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Theory of mind processing in expectant fathers: associations with prenatal oxytocin

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Theory of mind processing in expectant fathers: associations with prenatal oxytocin

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Content Copyright 2020 Sofia I. Cárdenas
Theory of Mind Processing in Expectant Fathers: Associations with Prenatal Oxytocin
by
Sofia I. Cárdenas
A Thesis Presented to the
FACULTY OF THE USC DANA AND DAVID DORNSIFE COLLEGE OF LETTERS, ARTS,
AND SCIENCES
UNIVERSITY OF SOUTHERN CALIFORNIA
In Partial Fulfillment of the
Requirements for the Degree
MASTER OF ARTS
(PSYCHOLOGY)
December 2020

ii
Table of Contents
List of Figures .................................................................................................................................. iv
List of Tables .................................................................................................................................... v
Abstract ........................................................................................................................................... vi
1. Introduction .................................................................................................................................. 1
1.1. Fathering behavior and theory of mind .................................................................................. 2
1.2. Measuring neural correlates of theory of mind ....................................................................... 3
1.3. The role of oxytocin ............................................................................................................... 4
1.4. The current study ................................................................................................................... 6
2. Methods ........................................................................................................................................ 7
2.1. Participants ............................................................................................................................ 7
2.2. Data availability statement ...................................................................................................... 8
2.3. Procedure ............................................................................................................................... 8
2.3.1. Oxytocin collection ......................................................................................................... 9
2.3.2. Oxytocin processing ........................................................................................................ 9
2.3.3. Why-How Task ............................................................................................................. 10
2.3.4. fMRI data acquisition .................................................................................................... 11
2.3.5. fMRI data analysis ......................................................................................................... 11
2.4. Measures .............................................................................................................................. 12
2.4.1. Demographics ............................................................................................................... 12

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3. Results ........................................................................................................................................ 12
3.1. Behavioral results ................................................................................................................. 13
3.2. fMRI results ......................................................................................................................... 13
3.2.1. Covariates: Associations between education and activation during theory of mind ........ 13
3.2.2. Hypothesis 1: Neural activation during main effects of task ........................................... 14
3.2.3. Hypothesis 2: Associations between oxytocin and BOLD signal during theory of mind 14
4. Discussion .................................................................................................................................. 15
References ...................................................................................................................................... 19
Figures ............................................................................................................................................ 25
Tables ............................................................................................................................................. 32

iv
List of Figures
Figure 1. Conditions of the Why-How Task. ................................................................................... 25
Figure 2. Group-level results of the Why>Rest contrast. ................................................................ 26
Figure 3. Group-level results of the Why>How contrast. ................................................................ 27
Figure 4. Group-level results of the Why>How contrast with education as a regressor. .............. 28
Figure 5. Group-level results of the Why>How contrast with education as a regressor. .................. 29
Figure 6. Group-level results of the Why>How contrast with oxytocin as a regressor. .................... 30
Figure 7. Group-level results of the Why>How contrast with oxytocin as a regressor. .................... 31

v
List of Tables
Table 1. Descriptive Statistics of Sample Demographics and Self-Report Measures ................... 32
Table 2. Bivariate Correlation of Main Study Variables ............................................................... 33
Table 3. Group-level Results of the Why>Rest Contrast .............................................................. 34
Table 4. Group-level Results of the Why>How Contrast ............................................................. 35
Table 5. Neural Regions Positively Correlated with Education During Why>Rest Contrast ....... 38
Table 6. Neural Regions Correlated with Education During the Why>How Contrast ................. 39
Table 7. Neural Regions Positively Correlated with Oxytocin During Why>How Contrast ....... 41

vi
Abstract

Social cognition may facilitate fathers’ sensitive caregiving behavior and promote father-
infant bonding. We administered the Why-How Task, an fMRI task which elicits theory of mind
processing, to expectant fathers (N =39) who also visited the lab during their partner’s pregnancy
and provided a plasma sample for oxytocin assay. When rating “Why” an action was being
performed vs. “How” the action was being performed, participants showed neural activity in regions
theorized to support theory of mind, including medial prefrontal cortex, right temporoparietal
junction, and posterior cingulate cortex. Fathers’ prenatal oxytocin levels predicted greater activation
to the Why condition in regions supporting theory of mind and embodied simulation. Fathers’
propensity to engage in theory of mind processing may support their transition to parenthood.
Keywords: theory of mind, oxytocin, fatherhood, neuroimaging

1
1. Introduction

Father caregiving behavior is associated with long-term benefits to children in social,
behavioral, and cognitive domains (Sarkadi et al., 2008). Extensive research exists on the complex
underpinnings of mothers’ attachment to infants, including biological, behavioral, and psychological
adaptations that serve to support the infants’ survival (Norholt, 2020). Infants also form attachments
with fathers (Cabrera et al., 2018), but less remains known about the neurocognitive mechanisms
that support the transition to fatherhood. Within the context of father-infant relationships, social
cognition – specifically, theory of mind – may support fathers’ ability to provide sensitive and
attuned caregiving, allowing fathers to infer what an infant feels and needs (Abraham et al., 2014).
Preliminary research suggests that the neural regions that support theory of mind (e.g., superior
temporal sulcus, ventromedial prefrontal cortex, temporal poles, and lateral frontopolar cortex) may
also be implicated in fathering behavior (Abraham et al., 2014). Given that sensitive caregiving
supports parent-infant attachment for both mothers and fathers (Grossman et al., 2002; van
IJzendoorn, 1995), fathers’ theory of mind ability may be a key building block that underlies father-
infant attachment.
The research literature on the fathering brain is scarce and preliminary. Most parenting-
focused neuroimaging research has measured parents’ neural responses to infant stimuli, such as
photographs, infant cry sound, or video clips of the infant (e.g., Atzil et al., 2012; Kuo et al., 2012).
Although these studies provide insight into the neural correlates of own-infant vs. other-infant
response, they do not specify the underlying cognitive processes that may occur when parents
witness infant stimuli. Further, though several studies have linked hormones with the neural systems
implicated in fathering (Abraham et al., 2014; Mascaro et al., 2014; Wittfoth-Schardt et al., 2012),
studies with more rigorous methodology would allow for isolation of specific cognitive processes
implicated in father neurobiology. Although emerging evidence suggests changes in men’s hormones

