/10.1016/j.neubiorev.2019.01.026
Worsley, K. J., Liao, C. H., Aston, J., Petre, V., Duncan, G. H., Morales, F., & Evans, A. C.
(2002). A general statistical analysis for fMRI data. NeuroImage, 15(1), 1–15.
https://doi.org/10.1006/nimg.2001.0933
Zeifman, D. M., & St James-Roberts, I. (2017). Parenting the crying infant. Current Opinion in
Psychology, 15, 149–154. https://doi.org/10.1016/j.copsyc.2017.02.009

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Table 1
Means and standard deviations for study variables
N Mean (SD)
Parental Bonding Questionnaire (PBQ) 34 .46 (.35)
Parental Stress Scale (PingSS) 34 1.90 (.43)
Emotional Reactions Questionnaire (ERQ) 40 12.03 (6.01)
Trait Rating Task 40 5.42 (1.86)

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78

79

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The Moderating Role of Testosterone on Sexual Satisfaction and Marital Satisfaction
Across the Transition to Parenthood
Hannah Khoddam,

Hannah F. Rasmussen, Geoffrey W. Corner, & Darby Saxbe

In Revision at Hormones and Behavior

Abstract
Declines in relationship satisfaction following the birth of a child have been consistently
found across multiple studies, but less is known about the prenatal factors that predict diminished
relationship satisfaction over the transition to parenthood. In a longitudinal study of couples
transitioning to first-time parenthood, we examined mothers’ and fathers’ prenatal sexual
dissatisfaction and fathers’ prenatal testosterone as predictors of both parents’ changes in
relationship satisfaction from pregnancy to the postpartum period. We found that relationship
satisfaction decreased across the transition to parenthood for fathers, but not mothers. We also
found that neither prenatal testosterone nor prenatal sexual dissatisfaction alone predicted
changes in relationship satisfaction for mothers or fathers. However, paternal prenatal
testosterone moderated the association between mothers’ and fathers’ prenatal sexual
dissatisfaction and mothers’ change in relationship satisfaction, but with opposite effects.
Specifically, in couples with higher-T fathers, greater sexual dissatisfaction in mothers was
related to greater declines in their own relationship satisfaction, and somewhat paradoxically,
greater sexual dissatisfaction in fathers led to less of a decline and in some cases even an
increase in relationship satisfaction in mothers across the transition to parenthood. Additionally,
paternal prenatal testosterone moderated the association between fathers’ prenatal sexual
satisfaction and their own change in relationship satisfaction from pre to postpartum. These

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results point to the importance of investigating the transition to parenthood dyadically and
considering both biological and psychosocial data together when investigating couple dynamics
during this critical life transition.
Introduction
Testosterone (T) is an androgenic steroid hormone that may play an important role in the
transition to fatherhood. Studies have found decreased levels of T both when men become
partnered and when they become fathers, which may reflect greater commitment to the romantic
and co-parenting relationships (Gettler et al., 2011; Gray et al., 2007). However, higher T has
also been associated with more competitive mating advantages, creating a potential tradeoff
between the benefits of high T when finding a mate and low T when becoming a partnered father
(Wingfield et al., 1990; Wingfield, 2017). Given this, paternal T may be especially dynamic in
expectant couples. Further research is needed to investigate how men’s T impacts different
aspects of relationship functioning (e.g., sexual and relationship satisfaction) during the
transition to parenthood, a period when there is greater risk for declines in relationship
satisfaction (Cowan & Cowan, 2000) and when sexual problems may impact relationship
functioning (Vannier & Rosen, 2017).
Sexual Satisfaction and Relationship Satisfaction Across the Transition to Parenthood
Sexual satisfaction is an important aspect of dyadic functioning and relationship
satisfaction (for reviews see: Impetti et al., 2014; Sprecher et al., 2004). It is cross-sectionally
and longitudinally associated with marital stability (Byers, 2005; Laumann et al., 1994; Santtila
et al., 2008; Sprecher, 2002; Yeh et al., 2006) and is a consistently linked with relationship
satisfaction, although the causal directionality of this relationship remains unclear (Impetti et al.,
2014). During pregnancy, biopsychosocial changes may interfere with the sexual and emotional

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relationship between expectant parents (Haugen et al., 2004; Vannier & Rosen, 2017).
Surprisingly, few studies have investigated associations between prenatal sexual satisfaction and
changes in relationship satisfaction across the transition to parenthood. Most of the current
literature focuses only on either prenatal or postpartum associations between sexual satisfaction
and relationship satisfaction. Some studies suggest that sexual distress after pregnancy impacts
relationship functioning in the postpartum period (Cappell et al., 2016; Gray et al., 2015; Vannier
& Rosen, 2017). Other studies have only found these associations in certain contexts. For
example, lower sexual satisfaction may only predict worse marital satisfaction in the postpartum
period in the presence of destructive communication (Litzinger & Gordon, 2005). Ahlborg and
colleagues (2009) found that the sexual functioning of the couple and the amount of cohesion the
couple felt after the birth of a child were the best predictors of marital quality four years later.
Results are mixed when looking at differences between men and women; one study found
that new mothers’ concerns about the sexual relationship predicted lower relationship
satisfaction in both parents, whereas fathers’ concerns were not associated with either partner’s
relationship satisfaction (Schlagintweit et al., 2016). Understanding the dyadic nature of sexual
problems and relationship variables in expectant couples is important; the complex interplay of
these factors may have long-lasting impacts on couple functioning.
Testosterone and Relationship Variables Across the Transition to Parenthood
Although prenatal T may be associated both with sexuality and with perinatal changes in
relationship satisfaction, paternal T levels have not been tested as a moderator of this
relationship. In partnered men lower T levels are cross-sectionally linked with greater
relationship satisfaction and lower likelihood of divorce (Booth & Dabbs, 1993; Edelstein et al.,
2011; Julian & McKenry, 1989; McIntyre et al., 2006; Perini et al., 2012). During the transition

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to parenthood, fathers with steeper declines in T across the prenatal period report higher
postpartum paternal involvement and investment in their relationship with their partner (Gray et
al., 2007; Saxbe et al., 2017). Additionally, men’s T may even be important for women’s
relationship outcomes. For example, men’s T levels appear to be more closely linked with
women’s relationship satisfaction in long term relationships or relationships with children than in
newer relationships (Edelstein et al., 2014). In the context of parenthood, lower postpartum
levels of T in fathers of an infant are associated with greater relationship satisfaction and less risk
of postpartum depression in their partners (Saxbe, et al., 2017). To our knowledge, no studies
have investigated prenatal T and changes in relationship satisfaction from pregnancy to
postpartum.
Testosterone may be an important biological marker for fathering and relationship
investment and behavior (Gray et al., 2007; Gray et al., 2017; Saxbe et al., 2017), but less is
known about the role of testosterone in sexual behaviors, satisfaction, and desire. Although
higher T is purported to increase successful mating challenges (i.e., likelihood of fathering a
child) in human and mammalian males (Wingfield et al., 1990), research focusing on naturally
occurring T in healthy men has often failed to find a relationship between high T and more
frequent sexual behaviors or increased sexual desire (van Anders, 2013; Van Anders, 2012). In
fact, dyadic desire (or the desire to be sexual with another person) is often negatively correlated
with T (van Anders & Dunn, 2009; Van Anders et al., 2007). Research on sexual behavior and
desire across the transition to parenthood is scarce, but there is some evidence to suggest that
fathers’ sexual satisfaction decreases during the first year postpartum (Ahlborg et al., 2009), that
mothers and fathers often have discrepant levels of satisfaction in their sexual relationship
(Condon et al., 2004), and that nearly half of parents state that childcare demands interfere with