2
before pregnancy may indicate preparation for caregiving (Khoddam et al., 2020; Saxbe et al., 2017),
no studies to our knowledge have looked specifically at prenatal levels of oxytocin, a neuropeptide
hormone that has been associated with social affiliative behavior, within expectant fathers.
The current study addresses these gaps in the literature by investigating the neural correlates
of theory of mind within expectant fathers. We also examined whether paternal prenatal oxytocin
levels predict neural activation to theory of mind.
1.1. Fathering behavior and theory of mind
The neurobiology of paternal caregiving is an emerging field of research that overlaps with
existing research on the neurobiology of motherhood. An extensive literature has evaluated the
neural basis of parenting behaviors in human maternal caregivers (for review, see Cárdenas et al.,
2019; Rilling & Mascaro, 2017). The human maternal brain recruits several subcortical regions
involved in motivation (Gregory et al., 2015) and cortical systems involved in cognitive and
emotional empathy (Rilling et al., 2013). One of the cortical systems involved in considering the
thoughts and feelings of others (e.g., temporo-parietal junction, dorsomedial prefrontal cortex
ventromedial prefrontal cortex, superior temporal sulcus/superior temporal gyrus, precuneus;
Gallagher & Frith, 2003; Molenberghs et al., 2016).
An emerging body of research has applied neurobiological methods for understanding
paternal caregiving behavior. Atzil and colleagues (2012) measured mothers’ and fathers’ brain
activity in response to their infants’ videos. While mothers had greater activity in limbic areas, such
as the right amygdala, fathers had greater activity in social-cognitive cortical areas, such as the dorsal
prefrontal cortex. A second study by the same research group (Abraham et al., 2014) presented first-
time mothers and fathers with videos of their infants while in an fMRI scanner. They compared
three groups of first-time parents with varying degrees of involvement in childrearing: mothers (i.e.,

3
primary caregivers), heterosexual fathers (i.e., secondary caregivers), and homosexual fathers (i.e.,
primary caregivers) raising infants without maternal involvement. Primary caregiver mothers showed
higher activation in the amygdala to own-infant stimuli versus other infant stimuli. In contrast, both
groups of fathers exhibited higher activation in the superior temporal sulcus, widely associated with
theory of mind. Findings from these two studies suggest that the neurobiology of fatherhood is
distinct from that of mothers and that the theory of mind network may play a particularly important
role in supporting fathering behavior.
1.2. Measuring neural correlates of theory of mind
Theory of mind is the consideration of the mental states and intentions of people (Weimer
et al., 2017). Theory of mind also allows individuals to perceive behavior as driven by unobservable
mental thoughts (Spunt & Lieberman, 2012). By age four, most typically developing children achieve
an understanding of the mind (Cutting & Dunn, 1999). However, knowledge about mental states
continues to increase after the age of four, including the ability to understand mistaken beliefs about
mental states and the role of pre-existing biases and expectations in influencing personal tastes
(Carpendale & Chandler, 1996). Theory of mind abilities improve in typically developing adults
between late adolescence and adulthood (Dumontheil et al., 2010). Differences in theory of mind
abilities for individuals correlate with higher general social competence (Lalonde & Chandler, 1995).
Thus, theory of mind has implications for social competence and interpersonal functioning (Hughes
& Leekman, 2004).
Extensive work has sought to develop accurate behavioral and neural measures that reflect
the multidimensional, evolving, and variable nature of the theory of mind construct (e.g., Keysar et
al., 2003). Meta-analyses of neuroimaging studies involving theory of mind revealed a consistent
activation of four brain regions: the dorsomedial prefrontal cortex, temporoparietal junction,
precuneus, and posterior superior temporal sulcus (e.g., Gallagher et al., 2000; for review, see Frith

4
& Frith, 2003). However, activation in the theory of mind network varies across individual studies
based on the mental state evaluated (belief vs. desire) and the stimuli used (verbal vs. non-verbal;
Spunt & Adolphs, 2014). Thus, when evaluating the theory of mind construct, one must consider
which measure accurately captures the relevant component of theory of mind.
The Why-How Task isolates the multiple cognitive processes underlying the day-to-day
theory of mind that occurs when individuals perceive others’ actions (Spunt & Lieberman, 2012).
The task is one of several variants created by a team of researchers studying social cognition (Spunt
& Lieberman, 2012; Spunt & Adolphs, 2014).). The “Why” process is supported by brain regions
associated with theory of mind, whereas the “How” process is underlined by areas related to action
perception (Spunt & Lieberman, 2012). In the task, participants watch sixteen blocks of eight
photographs. Each of the blocks leads to a response question meant to elicit processes associated
with action perception ("Is the person looking sideways?") or theory of mind ("Is the person
helping?"; Spunt & Adolphs, 2014).
1.3. The role of oxytocin
Many recent studies have investigated the neuroendocrine underpinnings of parenting
(Gordon et al., 2010; Scatliffe et al., 2019). Some studies have found associations between fathers’
levels of hormones, including oxytocin, testosterone, and vasopressin and their caregiving behavior
(Feldman & Baermans-Kranenburg, 2017; Gettler, 2014; Storey & Zeigler, 2016). Oxytocin has been
associated with social and affiliative behavior, including behavioral synchrony and responsiveness in
mothers and fathers (Feldman et al., 2010; Morris et al., under review; Scatliffe et al., 2019). In
fathers, oxytocin has been specifically implicated in stimulatory parenting behaviors (Feldman et al.,
2010; Scatliffe et al., 2019), which may support father-infant attachment (George et al., 2010).
Further, experimentally administered oxytocin levels elicit more stimulating play in fathers (Gordon
et al., 2010; Naber et al., 2010; Weisman et al., 2014). Increased levels of oxytocin around the

5
transition to parenthood may indicate the potential for enhanced attachment in fathers. Similarly,
men’s ability to engage in theory of mind may support father-infant attachment.
The role of oxytocin in modulating theory of mind, however, has received little attention.
Few studies have investigated the role of oxytocin, brain activity, and father caregiving behavior
(e.g., Abraham et al., 2014; Atzil et al., 2012; Li et al., 2017; Mascaro et al., 2014). One study found
that when mothers and fathers watched videos of infant play, mothers showed relatively greater
activation in subcortical regions (e.g., amygdala, nucleus accumbens), while fathers had greater
activation in social-cortical regions. Further, fathers’ postpartum plasma oxytocin levels negatively
correlated with activity in the left inferior and superior frontal gyri, left primary motor cortex, medial
prefrontal cortex, and left anterior cingulate cortex (Atzil et al., 2012). Building upon this literature,
Abraham and colleagues (2014) assessed oxytocin and brain activity in parent caregiving behavior.
All participants provided salivary oxytocin data, completed a functional MRI task in which they
watched videos of their children, and took part in a father-infant play task micro-coded for parent-
infant behavioral synchrony. Though oxytocin levels were similar across mothers and fathers’
groups, fathers showed greater activity in areas of the brain involved in theory of mind, perspective-
taking, and empathy (i.e., ventromedial prefrontal cortex, superior temporal sulcus). Fathers’
superior temporal sulcus activation while watching infant videos correlated with their oxytocin levels
as well as parent-infant synchrony. This study suggests that fathers’ social cognition, oxytocin levels,
and fathering behavior may be correlated. However, as this study used a non-standardized fMRI
task, further research should clarify how oxytocin interacts with specific social-cognitive processes.
Additionally, this study used salivary oxytocin assayed without extraction, a controversial approach
to measuring oxytocin that may not be considered reliable and has attracted controversy within the
literature (MacLean et al., 2019).

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Surprisingly, no studies to our knowledge have assessed prenatal oxytocin levels in expectant
fathers. Men and women have been found to show shifts in hormone levels from the prenatal to the
postpartum period, and some evidence suggests that these shifts in hormones support parenting
outcomes (Edelstein et al., 2017; Saxbe et al., 2017). For instance, Storey and colleagues (2000)
found that men and women had similar stage-specific shifts in cortisol, testosterone, and prolactin
across the prenatal and postpartum periods. However, some research suggests that oxytocin does
not shift similarly across pregnancy for mothers and fathers. An older study by Leake et al. (1981)
found that healthy men, non-pregnant women, and pregnant women had similar levels of plasma
oxytocin, suggesting that oxytocin levels during pregnancy may not change. In contrast, Gordon et
al. (2010) found intraindividual stability during the first six months postpartum and interrelated
plasma oxytocin levels in mother-father dyads at one and six months postpartum. Further, paternal
oxytocin correlated with stimulatory parenting behavior. These findings suggest that oxytocin levels
may be consistent across time and provide preliminary support for investigating the utility of
prenatal oxytocin in men as a predictor of postpartum father-infant outcomes.
1.4. The current study
Although neural and behavioral research suggests that oxytocin and theory of mind may
support fathers in the transition to parenthood, our paper uniquely adopts a standardized,
longitudinal approach to measuring theory of mind in tandem with oxytocin. Moreover, we used
immunoassay with extraction to measure oxytocin levels in plasma, considered to be the gold
standard for oxytocin assay. The present study used the Why-How Task (Spunt & Adolphs, 2014),
an fMRI task designed to elicit theory of mind processing within a sample of expectant fathers.
Given the evidence that education may be associated with parenting (Cabrera et al., 2014) and that
fathers undergo neural plasticity during pregnancy (Kim et al., 2014), we controlled for partners’
pregnancy stage and fathers’ education in all analyses. We tested four hypotheses:

7
(i) Consistent with Spunt and Adolphs (2014), we predicted that expectant fathers would
show greater neural activation during the Why versus How contrast in regions that have
been associated with theory of mind specifically (e.g., dorsomedial prefrontal cortex;
ventromedial prefrontal cortex; lateral orbitofrontal cortex; temporoparietal junction;
posterior cingulate cortex; temporal pole; anterior superior temporal sulcus). Though not
reported in the original Spunt and Adolphs (2014) study, we also predicted that the Why
versus Rest contrast would produce activation in regions that support social cognition. We
planned to use whole-brain analyses during the Why > How and Why > Rest contrasts to
test this hypothesis.
(ii) As discussed above, oxytocin influences fathering behavior and is associated with neural
activity in regions linked to theory of mind (Abraham et al., 2014). We hypothesized that
fathers with higher prenatal oxytocin levels would show more activation in brain regions
associated with theory of mind processing during the Why > How and Why > Rest
contrasts.
2. Methods
2.1. Participants
The current study uses data from the larger longitudinal Hormones and Attachment across
the Transition to Childrearing (HATCh) study, which follows couples from the mothers’ mid-to-late
pregnancy period through the first year postpartum. We recruited participants through social media
advertising (e.g., Facebook, online parenting groups) and also through flyering and word of mouth.
The current study used self-reported data provided by fathers from prenatal laboratory visits,
conducted with both members of the couple when mothers were in mid-to-late pregnancy,
subsequent father-only MRI visits that we scheduled within approximately two weeks of the prenatal
in-lab visit, and postpartum questionnaires collected online approximately three months after the

8
birth. Eligible couples were cohabitating, first-time parents of a singleton pregnancy, and did not
report any medications or conditions known to interfere with endocrine system hormones (e.g.,
steroid medicines, Cushing’s disease). We also excluded couples who reported psychiatric illness
requiring medication or illegal drug use. Users of tobacco, marijuana, and we allowed some
psychiatric medications if able to abstain for 24 hours before their study visit. Additional exclusion
criteria included contraindications for magnetic resonance imaging (MRI), such as left-handedness,
Neurological or movement disorders, claustrophobia, history of brain injury, psychotropic
medication, or severe learning disability. Additionally, participants needed to have sufficient English
language fluency to complete study measures and scanning procedures. University IRB approved all
procedures, and all participants signed informed consent forms before participation.
Data for the current study were available for 39 expectant fathers who provided prenatal
demographic and neuroimaging data. Of these 39 fathers, 34 provided prenatal blood samples for
oxytocin assay, and we excluded one oxytocin sample due to lab processing issues. The mean age of
fathers was 31.56 years (SD = 4.25 years). The sample was ethnically and racially diverse, with
fathers reporting European American (33.3%), Latino (30.8%), Black or African American (5.1%),
Asian-American (25.7%), and multiracial or other heritage (5.1%). The majority of expectant fathers
(76.9%) reported a college degree or higher.
2.2. Data availability statement
The data that support the findings of this study are available from the corresponding author upon
reasonable request.
2.3. Procedure
During the prenatal in-lab visits, fathers and mothers participated in several discussion tasks
regarding their feelings and expectations about the pregnancy and birth, their projected division of
childcare, and areas of conflict within the relationship. They also completed psychosocial