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opportunities for sex during the postpartum period (Pastore et al., 2007). It is also possible that a
bidirectional relationship exists between changes in T and sexual behavior across the transition
to fatherhood. Married fathers who showed larger declines in T during the transition to marriage
and first time fatherhood reported lower sexual activity in their committed relationships, and at
the same time, fathers who had more intercourse at baseline and follow-up showed more modest
declines in T (Gettler et al., 2013; Goldey & Van Anders, 2012). Therefore, it is important to
disentangle the relationship between T and sexual satisfaction, behavior, and desire across the
transition to parenthood.
In women, there appears to be a gradual decline in sexual satisfaction across pregnancy, a
sharp drop in the third trimester, a general increase from pregnancy to postpartum, and a gradual
increase from three to twelve months postpartum (Haugen et al., 2004; Moss et al., 1986). One
study found decreases in sexual desire and sexual satisfaction in approximately half of pregnant
woman, which were associated with declines in relationship satisfaction (Vannier & Rosen,
2017). Another cross-sectional assessment of 768 new parents revealed that 36% of mothers and
46% of fathers were sexually dissatisfied at 6 months postpartum (Ahlborg et al., 2009).
Therefore, there is likely substantial variability in the changes that new parents experience in
their sexual satisfaction and functioning.
Overall, it appears that changes in sexual satisfaction are often related to changes in
marital satisfaction. However, to our knowledge, no studies have investigated sexual satisfaction
in the prenatal period in order to predict changes in relationship satisfaction across the transition
to parenthood. Furthermore, few studies have investigated sexual satisfaction in men during their
partner’s pregnancy. Partners may contribute to each other’s levels of sexual and relationship
satisfaction (Fisher et al., 2015; Schlagintweit et al., 2016), which suggests that couples should

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be investigated dyadically, rather than individually. Lastly, lower T in the transition to
parenthood has been linked to greater postpartum relationship investment. However, the role of
fathers’ prenatal T levels has not been explored in the context of links between couples’ sexual
and relationship satisfaction across this critical transition.
Current Study
The current study seeks to investigate the longitudinal and dyadic associations between
fathers’ prenatal T levels, fathers’ and mothers’ sexual dissatisfaction during pregnancy, and
changes in the relationship satisfaction of both partners across the transition to parenthood, as
well as test paternal T as a moderator of the relationship between prenatal sexual dissatisfaction
and change in relationship satisfaction pre to postpartum. Given that both prenatal T and sexual
satisfaction during pregnancy may individually and together contribute to changes in relationship
satisfaction across the transition to parenthood, we tested four hypotheses to better understand
the associations between these aspects of couple functioning:
First, based on previous literature on relationship satisfaction and the transition to
parenthood (Cowan & Cowan, 2000), we hypothesized that relationship satisfaction declines in
mothers and fathers from the prenatal to the postpartum period (H1).
Second, we examined associations between prenatal sexual dissatisfaction and changes in
relationship satisfaction across the transition to parenthood, which included testing within-person
and cross-partner associations. Consistent with previous literature finding a robust and positive
relationship between sexual satisfaction and relationship satisfaction (Byers, 2005; Laumann et
al., 1994; Santtila et al., 2008; Sprecher, 2002; Yeh et al., 2006) we hypothesized that greater
prenatal sexual dissatisfaction would be associated with greater declines in relationship
satisfaction for both partners from the prenatal to the postpartum period (H2).

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Third, given the importance of T in relationship functioning, we examined associations
between fathers’ prenatal T and changes in relationship satisfaction across the transition to
parenthood. We hypothesized that higher prenatal T in fathers would predict decreases in
relationship satisfaction from pregnancy to postpartum in both mothers and fathers (H3).
Lastly, given that certain contexts may moderate the relationship between sexual
satisfaction and relationship satisfaction and testosterone is important in the transition to
parenthood, we predicted that fathers’ prenatal T would moderate the relationship between
prenatal sexual dissatisfaction and changes in relationship satisfaction from pregnancy to
postpartum in both mothers and fathers (H4). The current study’s findings will contribute to our
understanding of these potential prenatal risk factors with regard to declines in relationship
satisfaction across the transition to parenthood.
Methods
Participants were drawn from the larger longitudinal Hormones and Attachment Across
the Transition to Childrearing (HATCH) study. The study, based in a large U.S. West Coast city,
follows couples from mid-to-late pregnancy and across the first year postpartum. Recruiting
occurred via flyers, social media advertising, and word of mouth. The current study used data
from a prenatal laboratory visit conducted in mid-to-late pregnancy and a postpartum visit
occurring approximately six months postpartum. During the prenatal visit, informed consent was
obtained from all participants. All study procedures were approved by the authors’ university’s
Institutional Review Board. Exclusion criteria included non-cohabitating parents, multiparous
parents, parents expecting multiples, and inability to complete study procedures in English.
A total of 68 cohabitating, opposite-sex couples expecting their first child participated in
the study. On average, expectant mothers were 31.9 years old (SD = 4.4, range = 21 to 39), and

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expectant fathers were 34.1 years old (SD = 6.1, range = 22 to 57). The sample was highly
educated, with 80% of participants achieving a college degree or higher, and ethnically diverse,
reflecting the population from which it was recruited (47% White, 7% Black, 20% Hispanic or
Latinx, 18% Asian or Pacific Islander, and 7% other).
Procedure
Participating couples took part in two in-lab visits. The first visit occurred in late
pregnancy (M = 28.6 weeks of pregnancy, SD = 3.6 weeks, range = 24 to 35 weeks). At this visit,
fathers provided three saliva samples over 90 minutes to be assayed for levels of T, and both
partners completed a battery of self-report questionnaires, including measures assessing their
sexual and relationship satisfaction. The second visit occurred approximately 6-8 months after
the birth of the child (M = 7.3 months; SD = 0.9 months; range = 6 to 10 months postpartum),
and at this visit, both partners completed self-report measures of relationship satisfaction.
Measures
Relationship satisfaction. Relationship satisfaction was assessed using the total score
from the Dyadic Adjustment Scale (DAS; Spanier, 1976). The DAS is a 32-item measure of
relationship quality. Items have varying response scales that are scored on a Likert-type scale
and are summed to create a total score ranging from 0 to 151. This measure has demonstrated
strong reliability, including internal consistency (Cronbach’s α = .80 in the current study; Carey,
Spector, Lantinga, & Krauss, 1993). Higher total scores indicate greater dyadic adjustment, and
lower scores suggest conflict within the couple relationship. Changes in relationship satisfaction
across the transition to parenthood were calculated by subtracting parents’ prenatal scores from
their postpartum scores. This change score was used in all path analyses.

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Sexual dissatisfaction. Sexual dissatisfaction was assessed using the Index of Sexual
Satisfaction (ISS; Hudson, Harrison, & Crosscup, 1981). This scale measures the degree of
sexual discord or dissatisfaction in the respondent’s sexual relationship with a partner or spouse.
The score is determined by summing all items, subtracting the number of completed items,
multiplying this figure by 100, and dividing the number of items completed by 6. This will
produce a range from 0 to 100 with higher scores indicating higher levels of discord or
dissatisfaction. This scale has demonstrated strong reliability (Cronbach’s α = .90 in the current
study) and validity (Murphy, Hudson, & Cheung, 2012).
Testosterone. Three saliva samples were collected in cryosafe collection tubes from
fathers using passive drool over 90 minutes during the prenatal in-lab visit. Fathers were
instructed not to eat or drink anything besides water and not to chew gum before collection.
Samples were stored at -80°C before shipment on dry ice to the Technical University of Dresden
to be assayed for levels of T. Samples were assayed in duplicate using a Luminescence
immunoassay (IBL International, Hamburg, Germany) according to manufacturer’s protocol.
The intra- and inter-assay CVs were below 10%. The first 18 participants only provided one
sample of saliva before we modified collection procedures to incorporate more samples. Among
fathers who provided three samples, the level of T in the first sample collected was strongly
correlated with their average level of T across all three samples (r(71) = .76, p < .001). This
suggests that the single measure of T for the first 18 participating fathers is likely a good index of
their T level over the course of the visit. For the remaining fathers in the sample, T was averaged
across the three samples.
Overview of analyses. We calculated descriptive statistics and examined correlations
between key study variables. We conducted a paired samples t-test to test for changes in