9
questionnaires, including the measures described below. Licensed phlebotomists conducted blood
draws at the end of the lab visits. The majority of fathers (32 out of 39 expectant fathers; 82.05%)
completed MRI scan visits within two weeks of the in-lab visit, with a range of .14 weeks to 5.71
weeks. During the MRI visit, fathers completed the Why-How Task (Spunt & Adolphs, 2014) as
part of a larger MRI data collection protocol.
2.3.1. Oxytocin collection

We collected blood for the plasma oxytocin assay in a sterile EDTA vacutainer tube. We
added twenty microliters of a protease inhibitor (Amastatin; 10 µM final concentration) to the tube
to limit oxytocin degradation. Then, we centrifuged blood samples for 10 minutes to separate
plasma. We stored aliquots at -80 C and shipped on dry ice to the University of Miami School of
Medicine Diabetes Research Institute (Armando Mendez, PI) for oxytocin processing.
2.3.2. Oxytocin processing

We used enzyme-linked-immunosorbent assay (ELISA) kits (Arbor Assays; Ann Arbor, MI)
to calculate oxytocin immunoreactivity. In our sample, the lower limit of detection was 0.8 pg/ml. In
line with prior work (Szeto et al., 2011), we extracted 2.2 ml of plasma. Further, we reconstituted in
220 µl assay buffer leading to a 10-fold concentration compared to the starting plasma volume. We
performed assays in line with manufacturer instructions. We assessed samples in duplicate. The
inter-assay coefficient of variation was less than 10%. Of the MRI sample, nine fathers with
oxytocin (27% of the sample) had oxytocin levels that resulted as below detection levels following
extraction. Our rate is similar to prior research analyzing extracted oxytocin (e.g., Szeto et al., 2011;
Tabak et al., 2011). In line with common practice, we ascribed samples below detection level a value
of 0.4 pg/mL (Saxbe et al., 2019).
We analyzed the extracted oxytocin data for outliers and normality using SPSS Statistics 26
(IBM) (M = 1.99; SD = 1.90 Skewness = 1.87). Further, we truncated the data by dropping outliers

10
greater than 3 standard deviations from the sample (3%). We found that extracted oxytocin data
remained positively skewed after excluding outliers (M = 1.50; SD = 1.05; Skewness=.74). As a
consequence, we natural log-transformed the data to satisfy normality assumptions (M = .23;
SD=.88; Skewness = -.03).
2.3.3. Why-How Task

The Why-How Localizer Task is a standardized task for investigating social cognition (Spunt
& Adolphs, 2014). The task uses Psychophysics Toolbox Version 3 (PTB-3) and MATLAB (The
MathWorks, Inc.). The task presents individuals with 16 blocks of items, with each block consisting
of a question prompt and eight photographs. The task consists of 42 pictures of common hand
actions and 42 pictures of familiar facial expressions. The question prompts present as pre-tested
yes/no questions, with participants given a limited amount of time for response. The task requires
approximately six minutes and thirty seconds to complete. Participants complete the task in a suite
for a magnetic resonance imaging (MRI) scanner. Within the scanner, participants watch stimuli
while lying down in the scanner and can use a button box programmed to allow for yes/no
responses.
The Why-How Task features a 2 (condition: Why, How) X 2 (behavioral category: face,
hand) factorial design (see Figure 1). For the behavioral factor, participants watch a photograph with
either a face or a hand. For the stimulus factor, participants receive a stimulus prompt that asks
either a Why question (i.e., the question elicits theory of mind) or a How question (i.e., the question
evokes action-perception). In the present study, similar to Spunt & Adolphs (2014), we collapsed the
behavioral category conditions (i.e., face, hand) and focused on Why and How conditions. We
contrasted Why > How and Why > Rest to specifically examine the neural correlates of theory of
mind processing. We also scored the Why-How task results behaviorally for accuracy and response
time and entered this information in SPSS for further analysis.

11
2.3.4. fMRI data acquisition

We acquired whole-brain images on a Siemens 3 Tesla MAGNETON Prisma System
scanner, 20-channel matrix head coil. We acquired high-resolution, T1-weighted images using a 3D
Magnetization Prepared Rapid Acquisition Gradient Echo (repetition time, 2530 ms; echo time, 3.13
ms; flip angle, 10°), with an isotropic voxel resolution of 1mm
3
. We collected functional data using a
T2* weighted echo-planar imaging (EPI) with an interleaved sequence (194 2.5 mm transversal
slices; repetition time, 2000 ms; echo time, 25 ms; field of view, 192 mm
2
; 3.0 x 3.0 x 2.5 mm voxels,
flip angle, 90 degrees). We used the Flywheel platform to collect all our data and used the moco files
for all functional data.
We presented stimulus presentation and response recording using MATLAB (version 2012a;
MathWorks Inc., Natick, MA, USA). We used an LCD projector to show stimuli on a rear-
projection screen. Participants made responses using their right index and middle fingers on a
button box.
2.3.5. fMRI data analysis

We analyzed neural responses to the Why > How contrast Why > Rest contrast using FEAT
(FMRI Expert Analysis Tool) Version 6.00, part of FSL (FMRIB’s SoftwareLibrary,
www.FMRIb.ox.ac.uk/fsl). We used FLIRT (Jenkinson 2001, 2002) to register high resolution
structural and/or standard space images. For pre-processing, we conducted motion correction with
MCFLIRT [Jenkinson 2002], non-brain removal using BET [Smith 2002], spatial smoothing using a
Gaussian kernel of FWHM 5mm, and high pass temporal filtering with Gaussian-weighted least-
squares straight line fitting and a sigma=50.0s).
Next, we analyzed data using a General Linear Model with a multi-level mixed-effects
design. We entered each component of the task as a regressor (Why-Face trials, Why-Hand trials,
How-Face trials, Why-Hand trials) and modeled by convolving the task design with a double-gamma

12
hemodynamic response function with a phase of 0 s. We defined the task periods from the first
photograph in each block to the stimulus offset of the last photograph in each block. We created
contrasts as linear combinations of the explanatory variables to look at overall task effects as well as
condition effects (e.g., Why > How; Why > Rest). At the individual subject level, we generated
statistical maps. We then computed brain-activity maps for each of the task effects and condition
effects combining across all subjects using a higher-level mixed-effects design using FLAME to
produce contrast-level activity maps. This study focused on the Why > How and Why > Rest
contrasts. All higher-level analyses, except for the main effect analysis, included fathers’ education
level and weeks pregnant as covariates. To account for multiple comparisons, cluster thresholding
using Gaussian Random Field theory was applied with a cluster-forming z threshold of 2.3 and a
cluster size probability threshold of P<0.05. All coordinates reported are in the Montreal
Neurological Institute (MNI) standard brain space.
2.4. Measures
2.4.1. Demographics

At the prenatal visit, fathers self-reported their educational attainment with the following
options: high school/GED, some college, associate degree, bachelor’s degree, master’s degree,
professional/doctorate). Fathers also reported on age, and mothers reported their expected due date,
from which we calculated their pregnancy stage (number of days pregnant) at the time of their
prenatal study visit.
3. Results
We present variable descriptive statistics in Table 1. Zero-order correlations between study
variables are shown in Table 2. Fathers’ education and age were positively correlated (r (39) = .57, p
< .01). Age correlated with accuracy during How trials (r (39) = .34, p < .05). Fathers’ accuracy
during Why trials correlated with fathers’ accuracy during How trials (r (38) = .78, p < .01). Fathers’

13
response time during Why trials correlated with fathers’ response time during How trials (r (38) =
.93, p < .01). We also found a trending negative correlation between extracted oxytocin and response
time during How trials (r (32) = -.35, p = .05).
3.1. Behavioral results
Within this sample, we found behavioral results consistent with Spunt et al., 2014. The
percent accuracy on How (M = 94%, SD = 6.2%) trials differed from Why (M = 91%, SD = 7.4%)
trials, t (38) = 4.30, p < .01, such that participants responded less accurately to Why trials. Response
time to How (M = 820 ms., SD = 134 ms.) trials differed from Why (M = 890 ms., SD = 155) trials,
t (37) =-8.45, p <.01, such that participants responded more slowly during Why trials.
3.2. fMRI results
3.2.1. Covariates: Associations between education and activation during theory of mind

Higher educational attainment predicted greater activation in theory of mind regions during
the Why > How and Why > Rest contrasts. When we added education level as a regressor into the
Why > Rest contrast, signal changes associated with this covariate emerged in a cluster, including the
right middle temporal gyrus (see Table 5 and Figure 4). When we added education level as a
regressor into the Why > How contrast (see Table 6 and Figure 5), signal changes associated with
this covariate occurred in four clusters, including postcentral gyrus, precentral gyrus, and right
precuneus.
Given these findings, we included education as a covariate in all regressor analyses (i.e.,
hypothesis 2 and hypothesis 3). In line with the work of Khoddam et al. (2020), which used the
same sample as the current study, pregnancy stage was also included as a covariate in all analyses.
Thus, we mean-centered education level and pregnancy stage (i.e., weeks pregnant) sand included as
confound regressors in all analyses except for the main effect hypothesis (i.e., hypothesis 1).

14
3.2.2. Hypothesis 1: Neural activation during main effects of task
We partially replicated neural findings from the Spunt and Adolphs paper (2014). The full
list of brain regions activated by Why > How and Why > Rest contrast is listed in Table 3 and Table
4, respectively. A visual representation of the brain regions activated by the Why > Rest and Why >
How contrasts is shown in Figure 2 and Figure 3, respectively.
When contrasting activation during Why trials compared with Rest (i.e., Why > Rest
contrast), we found significant clusters in the right fusiform gyrus, bilateral anterior insula, right
precentral gyrus, and bilateral ventromedial prefrontal cortex.
When contrasting activation during Why trials compared with Rest (i.e., Why > Rest
contrast), we found activation in the right fusiform gyrus, bilateral anterior insula, right precentral
gyrus, bilateral paracingulate gyrus, and bilateral ventromedial prefrontal cortex.
When contrasting activation during Why trials, compared with How trials (i.e., Why > How
contrast), we found activation in the original group-level results of the study: left dorsomedial
prefrontal cortex, left ventromedial prefrontal cortex, left lateral orbitofrontal cortex, right
temporoparietal junction, bilateral posterior cingulate cortex, left temporal pole, and right anterior
superior temporal sulcus. These results largely replicate findings from the original Why-How paper
(Spunt & Adolphs, 2014). However, during the Why > How contrast, the present study found
additional activation in several areas not found in the original task study: left lateral occipital cortex,
left amygdala, left dorsolateral prefrontal cortex, left caudate, left thalamus, and right posterior
superior temporal gyrus.
3.2.3. Hypothesis 2: Associations between oxytocin and BOLD signal during theory of mind

Prenatal oxytocin was added as a regressor in the original Why > Rest and Why > How.
contrasts. Higher prenatal plasma oxytocin predicted greater signal in theory of mind regions during
the Why > Rest and Why > How contrasts. We added oxytocin levels as regressors into the Why >

15
Rest contrast, activation emerged in a left-lateralized cluster, including angular gyrus, supramarginal
gyrus, Heschel’s gyrus, and superior temporal gyrus (see Table 7 and Figure 6). When we added
prenatal oxytocin levels as regressors into the Why > How contrast, activation emerged in eight
clusters, including bilateral angular gyrus, bilateral supramarginal gyrus, bilateral middle frontal gyrus,
bilateral posterior cingulate gyrus, and right superior temporal gyrus (see Table 8 and Figure 7).
4. Discussion
In this study, we partially replicated the results from the Spunt and Adolphs (2014) paper
with a sample of expectant fathers. Specifically, we found that theory of mind processing during the
Why > How contrast revealed activation in left dorsomedial prefrontal cortex, left ventromedial
prefrontal cortex, left lateral orbitofrontal cortex, right temporoparietal junction, bilateral posterior
cingulate cortex, left temporal pole, and bilateral anterior superior temporal sulcus. Moreover, we
found that prenatal oxytocin in fathers during the Why > Rest contrast was associated with
activation in a left-lateralized cluster including that spans parts of angular gyrus, supramarginal gyrus,
Heschel’s gyrus, and superior temporal gyrus.
We found evidence to partially support our main effect hypothesis that areas of the brain
associated with social cognition and theory of mind appear active during the contrasts that isolate
neural activation during theory of mind: Why > Rest, Why > How. During the Why versus Rest
contrast, as mentioned above, activity emerged in the right fusiform gyrus, bilateral anterior insula,
right precentral gyrus, bilateral paracingulate gyrus, and bilateral ventromedial prefrontal cortex.
Since the original task study (Spunt & Adolphs, 2014) did not focus on the Why > Rest contrast, we
cannot compare current results to other Why-How papers. However, the anterior insula relates to
social cognition. For instance, the anterior insula is active when individuals experience pain or see
others in pain, suggesting anterior insula may be involved in empathy (Eisenberger & Cole, 2012).
Further, anterior insula activity has been found in studies where fathers watch infant imagery