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relationship satisfaction from the prenatal to the postpartum period in both mothers and fathers
(H1). The other study hypotheses were tested using path models and Actor-Partner
Interdependence Models (APIM; Kenny, 1996; Kenny & Judd, 1986). The APIM accounts for
the interdependence of dyadic data and allows for simultaneous, independent estimation of the
impact of a person’s characteristics on their own outcomes, actor effects, and the impact of a
person’s characteristics on their partner’s outcomes, partner effects. First, we fit an APIM to
examine the actor and partner effects of mothers’ and fathers’ prenatal sexual dissatisfaction on
mothers’ and fathers’ changes in relationship satisfaction from the prenatal to the postpartum
period (H2). Then, we ran a path model testing associations between fathers’ prenatal T and
mothers’ and fathers’ changes in relationship satisfaction (H3).
We tested Hypothesis 4 using the Actor-Partner Interdependence Moderation Model
(APIMoM; Garcia, Kenny, & Ledermann, 2015). The APIMoM is an extension of the APIM
with the addition of a moderating variable. The APIMoM can take different forms depending on
the type of dyadic moderator tested. The APIMoM with a between-dyads moderator (Figure 1)
has four moderator effects. Fathers’ prenatal T acted as a between-dyads moderator of all actor
and partner effects in the model. This approach was used to test fathers’ prenatal T as a
moderator of the association between sexual dissatisfaction and changes in relationship
satisfaction (H4).
Prior to testing the path models, variables were grand mean centered and standardized.
All models controlled for mothers’ and fathers’ prenatal relationship satisfaction and age,
mothers’ total days pregnant at the prenatal visit, and the infant’s age at the postpartum visit. We
elected to use change scores in order to aid our interpretations of the results within a dyadic
framework. In addition, models that accounted for prenatal relationship satisfaction and

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estimated change as the outcome ensured that both the actor and partner effects in our models
represented longitudinal change (Perry et al., 2017). Controlling for prenatal relationship
satisfaction was also important given the likelihood of cross-sectional associations between
prenatal levels of relationship and sexual satisfaction. This enabled us to isolate the impact of
sexual satisfaction above and beyond its shared variance with overall relationship functioning.
Of note, models run with residualized change scores and without controlling for prenatal
relationship satisfaction produced similar results in terms of magnitude, direction, and
significance of paths of interest. Therefore, for conceptual reasons and to aid in interpretation of
study results, models with change scores as the outcome and prenatal levels as a covariate are
reported. As all participants were members of opposite-sex couples, we treated the dyads as
theoretically distinguishable by sex (Gareau et al., 2016). All path coefficients were estimated
using path models with maximum likelihood estimation.
Results
Descriptive Analyses
The top half of Table 1 presents Ms and SDs for mother and father variables and results
from t-tests examining gender differences. There was a significant difference between mothers’
and fathers’ prenatal relationship satisfaction such that mothers reported higher prenatal
relationship satisfaction than fathers, t(74) = 2.27, p = .03. In addition, fathers were older than
mothers, t(74) = -4.82, p < .001. The bottom half of Table 1 presents Ms and SDs for dyadic
variables.
Bivariate correlations between main study variables are presented in Table 2. Prenatal
age and prenatal levels of sexual dissatisfaction were positively correlated between partners.
Fathers’ prenatal T was negatively correlated with their own postpartum relationship satisfaction.

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Mothers’ change in relationship satisfaction was negatively correlated with their own age.
Neither partner’s change in relationship satisfaction was correlated with fathers’ prenatal T or
either partner’s prenatal sexual dissatisfaction.
Hypothesis 1: Declines in Relationship Satisfaction
Results from two paired samples t-tests demonstrate that relationship satisfaction
declined significantly across the transition to parenthood for fathers t(67) = 3.12, p < .01, but not
for mothers, t(67) = 1.87, p = .07.
Hypothesis 2: Prenatal Sexual Dissatisfaction and Changes in Relationship Satisfaction
The saturated APIM examining associations between mothers’ and fathers’ prenatal
sexual dissatisfaction and their changes in relationship satisfaction is presented in Figure 2.
There were no significant actor or partner effects for either parent’s change in relationship
satisfaction.
Hypothesis 3: Testosterone and Changes in Relationship Satisfaction
Figure 3 presents the results from the path model examining fathers’ prenatal T as a
predictor of mothers’ and fathers’ changes in relationship satisfaction. Fathers’ prenatal T did not
predict fathers’ or mothers’ change in relationship satisfaction.
Hypothesis 4: Testosterone as a Moderator
The path coefficients for the unrestricted model are displayed in Table 3. Mothers’
moderated actor and partner effects were significant, meaning fathers’ prenatal T moderated the
association between mothers’ and fathers’ prenatal sexual dissatisfaction and mothers’ change in
relationship satisfaction.
Next, we examined a series of submodels with different patterns of effects to find the best
fitting model. The estimates of the interaction effects and fit statistics for the unrestricted model

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and the submodels are displayed in Table 4. As a first step, we examined a submodel in which
we constrained all the interaction effects to be equal to zero. The fit indices suggested this model
fit the data, χ
2
(4) = 8.30, p = .08, but the SABIC suggested it was not superior to the
unrestricted model. This indicates that the effects of fathers’ prenatal T as a moderator should
differ from zero. Although the dyads were theoretically distinguishable, we tested whether the
interactions differed by gender. We fit a submodel constraining the interaction effects to be equal
across partners. This model did not fit the data better than the unrestricted model, so we
continued to treat mother and father interaction effects as distinguishable.
To reduce the complexity and aid in the interpretation of the APIMoM in accordance
with guidelines outlined by the authors describing this approach (Garcia et al., 2015), we tested
the model fit for four theoretically relevant APIM patterns separately for mothers and fathers
(Kenny & Cook, 1999; Kenny & Ledermann, 2010). We found that the contrast model was the
best fitting model for mothers and the actor only model was the best fitting model for fathers, χ
2
(2) = 0.14, p = .93. These patterns suggest the pattern of the interaction effects for mothers are
of similar magnitude, but the actor and partner effects are in the opposite direction. This model
met our fit criteria (for a review, see Garcia et al., 2015): (1) it had the smallest SABIC, (2) one
of the coefficients was statistically significant, (3) it fit as well as the unrestricted model, (4) it fit
better than the model with no moderation coefficients, and (5) it fit better than the other three
pattern models. The structural equation of this simpler model for mothers is
(1) Ŷm = -0.15 − 0.09Xm + 0.17Xf − 0.13M − 0.34MXm + 0.34MXf
and for fathers is
(2) Ŷf = -0.57 – 0.32Xf + 0.18Xm − 0.24M + 0.21MXf + (0)MXm.

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As seen in Figure 5, fathers’ and mothers’ prenatal sexual dissatisfaction had little to no
effect on mothers’ change in relationship satisfaction when fathers’ prenatal T was half a
standard deviation below average, but as T increased or decreased, both effects increased in
strength in opposite directions. At high levels of fathers’ prenatal T (M + 1SD), there is a
negative effect of mothers’ prenatal sexual dissatisfaction on their own change in relationship
satisfaction and there is a positive effect of fathers’ prenatal sexual dissatisfaction on mothers’
change in relationship satisfaction. However, at low levels of father’s prenatal T (M - 1SD),
mothers’ prenatal sexual dissatisfaction has a small, positive effect on their own change in
relationship satisfaction whereas fathers’ prenatal sexual dissatisfaction has a small, negative
effect. At all levels of prenatal T, there was a negative effect of fathers’ prenatal sexual
dissatisfaction on their own change in relationship satisfaction. This effect was strongest at low
levels of prenatal T (M - 1SD); as T level increased, the effect of fathers’ prenatal sexual
dissatisfaction approached zero, getting weaker in strength.
Figure 6 depicts plots of the individual interactions and the results of simple slope
analyses. The top half of the figure displays plots for mothers’ change in relationship
satisfaction; panel a.1 shows the association between fathers’ sexual dissatisfaction at low (M -
1SD) and high (M + 1SD) levels and low (M - 1SD) and high (M + 1SD) levels of fathers’
prenatal T and panel a.2 shows the association between mothers’ sexual dissatisfaction at low (M
- 1SD) and high (M + 1SD) levels and fathers’ prenatal T at low (M – 1.24SD) and high (M +
1.24SD). The values of T for panel a.2 were chosen after region of significance analyses because
neither slope was significant at +/-1SD. In both cases, the slopes are significant at high, but not
low, levels of fathers’ prenatal T. However, at high levels of T, the association between fathers’
sexual dissatisfaction and mothers’ change in relationship satisfaction is positive whereas the