16
(Mascaro et al., 2014), and supports a fathers’ ability to resonate with infants’ emotions (Feldman et
al., 2015). The ventromedial prefrontal cortex is involved in theory of mind reasoning (Shamay-
Tsoory & Aharon-Peretz., 2007). Within the context of the father-infant relationship, the
ventromedial prefrontal cortex is considered to be part of the mentalizing network, thought to
support a caregivers’ ability to understand nonverbal signals and infer infant intentions (Feldman et
al., 2015).
As expected, contrasting activation during Why trials compared to How trials (i.e., Why >
How contrast) revealed activation in regions involved in social cognition and theory of mind: left
dorsomedial prefrontal cortex, left ventromedial prefrontal cortex, left lateral orbitofrontal cortex,
right temporoparietal junction, bilateral posterior cingulate cortex, left temporal pole, and right
anterior superior temporal sulcus. Though not expected, the Why > How contrast revealed
activation in parts of the left lateral occipital cortex, left amygdala, left dorsolateral prefrontal cortex,
left caudate, left thalamus, and right posterior superior temporal gyrus. The ventromedial prefrontal
cortex, temporoparietal junction, posterior cingulate cortex, temporal pole, and superior temporal
sulcus are part of the parental mentalizing network and often emerge in studies where fathers watch
infant stimuli while in an fMRI scanner (Abraham et al., 2014; Feldman et al., 2015).
Though not the focus of the study, fathers’ education level related to activation during both
theory of mind contrasts (i.e., Why > Rest, Why > How). When contrasting the Why trials versus
Rest trials, education was associated with greater activity in a cluster primarily composed of right
middle temporal gyrus. The middle temporal gyrus is involved in many cognitive processes,
including sensory integration (Mesulam, 1998) and theory of mind (for meta-analysis, see Schurz &
Perner, 2015). When we contrasted Why trials with the How trials (i.e., Why > How contrast),
education was also related with neural activity in areas involved in sensory integration (i.e., right
superior parietal gyrus, left supramarginal gyrus; Culham & Valyear, 2006; Sliwinska et al., 2012), the

17
mirror neuron system (i.e., precentral gyrus and left postcentral gyrus; Wu et al., 2017) and theory of
mind (i.e., right precuneus; Saxe et al., 2006). In the parenting network, the mirror neuron system
and the theory of mind network work together to support a parents’ ability to quickly resonate with
infant behaviors (i.e., mirror neuron system), consider intentions (i.e., theory of mind network), and
provide caregiving behavior that synchs with the child’s needs (Feldman et al., 2015). Research
suggests that education level supports fathers’ caregiving involvement (Cabrera et al., 2014). These
findings might also reflect that more educated fathers may exhibit more comfort or engagement
with the Why-How Task.
Additionally, fathers’ prenatal oxytocin levels predicted activation in areas of the brain that
support embodied simulation and theory of mind. Of note, we collected prenatal oxytocin following
a lab visit that included partner interaction and questions relevant to the transition to parenthood.
Activation during both the Why> Rest contrast and Why >How contrast was associated with
prenatal oxytocin in a region, including the left angular gyrus and left supramarginal gyrus, which
compose the inferior parietal lobule (IPL). The angular gyrus has been implicated in theory of mind
and mentalizing tasks (for review, see Seghier, 2013). With regard to parenting, the IPL is part of the
embodied simulation network as well as the mirror neuron system. The mirror neuron system
supports an individual’s ability to observe and mimic the emotions and behaviors of others (for
review, see Rizzolatti & Craighero, 2004). The embodied simulation network supports a parents’
ability to simulate the behavior of the infants and consider their intentions and needs (Feldman,
2015). The Why > How contrast was also related to oxytocin in areas that support social cognition,
including the posterior cingulate cortex, a key region of the Default Mode Network (Li et al., 2014),
Given the minimal research on prenatal oxytocin in expectant fathers, future studies would benefit
from sampling oxytocin and measures of social cognition across the transition to parenthood in

18
expectant fathers to understand whether changes in social cognition shift in preparation for
fatherhood.
The current study has a number of limitations. Although larger than other samples in the
neurobiology of parenting literature, the sample size was small (39 fathers with MRI data; 33 with
oxytocin data). Further, we collected oxytocin levels during a laboratory visit that preceded the
neuroimaging scan by approximately two weeks. Additionally, the oxytocin data collection and
neuroimaging scan both only occurred at one time point during pregnancy, despite evidence that
men show both hormonal and neural changes across the transition to parenthood; Kim et al., 2014;
Storey et al., 2000) and that neural regions involved in social cognition can develop an change over
time (Blakemore, 2008; Dumontheil et al., 2010). As previously noted, limited literature exists on the
role of oxytocin across the transition to parenthood. Future studies can extend our work by
collecting neural measures of social cognition, both pre and postpartum.
Despite the aforementioned limitations, the present study makes several important
contributions to research on parenting and child development. This study extends prior work on the
neurobiology of fatherhood by using a standardized task to measure social cognition in expectant
fathers. Also, the present study is novel as we measure prenatal hormones in tandem with brain
activation in expectant fathers and the first to focus specifically on prenatal oxytocin. Our sample of
expectant fathers was racially and ethnically diverse and contributed longitudinal, multimodal data,
including neuroimaging, oxytocin, and self-report measures. Our results suggest that expectant
fathers have overlapping neural activation related to oxytocin and social cognition during the
prenatal period. These findings contribute to understanding of neurobiological changes underlying
fathers’ preparation for parenthood.

19
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25
Figures

Figure 1. Conditions of the Why-How Task. The above diagram displays the 2 X 2factorial design of
the Why-How Task with type of question (condition: Why, How) and stimulus (condition: face,
hand).

26

Figure 2. Group-level results of the Why>Rest contrast. Sagittal, coronal, and axial view of whole
brain activation during the Why > How contrast. Analyses cluster corrected at z = 3.1. (N = 39).

27

Figure 3. Group-level results of the Why>How contrast. Sagittal, coronal, and axial view of whole
brain activation during the Why > How contrast. Analyses cluster corrected at z = 3.1. (N = 39).