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association between mothers’ sexual dissatisfaction and their own change in relationship
satisfaction is negative. In other words, for fathers that have higher prenatal T, high levels of
fathers’ sexual dissatisfaction and low levels of mothers’ sexual dissatisfaction are associated
with increases in mothers’ relationship satisfaction across the transition to parenthood, and as
levels of fathers’ sexual dissatisfaction decrease and mothers’ sexual dissatisfaction increase
mothers’ relationship satisfaction shows a greater decrease from the prenatal to postpartum
period. Panel b depicts associations between fathers’ sexual dissatisfaction at low (M - 1SD) and
high (M + 1SD) levels and fathers’ change in relationship satisfaction moderated by fathers’
prenatal T at low (M - 1SD) and high (M + 1SD) levels. The slope is significant and negative at
low levels of fathers’ prenatal T and is non-significant at high levels. In other words, for fathers
with low levels of prenatal T, low sexual dissatisfaction showed increases in relationship
satisfaction across the transition to parenthood, but higher levels of sexual dissatisfaction were
associated with greater declines in relationship satisfaction.
Discussion
The current study explored potential prenatal risk factors for declines in relationship
satisfaction across the transition to parenthood. First, we found that relationship satisfaction
significantly declined across the transition to parenthood for fathers, but not for mothers.
Notably, prenatal relationship satisfaction was significantly higher in mothers than fathers.
Additionally, higher levels of paternal T were correlated with lower prenatal relationship
satisfaction in mothers and lower postpartum satisfaction in fathers. However, neither prenatal
sexual dissatisfaction nor paternal T alone predicted changes in relationship satisfaction for
either expectant mothers or fathers. However, we found that paternal prenatal T moderated the
associations between mothers’ and fathers’ prenatal sexual dissatisfaction and mothers’ change

95
in relationship satisfaction, but with opposite effects. Specifically, in couples with higher-T
fathers, greater sexual dissatisfaction in mothers was related to greater declines in their own
relationship satisfaction, and somewhat paradoxically, greater sexual dissatisfaction in fathers led
to less of a decline and in some cases even an increase in relationship satisfaction in mothers
across the transition to parenthood. Additionally, we found that for fathers with low levels of
prenatal T, higher levels of sexual dissatisfaction were associated with greater declines in
relationship satisfaction. Low sexual dissatisfaction was associated with smaller declines or even
increases in relationship satisfaction across the transition to parenthood.
Consistent with our expectations and with the literature (Edelstein et al., 2011; Edelstein
et al., 2014; Saxbe et al., 2017), higher prenatal T was corrleated with greater risk to relationship
quality, both in mothers during pregnancy and in fathers at the postpartum assessment. However,
it is notable that there were no main effects of paternal T or prenatal sexual satisfaction in
predicting relationship declines. It is especially surprising given that men’s prenatal T was
correlated with fathers’ postpartum relationship satisfaction in this sample and has been
associated with the sexual and relationship functioning of men in other studies. However, given
the inconsistent findings between T and sexual satisfaction or desire that have been reported in
other previous studies (van Anders, 2013; Van Anders, 2012), it may be that social context is
important for understanding the relationship between T and partnering variables (van Anders,
2013; Van Anders, 2012; Van Anders et al., 2007.) Additionally, unlike prior studies (Perini et
al., 2012), we tested hypotheses using dyadic analyses, which may more accurately represent
how these variables work together in predicting each partner’s relationship decline. Lastly, this is
the first study to investigate prenatal T in conjunction with postpartum relationship outcomes,
which may be a more complicated story than cross-sectional T and relationship functioning.

96
We did find that T moderated the association between prenatal sexual dissatisfaction and
changes in women’s relationship satisfaction, but in opposite directions for mothers and fathers.
Our simple slope analyses revealed that the relationship between each partner’s levels of sexual
dissatisfaction and the mother’s change in relationship satisfaction was only significant when
prenatal T was high. Specifically, in couples with higher-T fathers, greater sexual dissatisfaction
in mothers was related to greater declines in their own relationship satisfaction, and somewhat
paradoxically, greater sexual dissatisfaction in fathers led to less of a decline and in some cases
even an increase in relationship satisfaction in mothers across the transition to parenthood.
Previous research is mixed, but some studies suggest that higher T may be associated with a
stronger sex drive (Gettler et al., 2013). Although highly speculative, it is possible that women’s
high sexual dissatisfaction in the presence of higher paternal T reflected the difficulties of having
intercourse during pregnancy in combination with a high demand from her partner. Thus, it could
be that in relationships with higher-T men, who may emphasize the importance of sex in their
relationship to a greater degree, higher levels of sexual dissatisfaction in women result in more
relationship discord, particularly in the context of a major medical event like childbirth. With
regard to the observed association between fathers’ sexual dissatisfaction and mothers’ changes
it relationship satisfaction for couples with higher-T men, it may be that these men who report
greater sexual dissatisfaction during pregnancy are dissatisfied due to the difficulties of having a
sexual relationship during this period (Weiss & Zverina, 2009). These sexual difficulties and
fathers’ feelings of dissatisfaction could be alleviated six months after the birth of the child, and
at that point, mothers may experience a renewed sense of relationship satisfaction. As the first
study to examine the associations between T, sexual satisfaction, and longitudinal changes in

97
relationship satisfaction in new parents, these findings require replication and additional research
to fully understand their implications.
Lastly, paternal T moderated the effect of fathers’ sexual satisfaction on their own
perinatal relationship satisfaction change, such that lower-T fathers with less sexual
dissatisfaction showed less of a decline, and in some cases even an increase, in their relationship
satisfaction. These results suggest that fathers with lower T who report fewer problems in the
sexual functioning of their relationship may fare better in their relationship as they become
fathers. This result is in line with previous investigations that have found associations between
low T and more commitment to the co-parenting relationship. However, it appears that low T
and satisfaction in other aspects of the relationship (i.e., the sexual relationship) are also
important in predicting changes in relationship satisfaction across the transition to parenthood.
Our study had a number of limitations. For example, our measure of sexual
dissatisfaction relied on self-report and did not cover all facets of sexual behavior and
satisfaction, such as desire and frequency, which may play important roles in disentangling the
relationship between T and sexual satisfaction in predicting changes in relationship satisfaction.
In addition, couples in this study only reported on prenatal sexual dissatisfaction at one timepoint
and were an average of 6.6 months pregnant at the assessment. This may not provide the most
complete picture of how sexual behavior and satisfaction changes from conception to birth
(Vannier & Rosen, 2017). Additionally, other factors not measured in the current investigation,
such as breastfeeding or a difficult childbirth, may additionally impact sexual functioning and
relationship decline in expectant couples. Future studies should work to disentangle prenatal T
and its impact on sexual behavior in expectant couples and how this contributes to relationship
discord between parents.

98
The current study has a number of strengths. First, the use of dyadic data and APIM
analyses allow for the investigation of the effect of each partner’s own sexual satisfaction on
their own change in relationship functioning as well as on their partner’s. An individual level of
analysis cannot sufficiently capture the inherently dyadic nature of couples transitioning to
parenthood (Fisher et al., 2012). Second, this study used a longitudinal design and incorporated
both hormonal and psychosocial data, enabling us to examine the transition to parenthood as it
unfolds and begin to understand how paternal biology may contribute to both mothers’ and
fathers’ changing relationship functioning during this important life transition. Few longitudinal
studies follow couples from pregnancy through the postpartum period and integrate multimodal
data.
In conclusion, we found that lower levels of prenatal paternal T were correlated with
higher relationship satisfaction in both members of the couple—for mothers during pregnancy,
and for fathers in the postpartum period. Additionally, testosterone appeared to moderate the
relationship between prenatal sexual satisfaction in both parents and mothers’ changes in
relationship satisfaction and fathers’ prenatal sexual satisfaction and their own changes in
relationship satisfaction pre to postpartum. The current investigation is an important first step in
understanding how biological factors and psychosocial indices of relationship functioning work
together for couples experiencing the transition to parenthood. It is important for future research
on expectant and new parents to include both fathers and mothers; integrate hormonal, self-
report, and behavioral data; and further explore the mechanisms by which T may shape or
interact with relationship functioning during a challenging and meaningful life transition.