28

Figure 4. Group-level results of the Why>How contrast with education as a regressor. Sagittal,
coronal, and axial view of whole brain activation during the Why > How contrast. Analyses cluster
corrected at z = 2.3. (N = 39).

29

Figure 5. Group-level results of the Why>How contrast with education as a regressor. Sagittal,
coronal, and axial view of whole brain activation during the Why > How contrast. Analyses cluster
corrected at z = 2.3. (N = 39).

30

Figure 6. Group-level results of the Why>How contrast with oxytocin as a regressor. Sagittal,
coronal, and axial view of whole brain activation during the Why > How contrast. Analyses cluster
corrected at z = 2.3. (N = 33).

31

Figure 7. Group-level results of the Why>How contrast with oxytocin as a regressor. Sagittal,
coronal, and axial view of whole brain activation during the Why > How contrast. Analyses cluster
corrected at z = 2.3. (N = 33).

32
Tables

Table 1. Descriptive Statistics of Sample Demographics and Self-Report Measures
Variable Subcategory Value/Mean (%/SD)
Education High school/GED 1 (2.6%)
Some college 7 (17.9%)
Associate degree 1 (2.6%)
Bachelor’s degree 12 (30.8%)
Master’s degree 11 (28.2%)
Professional/Doctorate 7 (17.9%)
Race European American 13 (33.3%)
Latino 12 (30.8%)
Black or African American 2 (5.1%)
Asian-American 10 (25.7%)
Multiracial or Other 2 (5.1%)
Age at prenatal visit 31.56 (4.25), 23-41
Extracted oxytocin .23 (.88), -.92-1.92
Weeks pregnant 29.50 (4.70), 21.81-38.94
Weeks between lab and MRI 1.63 (1.15), .14-5.71
Paternal postnatal attachment 4.30 (0.43), 3.07-4.89
Note. Weeks pregnant is the self-reported time fathers reported their female partners were pregnant
on the day of the lab visit. * indicates p < .05. ** indicates p < .01.

33
Table 2. Bivariate Correlation of Main Study Variables
Variable 1 2 3 4 5 6 7
1. Age
2. Education .57
***

3. Weeks Pregnant .22 .29
*

4. Extracted Oxytocin -.05 -.10 .23
5. Why trials accuracy .14 .12 .02 -.12
6. How trials accuracy .34
**
.18 .14 -.05 .78
***

7. Why trials RT .28
*
.12 -.04 -.28 .26 .05
8. How trials RT .17 .07 -.08 -.35
*
.30
*
.01 .93
***

Note. Weeks pregnant is the self-reported time fathers reported their female partners were pregnant
including the time between the lab visit and the MRI visit. * indicates p < .10. ** indicates p < .05.
*** indicates p < .01.

34
Table 3. Group-level Results of the Why>Rest Contrast
MNI coordinates
Region L/R k x y z Z
Fusiform Gyrus R
39878
30 -52 -10 8.56
R - 42 -46 -16 8.26
R - 38 -36 -20 8.17
Anterior Insular Cortex L - -30 24 -2 8.43
R - 34 24 -2 8.14
Precentral Gyrus R - 40 10 30 8.07
Paracingulate Gyrus R 3169 6 22 48 7.1
L - -4 8 54 7.1
R - 4 16 54 7.05
R - 8 20 44 6.99
R
-
4 8 56 6.96
R - 4 22 42 6.88
Ventromedial PFC L 144 -2 48 -16 4.15
R - 4 60 -14 4.02
R - 4 48 -16 3.89
L - -2 58 -14 3.89
Note. All peaks survived a whole-brain search thresholded at a voxel-wise family-wise error rate of
.05. PFC = prefrontal cortex; STS = superior temporal sulcus; TPJ = tempo parietal junction: x, y, z
= Montreal Neurological Institute (MNI) coordinates in the left-right anterior-posterior, and
inferior-superior dimensions respectively.

35
Table 4. Group-level Results of the Why>How Contrast
MNI coordinates
Region L/R k x y z Z
Lateral OFC L
6439
-40 28 -16 6.57
Anterior STS L - -58 -8 -14 6.56
Temporal pole L - -48 16 -22 6.44
Temporal pole L - -54 4 -16 6.44
Temporal pole L - -42 -62 24 6.33
Temporal pole L - -52 22 12 6.28
Dorsomedial PFC L
5937
-8 64 26 7.39
Dorsomedial PFC L - -4 56 38 7.37
Dorsomedial PFC L - -10 60 32 7.33
Dorsomedial PFC L - -10 44 46 7.25
Dorsomedial PFC L - -4 52 -14 7.18
Ventromedial PFC L - -2 48 -16 7.25
Posterior cingulate cortex
1844
0 -50 30 6.44
Posterior cingulate cortex L - -4 -60 34 6.22
Posterior cingulate cortex L - -2 -60 30 5.93
Posterior cingulate cortex L - -4 -50 22 5.82
Posterior cingulate cortex L - -10 -52 4 4.4
Posterior cingulate cortex R - 14 -48 6 3.68
Anterior STS R
709
58 2 -22 5.43
Anterior STS R - 60 -6 -18 5.24

36
Table 4 Continued
MNI coordinates
Region L/R k x Region L/R k
Anterior STS R - 62 -2 -22 5.15
Anterior STS R - 54 8 -22 5.05
Anterior STS
R - 50 -8 -12 4.99
Anterior STS R - 48 -14 -10 4.91
Amygdala L 270 -26 -18 -18 5.08
Amygdala L - -20 -10 -14 4.98
Amygdala L
-
-28 -6 -22 4.18
Amygdala L - -36 -16 -22 3.42
Dorsolateral PFC L 241 -42 14 54 4.38
Dorsolateral PFC L - -36 24 52 4.31
Dorsolateral PFC L
-
-34 18 50 4.31
Dorsolateral PFC L - -44 24 48 3.62
Temporoparietal junction R 171 48 -58 24 4.27
Temporoparietal junction R - 54 -62 26 4.08
Temporoparietal junction R
-
60 -56 28 3.63
Caudate L 151 -12 14 10 4.6
Caudate L - -10 -4 12 3.23
Posterior STG R
110
64 -34 4 4.04
Posterior STG R - 56 -28 4 3.91
Posterior STG R 46 -32 -2 3.66
Thalamus
L
104

-4 -12 6 4.28
Thalamus - 0 -18 10 4

37
Note. All peaks survived a whole-brain search thresholded at a voxel-wise family-wise error rate of
.05. OFC= Orbitofrontal cortex; PFC = prefrontal cortex; STS = superior temporal sulcus. x, y, z =
Montreal Neurological Institute (MNI) coordinates in the left-right anterior-posterior, and inferior-
superior dimensions respectively.