References
Ahlborg, T., Misvaer, N., & Möller, A. (2009). Perception of marital quality by parents with

99
small children: A follow-up study when the firstborn is 4 years old. Journal of Family
Nursing, 15(2), 237–263. https://doi.org/10.1177/1074840709334925
Booth, A., & Dabbs, J. M. (1993). Testosterone and Men’s Marriages. Social Forces, 72(2), 463.
https://doi.org/10.2307/2579857
Byers, E. S. (2005). Relationship satisfaction and sexual satisfaction: A longitudinal study of
individuals in long-term relationships. Journal of Sex Research, 42(2), 113–118.
https://doi.org/10.1080/00224490509552264
Cappell, J., MacDonald, T. K., & Pukall, C. F. (2016). For new mothers, the relationship matters:
Relationship characteristics and postpartum sexuality. The Canadian Journal of Human
Sexuality, 25(2), 126–137. https://doi.org/10.3138/cjhs.252-a5
Carey, M. P., Spector, I. P., Lantinga, L. J., & Krauss, D. J. (1993). Reliability of the Dyadic
Adjustment Scale. Psychological Assessment, 5(2), 238–240. https://doi.org/10.1037/1040-
3590.5.2.238
Condon, J. T., Boyce, P., & Corkindale, C. J. (2004). The first-time fathers study: A prospective
study of the mental health and wellbeing of men during the transition to parenthood.
Australian and New Zealand Journal of Psychiatry, 38(1–2), 56–64.
https://doi.org/10.1111/j.1440-1614.2004.01298.x
Cowan, C. P., & Cowan, P. A. (2000). When Partners Become Parents: The Big Life Change for
Couples.
Edelstein, R. S., Chopik, W. J., & Kean, E. L. (2011). Sociosexuality moderates the association
between testosterone and relationship status in men and women. Hormones and Behavior,
60, 248–255. https://doi.org/10.1016/j.yhbeh.2011.05.007
Fisher, W. A., Donahue, K. L., Long, J. S., Heiman, J. R., Rosen, R. C., & Sand, M. S. (2015).

100
Individual and Partner Correlates of Sexual Satisfaction and Relationship Happiness in
Midlife Couples: Dyadic Analysis of the International Survey of Relationships. Archives of
Sexual Behavior, 44(6), 1609–1620. https://doi.org/10.1007/s10508-014-0426-8
Garcia, R. L., Kenny, D. A., & Ledermann, T. (2015). Moderation in the actor-partner
interdependence model. Personal Relationships, 22(1), 8–29.
https://doi.org/10.1111/pere.12060
Gareau, A., Fitzpatrick, J., Gaudreau, P., & Lafontaine, M.-F. (2016). Analysing, Interpreting,
and Testing the Invariance of the Actor-Partner Interdependence Model. The Quantitative
Methods for Psychology, 12(2), 101–113. https://doi.org/10.20982/tqmp.12.2.p101
Gettler, L. T., McDade, T. W., Feranil, A. B., & Kuzawa, C. W. (2011). Longitudinal evidence
that fatherhood decreases testosterone in human males. Proceedings of the National
Academy of Sciences, 108(39), 16194–16199. https://doi.org/10.1073/pnas.1105403108
Gettler, Lee T., McDade, T. W., Agustin, S. S., Feranil, A. B., & Kuzawa, C. W. (2013). Do
testosterone declines during the transition to marriage and fatherhood relate to men’s sexual
behavior? Evidence from the Philippines. Hormones and Behavior, 64(5), 755–763.
https://doi.org/10.1016/j.yhbeh.2013.08.019
Goldey, K. L., & Van Anders, S. M. (2012). Sexual thoughts: Links to testosterone and cortisol
in men. Archives of Sexual Behavior, 41(6), 1461–1470. https://doi.org/10.1007/s10508-
011-9858-6
Gray, P B, Parkin, J. C., & Samms-Vaughan, M. E. (2007). Hormonal correlates of human
paternal interactions: a hospital-based investigation in urban Jamaica. Hormones and
Behavior, 52(4), 499–507. https://doi.org/10.1016/j.yhbeh.2007.07.005
Gray, Peter B., Reece, J. A., Coore-Desai, C., Dinnall-Johnson, T., Pellington, S., & Samms-

101
Vaughan, M. (2015). Sexuality among fathers of newborns in Jamaica. BMC Pregnancy
and Childbirth, 15(1). https://doi.org/10.1186/s12884-015-0475-6
Gray, Peter B., Reece, J., Coore-Desai, C., Dinall, T., Pellington, S., & Samms-Vaughan, M.
(2017). Testosterone and Jamaican Fathers: Exploring Links to Relationship Dynamics and
Paternal Care. Human Nature, 28(2), 201–218. https://doi.org/10.1007/s12110-016-9283-6
Haugen, E. N., Schmutzer, P. A., & Wenzel, A. (2004). Sexuality and the partner relationship
during pregnancy and the postpartum period. The Handbook of Sexuality in Close
Relationships., 411–435.
Hudson, W. W., Harrison, D. F., & Crosscup, P. C. (1981). A Short-Form Scale to Measure
Sexual Discord in Dyadic Relationships. The Journal of Sex Research, 17(2), 157–174.
https://doi.org/10.1080/00224498109551110
Impett, E. A., Muise, A., & Peragine, D. (2014). APA Handbook of Sexuality and Psychology.
Volume 1: Person-Based Approaches.
Julian, T., & McKenry, P. C. (1989). Relationship of testosterone to men’s family functioning at
mid‐life: A research note. Aggressive Behavior, 15(4), 281–289.
https://doi.org/10.1002/ab.2480150403
Kenny, D. A. (1996). Models of non-independence in dyadic research. Journal of Social and
Personal Relationships, 13(2), 279–294. https://doi.org/10.1177/0265407596132007
Kenny, D. A., & Cook, W. (1999). Partner effects in relationship research: Conceptual issues,
analytic difficulties, and illustrations. Personal Relationships, 6(4), 433–448.
https://doi.org/10.1111/j.1475-6811.1999.tb00202.x
Kenny, D. A., & Judd, C. M. (1986). Consequences of Violating the Independence Assumption
in Analysis of Variance. Psychological Bulletin, 99(3), 422–431.

102
https://doi.org/10.1037/0033-2909.99.3.422
Kenny, D. A., & Ledermann, T. (2010). Detecting, measuring, and testing dyadic patterns in the
actor-partner interdependence model. Journal of Family Psychology : JFP : Journal of the
Division of Family Psychology of the American Psychological Association (Division 43),
24(3), 359–366. https://doi.org/10.1037/a0019651
Laumann, E. O., Gagnon, J. H., Michael, R. T., & Michaels, S. (1994). The social organization
of sexuality. University of Chicago Press.
Litzinger, S., & Gordon, K. C. (2005). Exploring relationships among communication, sexual
satisfaction, and marital satisfaction. Journal of Sex and Marital Therapy, 31(5), 409–424.
https://doi.org/10.1080/00926230591006719
McIntyre, M., Gangestad, S. W., Gray, P. B., Chapman, J. F., Burnham, T. C., O’Rourke, M. T.,
& Thornhill, R. (2006). Romantic involvement often reduces men’s testosterone levels - But
not always: The moderating role of extrapair sexual interest. Journal of Personality and
Social Psychology, 91(4), 642–651. https://doi.org/10.1037/0022-3514.91.4.642
Moss, P., Bolland, G., Foxman, R., & Owen, C. (1986). Marital Relations During the Transition
to Parenthood. Journal of Reproductive and Infant Psychology, 4(1–2), 57–67.
https://doi.org/10.1080/02646838608408665
Murphy, G. J., Hudson, W. W., & Cheung, P. P. L. (2012). Marital and sexual discord among
older couples. Social Work Research and Abstracts, 16(1), 11–16.
https://doi.org/10.1093/swra/16.1.11
Pastore, L., Owens, A., & Raymond, C. (2007). Postpartum sexuality concerns among first-time
parents from one U.S. Academic Hospital. Journal of Sexual Medicine, 4(1), 115–123.
https://doi.org/10.1111/j.1743-6109.2006.00379.x

103
Perini, T., Ditzen, B., Fischbacher, S., & Ehlert, U. (2012). Testosterone and relationship quality
across the transition to fatherhood. Biological Psychology, 90(3), 186–191.
https://doi.org/10.1016/j.biopsycho.2012.03.004
Santtila, P., Wager, I., Witting, K., Harlaar, N., Jern, P., Johansson, A., Varjonen, M., &
Sandnabba, N. K. (2008). Discrepancies between sexual desire and sexual activity: Gender
differences and associations with relationship satisfaction. Journal of Sex and Marital
Therapy, 34(1), 31–44. https://doi.org/10.1080/00926230701620548
Saxbe, D.E., Edelstein, R. S., Lyden, H. M., Wardecker, B. M., Chopik, W. J., & Moors, A. C.
(2017). Fathers’ decline in testosterone and synchrony with partner testosterone during
pregnancy predicts greater postpartum relationship investment. Hormones and Behavior,
90. https://doi.org/10.1016/j.yhbeh.2016.07.005
Saxbe, Darby E., Schetter, C. D., Simon, C. D., Adam, E. K., & Shalowitz, M. U. (2017). High
paternal testosterone may protect against postpartum depressive symptoms in fathers, but
confer risk to mothers and children. Hormones and Behavior, 95, 103–112.
https://doi.org/10.1016/j.yhbeh.2017.07.014
Schlagintweit, H. E., Bailey, K., & Rosen, N. O. (2016). A New Baby in the Bedroom:
Frequency and Severity of Postpartum Sexual Concerns and Their Associations With
Relationship Satisfaction in New Parent Couples. Journal of Sexual Medicine, 13(10),
1455–1465. https://doi.org/10.1016/j.jsxm.2016.08.006
Spanier, G. B. (1976). Measuring Dyadic Adjustment: New Scales for Assessing the Quality of
Marriage and Similar Dyads. Journal of Marriage and the Family, 38(1), 15.
https://doi.org/10.2307/350547
Sprecher, S., Cate, R. ., Harvey, J. ., & Wenzel, A. (2004). Sexual satisfaction and sexual

104
expression as predictors of relationship satisfaction and stability. In The handbook of
sexuality in close relationships. (pp. 235–256).
Sprecher, Susan. (2002). Sexual satisfaction in premarital relationships: associations with
satisfaction, love, commitment, and stability. Journal of Sex Research, 39(3), 190–196.
https://doi.org/10.1080/00224490209552141
van Anders, S. M. (2013). Beyond masculinity: Testosterone, gender/sex, and human social
behavior in a comparative context. In Frontiers in Neuroendocrinology.
https://doi.org/10.1016/j.yfrne.2013.07.001
Van Anders, S. M. (2012). Testosterone and sexual desire in healthy women and men. Archives
of Sexual Behavior, 41(6), 1471–1484. https://doi.org/10.1007/s10508-012-9946-2
van Anders, S. M., & Dunn, E. J. (2009). Are gonadal steroids linked with orgasm perceptions
and sexual assertiveness in women and men? Hormones and Behavior, 56(2), 206–213.
https://doi.org/10.1016/j.yhbeh.2009.04.007
Van, S. M., Peter, A., Gray, B., & Anders, V. (n.d.). Hormones and Human Partnering.
Vannier, S. A., & Rosen, N. O. (2017). Sexual Distress and Sexual Problems During Pregnancy:
Associations With Sexual and Relationship Satisfaction. Journal of Sexual Medicine, 14(3),
387–395. https://doi.org/10.1016/j.jsxm.2016.12.239
Weiss, P., & Zverina, J. (2009). Human sexuality during pregnancy and the postpartum period.
In researchgate.net. www.bmj.sk.
Wingfield, J. C., Hegner, R. E., Dufty, A. M., & Ball, G. F. (1990). The “challenge hypothesis’’:
theoretical implications for patterns of testosterone secretion, mating systems, and breeding
strategies.” American Naturalist, 136(6), 829–846. https://doi.org/10.1086/285134
Wingfield, John C. (2017). The challenge hypothesis: Where it began and relevance to humans.

105
In Hormones and Behavior (Vol. 92, pp. 9–12). Academic Press Inc.
https://doi.org/10.1016/j.yhbeh.2016.11.008
Yeh, H. C., Lorenz, F. O., Wickrama, K. A. S., Conger, R. D., & Elder, G. H. (2006).
Relationships among sexual satisfaction, marital quality, and marital instability at midlife.
Journal of Family Psychology, 20(2), 339–343. https://doi.org/10.1037/0893-3200.20.2.339

106
Table 1
Means and standard deviations for study variables

Mothers Fathers
M (SD) M (SD) t-test
b

Partner Variables
Change in Relationship Satisfaction
a
-2.40 (10.57) -3.49 (9.21) 0.72
Prenatal Sexual Dissatisfaction 65.35 (21.01) 67.66 (21.62) -1.05
Prenatal Relationship Satisfaction 137.19 (11.35) 134.24 (13.45) 2.34*
Postpartum Relationship Satisfaction 134.79 (13.95) 130.75 (15.21) 2.83**
Prenatal Age 31.21 (4.42) 33.19 (5.87) -3.53**
Mean (SD)
Dyadic variables
Father’s Prenatal Testosterone 56.05 (25.82)
Days Pregnant 204.05 (26.75)
Baby Age at Postpartum visit 28.71 (2.80)
Note.
a
prenatal relationship satisfaction subtracted from postpartum relationship satisfaction, i.e.,
positive values indicate increases in relationship satisfaction;
b
paired samples t-test evaluating
gender differences.

107

108
Table 3

Unrestricted model with a between-dyads moderator

Effect Coefficient SE p
Intercept

Mother -0.14 0.13 .28
Father -0.56 0.11 .00
Actor effects of sexual dissatisfaction

Mother -0.09 0.16 .58
Father -0.31 0.16 .05
Partner effects of sexual dissatisfaction

Mother 0.17 0.18 .33
Father 0.18 0.14 .22
Prenatal Testosterone

Mother -0.12 0.13 .33
Father -0.24 0.11 .03
Actor sexual dissatisfaction by T

Mother -0.38 0.18 .04
Father 0.24 0.14 .10
Partner sexual dissatisfaction by T

Mother 0.35 0.16 .03
Father -0.06 0.16 .73
Note. N = 68 couples; Bold = significant paths. The outcome variable is change in relationship
satisfaction from prenatal to postpartum measurements. Model uses standardized x and y
variables and controls for prenatal relationship satisfaction, prenatal age, days pregnant and
baby age postpartum.

109

110
Figure 1
The actor-partner interdependence model with a between-dyads moderator.

111

112

113

114

115

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General Discussion
Despite a burgeoning literature on neuroendocrine and neurological factors purported to
underlie paternal readiness, many gaps in the literature exist. To this end, the current set of
studies aimed to (1) characterize the neurobiological, psychological, and behavioral response to
infant cry in expectant fathers, and test the moderating role of testosterone in these responses, (2)
test the relationship between reactivity to infant cry and parenting outcomes in new fathers at
three months postpartum and (3), investigate measurable prenatal factors (i.e. testosterone,
sexual satisfaction) that may predict postpartum relationship functioning in new mothers and
fathers.
Responses to an infant crying have been used as a proxy for risk of harsh, aggressive, or
insensitive parenting responses toward an infant, and have been suggested as a target for
interventions geared towards expectant parents (Barr et al., 2009). Study 1 aimed to further
examine and characterize the neural, psychological, and behavioral responses to infant cry in a
community sample of expectant fathers. We found that expectant fathers responded to infant cry
versus a frequency-matched white noise with increased activation in bilateral areas of the
temporal lobe involved in processing speech sounds and social and emotional stimuli. Handgrip
force, which has been used to measure parents’ reactivity to cry sounds in previous studies, did
not differentiate cry from white noise within this sample, nor was it related to any other
responses to an infant crying. Expectant fathers with higher prenatal testosterone additionally
showed greater activation in the supramarginal gyrus, left occipital lobe and precuneus cortex to
cry sounds. This is the first investigation to look at neural activation to infant cry in expectant
fathers and baseline prenatal T levels. Additionally, greater activation to infant cry in association