38
Table 5. Neural Regions Positively Correlated with Education During Why>Rest Contrast
MNI coordinates
Region L/R k x y z Z
Frontal Medial Cortex/Frontal
Pole R 498 20 28 -6 3.57
R 12 36 -10 3.38
R 12 34 -6 3.33
R 18 46 -16 3.31
R 12 54 -6 3.3
R 10 44 -18 3.3
Middle Temporal Gyrus R 380 62 -48 2 3.84
Middle Temporal Gyrus R 54 -54 -6 3.73
Middle Temporal Gyrus R 48 -64 0 3.03
Middle Temporal Gyrus R 50 -62 -6 3.01
Middle Temporal Gyrus R 50 -44 -4 2.83
Middle Temporal Gyrus R 50 -70 6 2.72
Note. All peaks survived a whole-brain search thresholded at a voxel-wise family-wise error rate of
.05. x, y, z = Montreal Neurological Institute (MNI) coordinates in the left-right anterior-posterior,
and inferior-superior dimensions respectively.

39
Table 6. Neural Regions Correlated with Education During the Why>How Contrast
MNI coordinates
Region L/R k x y z Z
Precuneus R 2127 18 -62 42 4.55
Superior Parietal Lobule R - 30 -40 46 4.38
Lateral Occipital Cortex R - 10 -66 60 4.05
Agular Gyrus R - 54 -52 50 3.93
Superior Parietal Lobule R - 32 -50 56 3.87
Lateral Occipital Cortex R - 34 -56 64 3.85
Precentral Gyrus R 824 10 -32 50 3.76
Lateral Occipital Cortex L - -22 -60 56 3.73
Precuneus R - 2 -42 54 3.71
Precuneus R - 4 -38 54 3.69
Posterior Cingulate Gyrus R - 8 -34 44 3.52
Precuneus L - -16 -58 52 3.17
Postcentral Gyrus L 632 -46 -28 52 4.02
Supramarginal Gyrus L - -62 -38 44 3.91
Supramarginal Gyrus L - -62 -42 44 3.8
Postcentral Gyrus L - -48 -26 44 3.52
Postcentral Gyrus L - -32 -26 42 3.45
Precentral Gyrus L - -40 -18 60 3.23
Inferior Temporal Gyrus R 494 50 -58 -8 3.84
Middle Temporal Gyrus R - 38 -56 2 3.46

40
Table 6 Continued
MNI coordinates
Region L/R k x y z Z
Lingual Gyrus R - 30 -64 0 3.36
Precuneus R - 18 -60 14 3.34
Lingual Gyrus R - 24 -74 2 3.24
Lingual Gyrus R - 28 -58 -2 3.24
Note. All peaks survived a whole-brain search thresholded at a voxel-wise family-wise error rate of
.05. x, y, z = Montreal Neurological Institute (MNI) coordinates in the left-right anterior-posterior,
and inferior-superior dimensions respectively

41
Table 7. Neural Regions Positively Correlated with Oxytocin During Why>How Contrast
MNI coordinates
Region L/R k x y z Z
Angular/Supramarginal
gyrus
L
3882 -48 -56 46 4.68
L - -58 -48 40 4.68
L - -50 -50 38 4.66
L - -42 -52 54 4.54
L - -52 -38 42 4.06
L - -20 -74 42 4.05
R 1481 44 -56 50 4.17
R - 46 -50 32 3.67
R - 34 -80 32 3.94
R - 40 -60 46 3.93
R - 48 -44 56 3.87
R - 52 -46 40 3.63
Middle Frontal Gyrus R 817 34 34 30 3.57
R - 44 30 26 3.45
R - 44 32 38 3.2
R - 52 36 26 3.2
R - 36 18 36 3.16
R - 22 50 24 3.25
L 811 -36 -2 54 3.76
L - -38 20 50 3.60

42
Table 7 continued

MNI coordinates
Region
L/R k x y z Z
L - -50 26 32 3.47
L - -36 26 44 3.47
L - -38 32 28 2.97
L - -50 32 24 2.96
Paracingulate Gyrus R 707 12 30 26 3.58
L - -10 42 30 3.29
R - 2 14 56 3.05
L - -6 28 44 3.14
R - 8 60 28 3.22
- 0 30 22 3.09
Intracalcarine Cortex R 561 2 -64 16 3.27
L - -16 -62 0 3.23
L - 12 -78 6 3.16
L - -12 -72 14 3.16
L - -8 -68 10 3.04
L - -18 -72 2 3.03
Posterior Cingulate Gyrus R 429 2 -36 46 3.71
R - 2 -44 50 3.57
L - -2 -30 44 3.29
L - -10 -24 42 3.25
L - -12 -20 40 3.22

43

Table 7 continued

MNI coordinates
Region
L/R k x y z Z
L - -4 -22 46 2.92
Superior Temporal Gyrus R 340 48 -20 -4 4.04
R - 62 -20 2 3.21
R - 62 -28 2 3.10
Middle Temporal Gyrus R - 70 -26 -4 3.88
R - 60 -12 -10 3.44
R - 60 -22 -18 3.28
Note. All peaks survived a whole-brain search thresholded at a voxel-wise family-wise error rate of
.05. x, y, z = Montreal Neurological Institute (MNI) coordinates in the left-right anterior-posterior,
and inferior-superior dimensions respectively.

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Asset Metadata

Creator Cárdenas, Sofia I. (author)

Core Title Theory of mind processing in expectant fathers: associations with prenatal oxytocin

School College of Letters, Arts and Sciences

Degree Master of Arts

Degree Program Psychology

Publication Date 09/21/2020

Defense Date 09/18/2020

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

Tag fatherhood,neuroimaging,OAI-PMH Harvest,oxytocin,theory of mind

Language English

Contributor Electronically uploaded by the author (provenance)

Advisor Saxbe, Darby (committee chair), Bechara, Antoine (committee member), Kaplan, Jonas (committee member), Margolin, Gayla (committee member)

Creator Email sicarden@usc.edu

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c89-373657

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Rights Cárdenas, Sofia I.; Cardenas, Sofia I.

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