117
with higher T level may indicate a hyperreactivity to infant cry in line with previous
investigations that have found a positive relationship between neural activation and greater
reports of irritation to infant cry in new fathers (Li et al., 2008). This study suggests that infant
cry is a salient stimulus for expectant fathers, even though they have not yet participated in
caregiving for their own infants. Fine-tuning of these processes may occur with socialization
with the infant as has been previously hypothesized (Abraham et al., 2014; Atzil et al., 2012,
Feldman, 2015). However, the current study suggests that expectant fathers show processing of
infant cry that is similar to mothers and fathers whose child is already born. Future studies would
benefit from including expectant fathers at different stages of pregnancy, or age-matched non-
fathers to understand whether similar brain responses emerge in non-parents and if not, when and
how these brain responses come “online” during pregnancy.
Study 2 builds on the cross-sectional design of Study 1 and includes a longitudinal
assessment of the responses to infant cry measured in Study 1. Previous studies have found that a
caregiver’s response to a crying infant may indicate risk in the parenting relationship, with the
harshest outcome being child abuse or neglect (Crouch et al., 2008; Frodi & Lamb, 1980).
However, less severe yet more common outcomes in the parenting relationship, such as
difficulties bonding and parenting stress may also be associated with responses to a crying infant
(Beebe et al., 1993). Therefore, this study examined the association between neural and
psychological responses to infant cry and parental bonding and parenting stress in new fathers at
three months postpartum. We found that expectant fathers who rated the crying infant as more
hostile, difficult, and negative showed greater parenting stress and distressed bonding in the
father-child relationship approximately six months later. However, fathers’ self-reported

118
negative emotions while listening to cry sounds and their neural responses to infant cry sounds
did not predict their postpartum stress or bonding.
This is the first study, to our knowledge, to examine prenatal multi-modal responses to
infant cry and postpartum parenting across the transition to parenthood. The present findings
suggest that a father who views a crying infant more negatively may be at risk for developing
problems in the father-child relationship, whereas the neural responses to infant cry may not be
as sensitive a marker of these specific risks. Future studies should endeavor to include self-report
measures of reactivity to infant cry in predicting postpartum parenting along with neural or
physiological measures of reactivity and include observable measures of parenting behavior to
expand on the current results. Additionally, expectant fathers who may interpret infant cry as
more hostile or negative may be targeted for intervention before the birth of their child to
facilitate their adjustment to fatherhood.
Study 3 investigated measurable prenatal factors (i.e. paternal testosterone level, and
couple sexual satisfaction) that may predict declines in relationship satisfaction over the
transition to parenthood, as well as tested paternal prenatal T as a moderator of the relationship
between prenatal sexual satisfaction and changes in relationship satisfaction pre to postpartum.
Using Actor-Partner Interdependence Modeling (APIM) and Actor-Partner Interdependence
Moderation Modeling (APIMoM), both couple’s data was explored dyadically rather than
individually. As expected, both mother’s and father’s relationship satisfaction declined pre to
postpartum, however only fathers’ declines were significant. Notably, no individual variables
predicted declines in relationship satisfaction pre to postpartum in mothers or fathers. However,
we found that paternal prenatal T moderated the associations between mothers’ and fathers’
prenatal sexual dissatisfaction and mothers’ change in relationship satisfaction, but with opposite

119
effects. In addition, we found that for fathers with low levels of prenatal T, higher levels of
sexual dissatisfaction were associated with greater declines in relationship satisfaction. Low
sexual dissatisfaction was associated with smaller declines or even increases in relationship
satisfaction across the transition to parenthood. The current investigation is an important first
step in understanding how biological factors and psychosocial indices of relationship functioning
work together for couples experiencing the transition to parenthood. It is important for future
research on expectant and new parents to include both fathers and mothers; integrate hormonal,
self-report, and behavioral data; and further explore the mechanisms by which T may shape or
interact with relationship functioning during a challenging and meaningful life transition.
Conclusions
The transition to parenthood is one of the most transformative developmental periods in
an adult’s life (Cowan & Cowan, 2000; Rossi, 1968). Over the last few decades fatherhood has
taken on a new meaning as men become more involved in the caretaking of their children and
play a vitally important role in their children’s development. Although there is a growing
literature on the neurobiological and neuroendocrine factors that may underlie paternal readiness,
their exist many gaps in understanding how, when, and why a father becomes a father. The
current dissertation portfolio is an attempt to close a few of the gaps in this literature, namely
which measurable prenatal factors may influence postpartum relationship functioning between
new fathers and their partner’s and new fathers and their children. The current findings elucidate
the moderating role of men’s prenatal T on sexual satisfaction and declines in relationship
satisfaction across the transition to parenthood, as well as in prenatal neural reactivity to infant
cry. Additionally, the current set of studies demonstrated that neural reactivity to infant cry may
be similar in expectant fathers as new mothers and fathers and may also be related to how

120
negatively fathers view an infant crying in the prenatal period. Handgrip modulation was not
found to be a sensitive marker of response to infant cry, but negative interpretations of an infant
crying were found to have robust associations with parenting stress and difficulty bonding in the
postpartum period.
References for General Introduction and Discussion

Ahlborg, T., Misvaer, N., & Möller, A. (2009). Perception of marital quality by parents with
small children: A follow-up study when the firstborn is 4 years old. Journal of Family
Nursing, 15(2), 237–263. https://doi.org/10.1177/1074840709334925
Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Riem, M. M. E., Tops, M., & Alink, L. R.
A. (2012a). Oxytocin decreases handgrip force in reaction to infant crying in females
without harsh parenting experiences. Social Cognitive and Affective Neuroscience, 7(8),
951–957. https://doi.org/10.1093/scan/nsr067
Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Riem, M. M. E., Tops, M., & Alink, L. R.
A. (2012b). Oxytocin decreases handgrip force in reaction to infant crying in females
without harsh parenting experiences. Social Cognitive and Affective Neuroscience, 7(8),
951–957. https://doi.org/10.1093/scan/nsr067
Barr, R. G., Rivara, F. P., Barr, M., Cummings, P., Taylor, J., Lengua, L. J., & Meredith-Benitz,
E. (2009). Effectiveness of Educational Materials Designed to Change Knowledge and
Behaviors Regarding Crying and Shaken-Baby Syndrome in Mothers of Newborns: A
Randomized, Controlled Trial. Pediatrics, 123(3), 972–980.
https://doi.org/10.1542/peds.2008-0908
Brown, W. A., Monti, P. M., & Corriveau, D. P. (1978). Serum testosterone and sexual activity

121
and interest in men. Archives of Sexual Behavior, 7(2), 97–103.
Compier-de Block, L. H. C. G., Alink, L. R. A., Reijman, S., Werner, C. D., Maras, A., Rijnberk,
C., van IJzendoorn, M. H., & Bakermans-Kranenburg, M. J. (2015). Handgrip force of
maltreating mothers in reaction to infant signals. Child Abuse and Neglect, 40, 124–131.
https://doi.org/10.1016/j.chiabu.2014.03.006
Condon, J. T., Boyce, P., & Corkindale, C. J. (2004). The first-time fathers study: A prospective
study of the mental health and wellbeing of men during the transition to parenthood.
Australian and New Zealand Journal of Psychiatry, 38(1–2), 56–64.
https://doi.org/10.1111/j.1440-1614.2004.01298.x
Cowan, C. P., & Cowan, P. A. (2000). When Partners Become Parents: The Big Life Change for
Couples.
Crouch, J. L., Skowronski, J. J., Milner, J. S., & Harris, B. (2008). Parental responses to infant
crying: The influence of child physical abuse risk and hostile priming. Child Abuse and
Neglect, 32(7), 702–710. https://doi.org/10.1016/j.chiabu.2007.11.002
Garcia, R. L., Kenny, D. A., & Ledermann, T. (2015). Moderation in the actor-partner
interdependence model. Personal Relationships, 22(1), 8–29.
https://doi.org/10.1111/pere.12060
Gettler, L. T., McDade, T. W., Feranil, A. B., & Kuzawa, C. W. (2011). Longitudinal evidence
that fatherhood decreases testosterone in human males. Proceedings of the National
Academy of Sciences, 108(39), 16194–16199. https://doi.org/10.1073/pnas.1105403108
Gettler, Lee T., McDade, T. W., Agustin, S. S., Feranil, A. B., & Kuzawa, C. W. (2013). Do
testosterone declines during the transition to marriage and fatherhood relate to men’s sexual
behavior? Evidence from the Philippines. Hormones and Behavior, 64(5), 755–763.

122
https://doi.org/10.1016/j.yhbeh.2013.08.019
Gray, P. B., Parkin, J. C., & Samms-Vaughan, M. E. (2007). Hormonal correlates of human
paternal interactions: a hospital-based investigation in urban Jamaica. Hormones and
Behavior, 52(4), 499–507. https://doi.org/10.1016/j.yhbeh.2007.07.005
Joosen, K. J., Mesman, J., Bakermans-Kranenburg, M. J., Pieper, S., Zeskind, P. S., & Van
Ijzendoorn, M. H. (2013). Physiological Reactivity to Infant Crying and Observed Maternal
Sensitivity. Infancy, 18(3), 414–431. https://doi.org/10.1111/j.1532-7078.2012.00122.x
Kenny, D. A. (1996). Models of non-independence in dyadic research. Journal of Social and
Personal Relationships, 13(2), 279–294. https://doi.org/10.1177/0265407596132007
Kenny, D. A., & Judd, C. M. (1986). Consequences of Violating the Independence Assumption
in Analysis of Variance. Psychological Bulletin, 99(3), 422–431.
https://doi.org/10.1037/0033-2909.99.3.422
Li, T., Horta, M., Mascaro, J. S., Bijanki, K., Arnal, L. H., Adams, M., Barr, R. G., & Rilling, J.
K. (2018). Explaining individual variation in paternal brain responses to infant cries.
Physiology and Behavior, 193(December 2017), 43–54.
https://doi.org/10.1016/j.physbeh.2017.12.033
Magill-Evans, J., & Harrison, M. J. (2001). Parent-child interactions, parenting stress, and
developmental outcomes at 4 years. Children’s Health Care, 30(2), 135–150.
https://doi.org/10.1207/S15326888CHC3002_4
Molfese, V. J., Rudasill, K. M., Beswick, J. L., Jacobi-Vessels, J. L., Ferguson, M. C., & White,
J. M. (2010). Infant Temperament, Maternal Personality, and Parenting Stress as
Contributors to Infant. In Source: Merrill-Palmer Quarterly (Vol. 56, Issue 1).
Oldbury, S., & Adams, K. (2015). The impact of infant crying on the parent-infant relationship.

123
Community Practitioner, 88(3), 29–34.
Pastore, L., Owens, A., & Raymond, C. (2007). Postpartum sexuality concerns among first-time
parents from one U.S. Academic Hospital. Journal of Sexual Medicine, 4(1), 115–123.
https://doi.org/10.1111/j.1743-6109.2006.00379.x
Persky, H., Lief, H. I., Strauss, D., Miller, W. R., & O’Brien, C. P. (1978). Plasma testosterone
level and sexual behavior of couples. Archives of Sexual Behavior, 7(3), 157–173.
https://doi.org/10.1007/BF01542376
Petts, R. J., Shafer, K. M., & Essig, L. (2018). Does Adherence to Masculine Norms Shape
Fathering Behavior? Journal of Marriage and Family, 80(3), 704–720.
https://doi.org/10.1111/jomf.12476
Raboch, J., & Stárka, L. (1973). Reported coital activity of men and levels of plasma
testosterone. Archives of Sexual Behavior, 2(4), 309–315.
https://doi.org/10.1007/BF01541004
Riem, M. M. E., Bakermans-Kranenburg, M. J., van IJzendoorn, M. H., Out, D., & Rombouts, S.
A. R. B. (2012). Attachment in the brain: Adult attachment representations predict
amygdala and behavioral responses to infant crying. Attachment and Human Development,
14(6), 533–551. https://doi.org/10.1080/14616734.2012.727252
Rodriguez, C. M., & Green, A. J. (1997). Parenting stress and anger expression as predictors of
child abuse potential. Child Abuse and Neglect, 21(4), 367–377.
https://doi.org/10.1016/S0145-2134(96)00177-9
Rodriguez, C. M., Russa, M. B., & Kircher, J. C. (2015). Analog assessment of frustration
tolerance: Association with self-reported child abuse risk and physiological reactivity. Child
Abuse and Neglect, 46, 121–131. https://doi.org/10.1016/j.chiabu.2015.02.017

124
Rossi, A. S. (1968). Transition to Parenthood. Journal of Marriage and the Family, 30(1), 26.
https://doi.org/10.2307/350219
Saxbe, D. E., Edelstein, R. S., Lyden, H. M., Wardecker, B. M., Chopik, W. J., & Moors, A. C.
(2017). Fathers’ decline in testosterone and synchrony with partner testosterone during
pregnancy predicts greater postpartum relationship investment. Hormones and Behavior,
90. https://doi.org/10.1016/j.yhbeh.2016.07.005
van Anders, S. M. (2013). Beyond masculinity: Testosterone, gender/sex, and human social
behavior in a comparative context. In Frontiers in Neuroendocrinology.
https://doi.org/10.1016/j.yfrne.2013.07.001
Van Anders, S. M. (2012). Testosterone and sexual desire in healthy women and men. Archives
of Sexual Behavior, 41(6), 1471–1484. https://doi.org/10.1007/s10508-012-9946-2
Van Anders, S. M., Tolman, R. M., & Volling, B. L. (2012). Baby cries and nurturance affect
testosterone in men. Hormones and Behavior, 61(1), 31–36.
https://doi.org/10.1016/j.yhbeh.2011.09.012
Wilson, K. R., Hansen, D. J., & Li, M. (2011). The traumatic stress response in child
maltreatment and resultant neuropsychological effects. Aggression and Violent Behavior,
16(2), 87–97. https://doi.org/10.1016/j.avb.2010.12.007
Wingfield, J. C., Hegner, R. E., Dufty, A. M., & Ball, G. F. (1990). The “challenge hypothesis’’:
theoretical implications for patterns of testosterone secretion, mating systems, and breeding
strategies.” American Naturalist, 136(6), 829–846. https://doi.org/10.1086/285134
Wingfield, John C. (2017). The challenge hypothesis: Where it began and relevance to humans.
In Hormones and Behavior (Vol. 92, pp. 9–12). Academic Press Inc.
https://doi.org/10.1016/j.yhbeh.2016.11.008

125
Witteman, J., Van IJzendoorn, M. H., Rilling, J. K., Bos, P. A., Schiller, N. O., & Bakermans-
Kranenburg, M. J. (2019). Towards a neural model of infant cry perception. In
Neuroscience and Biobehavioral Reviews (Vol. 99, pp. 23–32). Elsevier Ltd.
https://doi.org/10.1016/j.neubiorev.2019.01.026

Abstract (if available)

Abstract The transition to fatherhood is a complex biological and psychological process with myriad neuroendocrine changes which prepare the father for sensitive and effective parenting. Biological mothers have nine months of physical and emotional changes to prepare for the impending child, but when and how this process unfolds in biological fathers remains understudied. Understanding how a father becomes a father, and which biological and psychological factors contribute to this evolution, is an important first step in creating targeted interventions to bolster father’s parenting efforts. Collectively, the included set of studies will add substantially to our understanding of the prenatal psychological and physical factors associated with the transition to fatherhood. Results from these studies may pinpoint important prenatal variables in predicting difficulties in the co-parenting relationship and the father-child relationship post birth and act as a first step in creating effective interventions for paternal readiness.

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Asset Metadata

Creator Khoddam, Hannah Marie Adams (author)

Core Title Predictors and outcomes across the transition to fatherhood

School College of Letters, Arts and Sciences

Degree Doctor of Philosophy

Degree Program Psychology

Publication Date 07/27/2020

Defense Date 05/05/2020

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag Fathers,infant cry,OAI-PMH Harvest,testosterone,transition to parenthood

Language English

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Advisor Saxbe, Darby (committee chair), John, Richard (committee member), Kaplan, Jonas (committee member), Margolin, Gayla (committee member), Miller, Lynn (committee member)

Creator Email hannah.marieadams@gmail.com,hmkhoddam@gmail.com

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